Plasmodium

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Red blood cell infected with malaria
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Red blood cell infected with malaria
A plasmodium[[1]] is also the macroscopic form of the unusual protozoa known as slime moulds.
iPlasmodium

Scientific classification
Kingdom: Protista
Phylum: Apicomplexa
Class: Aconoidasida
Order: Haemosporida
Family: Plasmodiidae
Genus: Plasmodium
Species

Plasmodium achiotense
Plasmodium aegyptensis
Plasmodium aeuminatum
Plasmodium agamae
Plasmodium anasum
Plasmodium atheruri
Plasmodium azurophilum
Plasmodium balli
Plasmodium bambusicolai
Plasmodium basilisci
Plasmodium berghei
Plasmodium brasilianum
Plasmodium booliati
Plasmodium brucei
Plasmodium bubalis
Plasmodium bucki
Plasmodium canis
Plasmodium carinii
Plasmodium coatneyi
Plasmodium cathemerium
Plasmodium cephalophi
Plasmodium chabaudi
Plasmodium chiricahuae
Plasmodium cinemidophori
Plasmodium circularis
Plasmodium circumflexum
Plasmodium clelandi
Plasmodium cnemidophori
Plasmodium coatnexi
Plasmodium coggeshalli
Plasmodium colombiense
Plasmodium corradettii
Plasmodium coturnix
Plasmodium coulangesi
Plasmodium cuculus
Plasmodium cyclopsi
Plasmodium cynomolgi
Plasmodium diminutivum
Plasmodium diploglossi
Plasmodium dissanaikei
Plasmodium dominicana
Plasmodium draconis
Plasmodium durae
Plasmodium egerniae
Plasmodium elongatum
Plasmodium eylesi
Plasmodium equi
Plasmodium fabesia
Plasmodium fairchildi
Plasmodium falciparum
Plasmodium fallax
Plasmodium fieldi
Plasmodium foleyi
Plasmodium forresteri
Plasmodium floridense
Plasmodium fornosanum
Plasmodium fragile
Plasmodim gabaldoni
Plasmodium garnhami
Plasmodium gallinaceum
Plasmodium giganteum
Plasmodium giovannolai
Plasmodium girardi
Plasmodium gonatodi Plasmodium gonderi
Plasmodium georgesi
Plasmodium gracilis
Plasmodium griffithsi
Plasmodium guanggong
Plasmodium gundersi
Plasmodium guyannense
Plasmodium heischi
Plasmodium hegneri
Plasmodium hermani
Plasmodium heteronucleare
Plasmodium hexamerium
Plasmodium holaspi
Plasmodium huffi
Plasmodium hylobati
Plasmodium icipeensis
Plasmodium inopinatum
Plasmodium inui
Plasmodium jefferi
Plasmodium juxtanucleare
Plasmodium kempi
Plasmodium kochi
Plasmodium knowlesi
Plasmodium kentropyxi
Plasmodium leanucteus
Plasmodium lemuris
Plasmodium lophurae
Plasmodium lepidoptiformis
Plasmodium lygosomae
Plasmodium mabuiae
Plasmodium mackerrasae
Plasmodium maculilabre
Plasmodium malariae
Plasmodium marginatum
Plasmodium matutinum
Plasmodium mexicanum
Plasmodium minasense
Plasmodium morulum
Plasmodium nucleophilium
Plasmodium octamerium
Plasmodium odocoilei
Plasmodium ovale
Plasmodium papernai
Plasmodium paranucleophilum
Plasmodium parvulum
Plasmodium pedioecetii
Plasmodium pelaezi
Plasmodium percygarnhami
Plasmodium petersi
Plasmodium pifanoi
Plasmodium pinotti
Plasmodium pinorrii
Plasmodium pitheci
Plasmodium pitmani
Plasmodium polare
Plasmodium praecox
Plasmodium reichenowi
Plasmodium relictum
Plasmodium rhadinurum
Plasmodium rhodaini
Plasmodium robinsoni
Plasmodium rouxi
Plasmodium sandoshami
Plasmodium sasai
Plasmodium schweitzi
Plasmodium silvaticum
Plasmodium simium
Plasmodium semiovale
Plasmodium shortii
Plasmodium subpraecox
Plasmodium tenue
Plasmodium tejerai
Plasmodium torrealbai
Plasmodium traguli
Plasmodium tribolonoti
Plasmodium tropiduri
Plasmodium uilenbergi
Plasmodium watteni
Plasmodium vacuolatum
Plasmodium vastator
Plasmodium vaughani
Plasmodium vinckei
Plasmodium vivax
Plasmodium volans
Plasmodium yoelii
Plasmodium youngi
etc.

Plasmodium is a genus of parasitic protozoa. Infection with this genus is known as malaria. The parasite always has two hosts in its life cycle: a mosquito vector and a vertebrate host. At least ten species infect humans. Other species infect other animals, including birds, reptiles and rodents.

There are over 175 species recognised in this genus. The genus is currently (2006) in need of reorganisation as it has been shown that other parasites belonging to the genera Haemocystis and Hepatocystis appear to be closely related to this genus. It is likely that other species such as Haemoproteus meleagridis will be included in this genus once it is revised.

Host range amoung the mammalian orders is non uniform. At least 25 species infect primates; rodents outside the tropical parts of Africa are rarely affected; a few species are known to infect bats, porcupines and squirrels; while carnivores, insectivores and marsupials are not known to act as hosts.

Contents

[edit] Discovery of the life cycle

In 1898 Ronald Ross demonstrated the existence of Plasmodium in the stomach of the Anopheles mosquito. For this discovery he won the Nobel Prize in 1902. However some credit must also be given to the Italian professor Giovanni Battista Grassi, who showed that human malaria could only be transmitted by Anopheles mosquitoes.

The species of Plasmodium that parasitise humans include:

The first four listed here are the most common species that infect humans. With the use of the polymerase chain reaction additional species have been and are still being identified that infect humans.

[edit] Taxonomy

Plasmodium belongs to the order Haemosporidia. The taxonomy is shown bwlow.

  • Order Haemosporida


  • Family Haemoproteidae
  • Genus Haemoproteus
  • Subgenus Parahaemoproteus
  • Subgenus Haemoproteus


  • Family Plasmodiidae
  • Genus Plasmodium
  • Subgenus Asiamoeba
  • Subgenus Bennettinia
  • Subgenus Carinamoeba
  • Subgenus Giovannolaia
  • Subgenus Haemamoeba
  • Subgenus Huffia
  • Subgenus Laverania
  • Subgenus Novyella
  • Subgenus Paraplasmodium
  • Subgenus Sauramoeba
  • Subgenus Vinckeia


  • Family Garniidae
  • Genus Fallisia
  • Subgenus Plasmodioides


  • Family Leucocytozoidae
  • Genus Leucocytozoon
  • Subgenus Leucocytozoon
  • Subgenus Akiba


Two other subgenera have been described in Plasmodium but are no longer used: Lacertamoeba and Ophidiella

[edit] Subgenera

The full taxonomic name of a species includes the subgenus but this is often omitted. The full name indicates some features of the morphology and type of host species. Examples of the use of full names include:

Plasmodium (Asiamoeba) draconis
Plasmodium (Asiamoeba) vastator

Plasmodium (Carinamoeba) basilisci
Plasmodium (Carinamoeba) minasense
Plasmodium (Carinamoeba) rhadinurum
Plasmodium (Carinamoeba) volans

Plasmodium (Giovannolaia) circumflexum
Plasmodium (Giovannolaia) dissanaikei
Plasmodium (Giovannolaia) durae
Plasmodium (Giovannolaia) fallax
Plasmodium (Giovannolaia) garnhami
Plasmodium (Giovannolaia) gundersi
Plasmodium (Giovannolaia) lophurae
Plasmodium (Giovannolaia) pedioecetii
Plasmodium (Giovannolaia) polare

Plasmodium (Haemamoeba) cathemerium
Plasmodium (Haemamoeba) coggeshalli
Plasmodium (Haemamoeba) gallinacium
Plasmodium (Haemamoeba) giovannolai
Plasmodium (Haemamoeba) matutinum
Plasmodium (Haemamoeba) relictum
Plasmodium (Haemamoeba) subpraecox
Plasmodium (Haemamoeba) tejera

Plasmodium (Huffia) elongatum
Plasmodium (Huffia) hermani

Plasmodium (Laverania) falciparum
Plasmodium (Laverania) reichenowi

Plasmodium (Novyella) bambusicolai
Plasmodium (Novyella) corradettii
Plasmodium (Novyella) hexamerium
Plasmodium (Novyella) juxtanucleare
Plasmodium (Novyella) kempi
Plasmodium (Novyella) nucleophilum
Plasmodium (Novyella) papernai
Plasmodium (Novyella) rouxi
Plasmodium (Novyella) vaughani

Plasmodium (Paraplasmodium) mexicanum
Plasmodium (Paraplasmodium) chiricahuae

Plasmodium (Plasmodium) brasilianum
Plasmodium (Plasmodium) cynomolgi
Plasmodium (Plasmodium) eylesi
Plasmodium (Plasmodium) fragile
Plasmodium (Plasmodium) georgesi
Plasmodium (Plasmodium) gonderi
Plasmodium (Plasmodium) inui
Plasmodium (Plasmodium) jefferyi
Plasmodium (Plasmodium) knowlei
Plasmodium (Plasmodium) hyobati
Plasmodium (Plasmodium) malariae
Plasmodium (Plasmodium) ovale
Plasmodium (Plasmodium) petersi
Plasmodium (Plasmodium) pitheci
Plasmodium (Plasmodium) rhodiani
Plasmodium (Plasmodium) schweitzi
Plasmodium (Plasmodium) semiovale
Plasmodium (Plasmodium) shortii
Plasmodium (Plasmodium) silvaticum
Plasmodium (Plasmodium) simium
Plasmodium (Plasmodium) vivax
Plasmodium (Plasmodium) youngi

P[asmndium (Sauramoeba)achiotense
Plasmodium (Sauramoeba) aeuminatum
Plasmodium (Sauramoeba) giganteum
Plasmodium (Sauramoeba) heischi
Plasmodium (Sauramoeba) pelaezi
Plasmodium (Sauramoeba) tropiduri

Plasmodium (Vinckeia) atheruri
Plasmodium (Vinckeia) berghei
Plasmodium (Vinckeia) booliati
Plasmodium (Vinckeia) brucei
Plasmodium (Vinckeia) bubalis
Plasmodium (Vinckeia) bucki
Plasmodium (Vinckeia) cephalophi
Plasmodium (Vinckeia) chabaudi
Plasmodium (Vinckeia) coulangesi
Plasmodium (Vinckeia) foleyi
Plasmodium (Vinckeia) girardi
Plasmodium (Vinckeia) lemuris
Plasmodium (Vinckeia) percygarnhami
Plasmodium (Vinckeia) sandoshami
Plasmodium (Vinckeia) uilenbergi
Plasmodium (Vinckeia) vinckei
Plasmodium (Vinckeia) watteni
Plasmodium (Vinckeia) yoelli


  • Notes


Species of Novyella are characterized by very small schizonts. Sexual stages in this subgenus resemble those of Haemoproteus.

The only two species in the sub genus Laverania and P. falciparum and P. reichenowi.

[edit] Infections in primates

The species that infect primates other than humans include: P. brasilianum, P. bucki, P. coulangesi, P. cynomolgi, P. coatneyi, P. fieldi, P. foleyi, P. fragile, P. girardi, P. georgesi, P. gonderi, P. hylobati, P. inui, P. knowlesi, P. lemuris, P. percygarnhami, P. petersi, P. reichenowi, P. rodhaini, P. sandoshami, P shortii, P. silvaticum, P. simiovale, P. simium and P. uilenbergi.


  • Host record


Most if not all Plasmodium species infect more than one host: the host records shown here should be regarded as being incomplete.


P. bucki - Lemur macaco macaco

P. coatneyi - the macaque (Macaca irus).

P. coulangesi - Lemur macaco macaco

P. foleyi - Lemur fulvus rufus

P. fragile - the macaque Macaca sinica

P. georgesi - Cercocebus albigena

P. girardi - Lemur fulvus rufus

P. gonderi - Cercocebus albigena

P. hylobati - gibbons.

P. knowlesi - the macaque Macaca fascicularis

P. knowlesi edesoni - the Javanese long-tailed macaque (Macaca irus)

P. lemuris - the lemur Lemur collaris

P. percygarnhami - Lemur macaco macaco

P. petersi - Cercocebus albigena

P. pitheci - orangutans

P. reichenowi - chimpanzees.

P. sandoshami - the Malayan flying lemur (Cynocephalus variegatus)

P. semiovale - Macaca sinica

P. schwetzi - gorillas.

P. uilenbergi - Lemur fulvus fulvus


  • Mosquito vectors


  • Subspecies

P. cynomolgi - P. cynomolgi bastianelli and P. cynomolgi ceylonensis.

P. inui - P. inui inui and P. inui shortii

P. knowlesi - P. knowlesi edesoni and P. knowlesi knowlesi.

P. vivax - P. vivax hibernans, P. vivax chesson and P. vivax multinucleatum.


  • Interrelatedness

The evolution of these species is still being worked out and the relationships given here should be regarded as tentative.


P. brasilianum and P. rodhaini are very similar to P. malariae

P. cynomolgi, P. fragile, P. simium and P. schwetzi are similar to P. vivax

P. fieldi and P. simiovale are similar to P. ovale

P. falciparum is closely related to P. reichenowi.

This last grouping while originally made on morphological grounds now has considerable support at the DNA level.

[edit] Infections in non primate mammals

The subgenus Vinckeia was created by Garnham to accomadate the mammalian parasites other than those infecting primates. Species infecting lemurs have also been incluced in this subgenus.

P. bergei, P. chabaudi, P. yoelli and P. vinckei infect rodents and have been used to study malarial infections in the laboratory. Other members of this subgenus infect other mammalian hosts.


  • Host records

P. booliati - Malayan giant flying squirrel [1]

P. cephalophi - the grey duiker (Sylvicapra grimmia) [2]

P. watteni - Formosan giant flying squirrel (Petaurista petaurista grandis)[3]

P. atheruri - the African porcupine (Atherurus africanus).


  • Subspecies


P. yoellii - P. yoelli nigeriensis and P. yoelli yoelli.

P. vinkei - P. vinckei brucechwatti, P. vinckei petteri and P. vinckei vinckei.

P. chabaudi - P. chabaudi adami and P. chabaudi chabaudi

[edit] Infections in birds

34 species in five of these subgenera infect birds - Bennettinia, Giovannolaia, Haemamoeba, Huffia and Novyella. [4] Giovannolaia appears to be a polyphytic group and may be sudivided in the future. [5]

Species infecting birds include: P. anasum, P. bambusicolai, P. cathemerium, P. circumflexum, P. corradettii, P. coturnix, P. dissanaikei, P. durae, P. elongatum, P. fallax, P. fornosanum, P. gallinacium, P. giovannolai, P. griffithsi, P. gabaldoni, P. gundersi, P. hegneri, P. hermani, P. hexamerium, P. juxtanucleare, P. kempi, P. lophurae, P. matutinum, P. nucleophilum, P. papernai, P. pediocetti, P. pinotti, P. polare, P. praecox, P. relictum, P. rouxi, P. tejerai and P. vaughani.


  • Subspecies


P. relictum has been divded into two subspecies: P. relictum relictum and P. relictum capistranoae.


  • Notes

Plasmodium relictum is probably responsible for more bird extinctions than any other protist.

[edit] Infections in reptiles

Species in the subgenera Carinamoeba, Paraplasmodium and Sauramoeba infect reptiles. [6]


  • Host records


P. heischi - skinks (Mabuya striata) [7]

P. giganteum - Agama cyanogaster. [8]

P. siamense - lizards. [9]

P. tribolonoti - skinks

P. gracilis - skinks

P. lygosomae - skink (Lygosoma moco)


  • Subspecies


P. minasense - P. minasense minasense, P. minasense carinii, P. minasense anolisi, P. minasense capitoi, P. minasense plicae, P. minasense tegui and P. minasense diminutivum. [10] An additional subspecies P. minasense calcaratae has also been decribed. [11]

P. traguli - P. traguli traguli and P. traguli memmina.

P. tropiduri - P. tropiduri tropiduri, P. tropiduri panamense and P. tropiduri aquaticum. [10]

P. lygosomae - P. lygosomae nucleoversans.

[edit] Life cycle

The life cycle of Plasmodium is very complex. Sporozoites from the saliva of a biting female mosquito are transmitted to either the blood or the lymphatic system[12] of the recipient. Mosquitoes of the genera Culex, Anopheles, Culiceta, Mansonia and Aedes may act as vectors. The currently known vectors for human malaria (> 100 species) all belong to the genus Anopheles. The life cycle of Plasmodium was discovered by Ross who worked with species from the genus Culex.

The sporozoites migrate to the liver and invade hepatocytes. The so-called latent or dormant stage of the Plasmodium sporozoite in the liver is called the hypnozoite. From the hepatocytes, the parasite replicates into thousands of merozoites, which then invade red blood cells. Here the parasite grows from a ring-shaped form to a larger trophozoite form. In the schizont stage, the parasite divides several times to produce new merozoites, which leave the red blood cells and travel within the bloodstream to invade new red blood cells. Most merozoites continue this replicative cycle, but some merozoites differentiate into male or female sexual forms (gametocytes) (also in the blood), which are taken up by the female Anopheles mosquito. In the mosquito's midgut, the gametocytes develop into gametes and fertilize each other, forming motile zygotes called ookinetes. The ookinetes penetrate and escape the midgut, then embed themselves onto the exterior of the gut membrane. Here they divide many times to produce large numbers of tiny elongated sporozoites. These sporozoites migrate to the salivary glands of the mosquito where they are injected into the blood of the next host the mosquito bites. The sporozoites move to the liver where they repeat the cycle.

This life cycle is best understood in terms of its evolution. It is thought that Plasmodium evolved from a parasite spread by the orofaecal route which infected the intestinal wall. At some point this parasite evolved the ability to infect the liver. This pattern is seen in the genus Cryptosporidium to which Plasmodium is distantly related. At some later point this ancestor developed the ability to infect blood cells and to survive and infect mosquitoes. Once mosquito transmission was firmly established the previous orofecal route of transmission was lost.

[edit] Molecular biology

On a molecular level, the parasite damages red blood cells using plasmepsin enzymes. Plasmepsins are aspartic acid proteases which degrade hemoglobin.

[edit] Notes

  1. ^ Sandosham AA, Yap LF, Omar I. (1965) A malaria parasite, plasmodium (Vinckeia) booliati sp.nov., from a Malayan giant flying squirrel. Med J Malaya. 20(1):3-7
  2. ^ Keymer IF. (1966) Studies on Plasmodium (Vinckeia) cephalophi of the grey duiker (Sylvicapra grimmia). Ann Trop Med Parasitol. 60(2):129-138
  3. ^ Lien JC, Cross JH. (1968) Plasmodium (Vinckeia) watteni sp. n. from the Formosan giant flying squirrel, Petaurista petaurista grandis. J Parasitol. 54(6):1171-1174
  4. ^ Wiersch SC, Maier WA, Kampen H. Plasmodium (Haemamoeba) cathemerium gene sequences for phylogenetic analysis of malaria parasites. Parasitol Res. 96(2): 90-94
  5. ^ Martinsen ES,Waite JL,Schall JJ Morphologically defined subgenera of Plasmodium from avian hosts: test of monophyly by phylogenetic analysis of two mitochondrial genes (2006) Parasitology 1 - 8
  6. ^ Schall JJ (2000) Transmission success of the malaria parasite Plasmodium mexicanum into its vector: role of gametocyte density and sex ratio. Parasitology. 121 (6):575-580
  7. ^ Garnham PC, Telford SR Jr. (1984) A new malaria parasite Plasmodium (Sauramoeba) heischi in skinks (Mabuya striata) from Nairobi, with a brief discussion of the distribution of malaria parasites in the family Scincidae. J Protozool. 31(4):518-521.
  8. ^ Southgate BA. (1970) Plasmodium (Sauramoeba) giganteum in Agama cyanogaster: a new host record. Trans R Soc Trop Med Hyg. 64(1):12-13
  9. ^ Telford SR (1986) Fallisia parasites (Haemosporidia: Plasmodiidae) from the flying lizard, Draco maculatus (Agamidae) in Thailand. J Parasitol. 72(5):766-769
  10. ^ a b Telford SR Jr. (1979) A taxonomic revision of small neotropical saurian Malarias allied to Plasmodium minasense. Ann Parasitol Hum Comp. 54(4):409-422
  11. ^ Telford SR Jr and Telford SR 3rd. Rediscovery and redescription of Plasmodium pifanoi and description of two additional Plasmodium parasites of Venezuelan lizards. Journal of Parasitology (2003) 89(2):362-368
  12. ^ [http://www.hhmi.org/news/menard20060122.html HHMI Staff (22 January 2006) "Malaria Parasites Develop in Lymph Nodes" HHMI News Howard Hughes Medical Institute

[edit] References

[edit] External links