Leucocytozoon
Leucocytozoon is a genus of parasitic alveolates belonging to the phylum Apicomplexa.
The species of this genus use blackflies as their definitive host and birds as their intermediate host. There are over 100 species in this genus. Over 100 species of birds have been recorded as hosts to these parasites.
Life cycle
Parasites in the genus Leucocytozoon have a life cycle that involves both a bird host, and a black fly. Parasites enter the bird host in a form called a sporozoite through the bite of the blood-sucking black fly. The sporozoites invade host cells in the liver where they undergo asexual replication, forming numerous daughter cells called merozoites within 4–5 days. The duration of this stage depends in part upon the species. In some species this stage may occur in the endothelial cells instead of the liver.The newly released merozoites infect either erythrocytes, leukocytes, macrophages or endothelial cells. Those infecting the macrophages or endothelial cells develop into megaloschizonts. The megaloschizonts divide into primary cytomeres, which in turn multiply into smaller cytomeres, which mature into schizonts, which in turn divide into merozoites. In erythrocytes or leukocytes, merozoites develop into gametocytes.
The gametocytes are taken up by a blood-sucking fly as it bites the bird host. Gametocytes then mature in the insect midgut into macrogametocytes with red-staining nuclei and microgametocytes with pale-staining diffuse nuclei: these fuse to form an ookinete. The male gametocytes normally give rise to eight microgametes. The ookinete penetrates an intestinal cell of the fly and matures into an oocyst. After several days the oocyst produces ~100 sporozoites that leave and migrate to the salivary glands of the fly.
Description
The nuclei of the schizonts is enclosed in a trilaminar membrane with peripherally arranged chromatin. The schizonts also contain numerous cytomers also with trilaminar membranes and containing multiple ribosomes. Repeated invagination of the cytomeres gives rise to the merozoites which also have a trilaminar membrane.The merozoites have rhoptries, micronemes and three apical rings. The mitochondrion contains vesicular cristae. There may be one or two paranuclear bodies in some species. Their function is unknown.
Two gametocyte forms are recognised: an elongated form and a compact spherical form. These are usually 12-14 micrometres long. The large gametocytes tend to grossly distort the infected cells and make cell identification difficult. A pseudopigment known as volutin may be present.
Evolution
It has been suggested that this genus arose in the late Oligocene or early Eocene at about the same time as Piciformes and Coraciiformes.Taxonomy
The genus is divided into two subgenera: Akiba and Leucocytozoon — based on the vector species.The only known member of the subgenus Akiba is Leucocytozoon caulleryi which uses members of the genus Culicoides as its vectors.
The remaining species in the genus use members of the genus Simulium as their vectors.
In 1977, Greiner and Kocan in an extensive examination of species in the order Falconiformes declared that the only valid species infecting this order was L. toddi.
L. dubreuili is considered to be restricted to the family Turdidae; L. fringillinarum is considered to be restricted to several passiform families; and L. majoris is considered restricted to the family Paridae.
L. ziemanni infects owls.
Diagnostic criteria
Form gamonts in white blood cells and/or erythrocytes. Gametocytes cause marked enlargement and distortion of the infected cell producing a football-like appearance.No merogony occurs in either leucocytes or erythrocytes.
Merogony occurs in the parenchyma of liver, heart, kidney, or other organs. Meronts may form large bodies divided into cytomeres.
Hemozoin deposits are not formed — a useful distinguishing feature for Leucocytozoon from Haemoproteus and Plasmodium.
Oocysts develop rapidly in 3–5 days. The oocysts are small and nonexpanding, reaching 13 micrometres in diameter and typically have less than 100 short, thick sporozoites.
The vectors are Simulium or Culicoides species.
The vertebrate hosts are birds.
Type species: Leucocytozoon ziemanni
Pathology
The typical pathology of infection with these parasites includes anaemia and enlargement of the liver and spleen. Gross lesions also include pulmonary congestion and pericardial effusion.Megaloschizonts appear as grey-white nodules found in the heart, liver, lung or spleen. Microscopically there is ischemic necrosis and associated inflammation in the heart, brain, spleen and liver due to occlusion of blood vessels by megaloschizonts in endothelial cells. Ruptured schizonts may induce granulomatous reactions in the surrounding tissues.
Clinically the majority of birds affected with leucocytozoonosis exhibit no signs. Among those that do the signs include mild to severe signs of anorexia, ataxia, weakness, anemia, emaciation and difficulty breathing.
The excess mortality due to Leucocytozoon in adult birds seems to occur as a result of debilitation and increased susceptibility to secondary infection.
Epidemiology
L. simondi is suspected to be a major parasite of Canada geese in some areas, including the upper Midwestern United States and Canada. L. smithi affects turkey farms in the southeastern United States.Host range
Bird hosts- L. anatis — ducks
- L. andrewsi — chicken
- L. anseris — geese
- L. artamidis — wood-swallows
- L. atkinsoni — common jery
- L. balmorali — shrikes, bushshrikes
- L. bennetti — hook-billed vanga
- L. berestneffi — yellow-billed magpie
- L. bishopi — parrotbills
- L. bonasae — ruffed grouse, spruce grouse, ptarmigan
- L. bouffardi — weaver birds
- L. brimonti — bulbuls
- L. caprimulgi — nightjars
- L. caulleryi Mathis and Leger 1909 — chicken
- L. coracinae — swifts, cuckoo-shrikes
- L. danilewskyi — owl
- L. deswardti — Sugarbirds
- L. dubreuili — American robin
- L. dutoiti — finches
- L. frascai — rufous-headed ground-roller
- L. fringillinarum — finches, dark-eyed Juncos, white winged crossbills, yellow wagtail, house sparrow
- L. gentili — house sparrow
- L. greineri — common sunbird asity, velvet asity
- L. grusi — sandhill crane
- L. hamiltoni — Bukharan great tit
- L. lairdi — blue vanga
- L. liothricis — Old World babblers
- L. lovati — greater sage-grouse, blue grouse, willow ptarmigan, wild rock ptarmigan
- L. maccluri — Thailand thrush
- L. macleani Sambon 1908 — chickens
- L. majoris — yellow wagtails
- L. marchouxi — Mauritian pink pigeon, doves
- L. nectariniae — sunbird
- L. neavei — guinea fowl
- L. pogoniuli — tinker barbets
- L. pycnonoti — bulbuls
- L. roubaudi — estrildid finches
- L. sabrazesi Mathis and Leger 1910 — chicken
- L. schoutedeni Rodhain, Pons, Vandenbranden and Bequaert 1913 — chicken
- L. schufneri Prowazek 1912- chicken
- L. shaartusicum — redstart
- L. simondii — ring-necked ducks, wood duck, white Pekin duck, Mexican duck, mallard duck, black duck, geese, Emperor Goose, mergansers, trumpeter swans, eider duck
- L. smithi — turkeys
- L. sturni — starlings
- L. tawaki — penguins
- L. timaliae — Old World babblers
- L. toddi — Coopers hawk, sharp-shinned hawks, Eurasian buzzard, Red-shouldered hawks, white-tailed sea eagle, bald eagles, Chimango Caracara, grouse
- L. trachyphoni — barbet
- L. ziemanni — eagle owl, great horned owls, snowy owls, spotted owls
- L. zosteropis — white-eyes
Vectors
- L. bonasae — Simulium aureum, Simulium latipes
- L. caulleryi — Culicoides arakawae
- L. lovati — Prosimulium hirtipes, Simulium japonicum, Simulium uchidai
- L. simondi — Cnephia ornithophilia, Simulium rugglesi
- L. smithi — Simulium nigritarse
- L. tawaki — Austrosimulium ungulatum
History