**Acknowledgement**

The authors thank Aurélien Dumètre, René Ecochard, Michel Langlais, Dominique Pontier, Philippe Thulliez and Stéphane Romand for their help in elaborating the 10-year research period that produced part of the results presented here. This project has been supported by the Agence Française de Sécurité Sanitaire de L'Environnement et du Travail (AFSSET) and by the Agence De l'Environnement et de la Maitrise de l'Energie (ADEME), with additional grants from Grünenthal France Laboratory (EA), Institut National de la Recherche Agronomique (INRA, CR), Région Champagne-Ardenne (EA, ML and CG), Département des Ardennes (ML), Communauté de Communes de l'Argonne Ardennaise (ML) and the National Institute for Mathematical and Biological Synthesis (NIMBioS, sponsored by the National Science Foundation, the U.S. Department of Homeland Security, and the U.S. Department of Agriculture, ML).

#### **7. References**

[1] Dubey JP (2010) Toxoplasmosis of animals and humans, second edition. Boca Raton: CRC Press. 313 p.

<sup>\*</sup> Corresponding Author

[2] Tenter AM, Heckeroth AR, Weiss LM (2000) *Toxoplasma gondii*: from animals to humans. Int. j. parasitol. 30: 1217-1258.

The Life Cycle of *Toxoplasma gondii* in the Natural Environment 25

[20] Liberg O (1980) Spacing pattern in a population of rural free-roaming domestic cats.

[21] Pontier D, Rioux N, Heizmann A (1995) Evidence of selection on the orange allele in the

[22] Fromont E, Artois M, Pontier D (1998) Epidemiology of feline leukemia virus (FeLV)

[23] Liberg O (1981) Predation and social behaviour in a population of domestic cats. An

[24] Liberg O (1984) Food habits and prey impact by feral and house-based domestic cats in

[25] Say L, Gaillard JM, Pontier D. (2002) Spatiotemporal variation in cat population density

[26] Say L, Devillard S, Natoli E, Pontier D (2002) The mating system of feral cats (*Felis catus* 

[27] Le Louarn H, Quéré JP (2003) Les Rongeurs de France. Faunistique et biologie. INRA

[28] Dickman CR, Doncaster CP (1987) The Ecology of Small Mammals in Urban Habitats. I.

[29] Baker PJ, Ansell RJ, Dodds PAA, Webber CE, Harris S (2003) Factors affecting the

[30] Dubey JP, Saville WJA, Stanek J, Reed SM (2002) Prevalence of *Toxoplasma gondii*

[31] Miró G, Montoya A, Jimenez S, Frisuelos C, Mateo M, Fuentes I (2004) Prevalence of antibodies to *Toxoplasma gondii* and intestinal parasites in stray, farm and household

[32] Afonso E, Thulliez P, Gilot-Fromont E (2010) Local meteorological conditions, dynamics of seroconversion to *Toxoplasma gondii* in cats (*Felis catus*) and oocyst burden in a rural

[33] Wu SM, Zhu XQ, Zhou DH, Fu BQ, Chen J, Yang JF, Song HQ, Weng YB, Ye DH (2011) Seroprevalence of *Toxoplasma gondii* infection in household and stray cats in Lanzhou,

[34] Zhang H, Zhou DH, Zhou P, Lun ZR, Chen XG, Lin RQ, Yuan ZG, Zhu XQ (2009) Seroprevalence of *Toxoplasma gondii* infection in stray and household cats in

[35] Barratt DG (1998) Predation by house cats, *Felis catus* (L.), in Canberra, Australia. II. Factors affecting the amount of prey caught and estimates of the impact on wildlife.

[36] Baker PJ, Bentley AJ, Ansell RJ, Harris S (2005) Impact of predation by domestic cats

[37] Woods M, McDonald RA, Harris S (2003) Predation of wildlife by domestic cats *Felis* 

[38] Gillies C, Clout M (2003) The prey of domestic cats (*Felis catus*) in two suburbs of

distribution of small mammals in an urban area. Mammal rev. 33: 95-100.

antibodies in domestic cats From rural Ohio. J. parasitol. 88: 802-803.

domestic cat *Felis catus*: the role of social structure. Oikos 73: 299-308.

evolutionary perspective. Ph. D. Thesis, Univ. Lund (Sweden), 135 p.

a rural area in southern Sweden. J. mammal. 65: 424-432.

L.) in a sub-Antarctic environment. Polar biol. 25: 838–842.

Populations in a Patchy Environment. J. anim. ecol. 56: 629-640.

in a sub-Antarctic environment. Polar biol. 25: 90–95.

cats in Spain. Vet. parasitol. 126: 249-255.

environment. Epidemiol. infect. 138: 1105–1113.

northwest China. Parasites & Vectors 4: 214-218.

Guangzhou, China. Zoonoses public health 56: 502-505.

*Felis catus* in an urban area. Mammal rev. 35: 302-312.

*catus* in Great Britain. Mammal rev. 33: 174–188.

Auckland City, New Zealand J. zool. 259: 309-315.

and structure of domestic cat populations. J. wildl. manage. 62: 978-988.

Oikos 35: 336-349.

éditions, Paris, 256pp.

Wildlife res. 25: 475-487.


[20] Liberg O (1980) Spacing pattern in a population of rural free-roaming domestic cats. Oikos 35: 336-349.

24 Toxoplasmosis – Recent Advances

humans. Int. j. parasitol. 30: 1217-1258.

otters. Int. j. parasitol 58: 928-937.

of males. PLoS ONE 6: e27229.

generations of mice. Nature 183: 1348-1349.

toxoplasmosis. Int. j. parasitol. 39: 1385-1394.

(United Kingdom). Rev. sci. tech. OIE 12: 23-26.

and supplemental feeding. J. anim. ecol. 58: 321-328.

Cambridge University Press.

of *Echinococcus multilocularis*. Trends parasitol 20 : 77-84.

[2] Tenter AM, Heckeroth AR, Weiss LM (2000) *Toxoplasma gondii*: from animals to

[3] Miller MA, Miller WA, Conrad PA, James ER, Melli AC, Leutenegger CM, Dabritz HA, Packham AE, Paradies D, Harris M, Ames J, Jessup DA, Worcester K, Grigg ME (2008) Type X *Toxoplasma gondii* in a wild mussel and terrestrial carnivores from coastal California: new linkages between terrestrial mammals, runoff and toxoplamosis of sea

[4] Deplazes P, Hegglin D, Gloor S, Romig T (2004) Wilderness in the city: the urbanization

[5] House PK, Vyas A, Sapolsky R (2011) Predator cat odors activate sexual arousal

[6] Dass SAH, Vasudevan A, Dutta D, Soh LJT, Salposky RM, Vyas A (2011) Protozoan parasite *Toxoplasma gondii* manipulates mate choice in rats by enhancing attractiveness

[7] Cook AJC, Gilbert RE, Buffolano W, Zufferey J, Petersen E, Jenum PA, Foulon W, Semprini AE, Dunn DT (2000) Sources of *Toxoplasma* infection in pregnant women:

[8] Beverly JKA (1959) Congenital transmission of toxoplasmosis through successive

[9] Dubey JP (2006) Comparative infectivity of oocysts and bradyzoites of *Toxoplasma gondii*

[10] Lélu M, Langlais M, Poulle ML, Gilot-Fromont E (2010) Transmission dynamics of *Toxoplasma gondii* along an urban-rural gradient. Theor. popul. biol. 78: 139–147. [11] Berenreiterova M, Flegr J, Kubena AA, Nemec P (2011) The Distribution of *Toxoplasma gondii* Cysts in the Brain of a Mouse with Latent Toxoplasmosis: Implications for the

[12] Pappas G, Roussos N, Falagas ME (2009) Toxoplasmosis snapshots: Global status of *Toxoplasma gondii* seroprevalence and implications for pregnancy and congenital

[13] Berger F, Goulet V, Le Strat Y, Desenclos JC (2008) Toxoplasmose chez les femmes enceintes en France : évolution de la séroprévalence et de l'incidence et facteurs

[17] Macdonald DW (1983) The ecology of carnivore social behaviour. Nature 301: 379-384. [18] Page RJC, Ross J, Bennett DH (1993) Home ranges of feral cats at Avonmouth Docks

[19] Calhoon RE, Haspel C (1989) Urban cat populations compared by season, subhabitat

associés, 1995-2003. Bulletin Epidemiologique Hebdomadaire 14-15: 117-121. [14] Gilot-Fromont E, Riche B, Rabilloud M (2009) Seroprevalence of *Toxoplasma gondii* in a rural population in France: detection of a household effect. BMC infect. dis. 9: 76. [15] Haydon DT, Cleaveland S, Taylor LH, Laurenson K (2002) Identifying Reservoirs of Infection: A Conceptual and Practical Challenge. Emerg. infect. dis. 8: 1468-1473. [16] Turner DC, Bateson PB (2000) The Domestic cat: the biology of its behaviour.

for intermediate (mice) and definitive (cats) hosts. Vet. parasitol. 140: 69-75.

pathways in brains of *Toxoplasma gondii* infected rats. PLoS ONE 6: e23277.

European multicentre case-control study. Br. med. j. 321: 142-147.

Behavioral Manipulation Hypothesis. PLoS One 6: e28925.


[39] Kays RW, DeWan AA (2004) Ecological impact of inside/outside house cats around a suburban nature reserve. Anim. conserv. 7: 273-283.

The Life Cycle of *Toxoplasma gondii* in the Natural Environment 27

[55] Bradley CA, Altizer S (2007) Urbanization and the ecology of wildlife diseases. Trends

[56] Zaunbrecher KI, Smith RE (1993) Neutering of feral cats as an alternative to eradication

[57] Neville PF, Remfry J (1984) Effect of neutering on two groups of feral cats. Vet. rec. 114:

[58] Natoli E (1985) Spacing pattern in a colony of urban stray cats (*Felis catus* L.) in the

[59] Haspel C, Calhoon RE (1989) Home ranges of free ranging cats (*Felis catus*) in Brooklyn,

[60] Salant H, Spira DT (2004) A cross-sectional survey of anti-*Toxoplasma gondii* antibodies

[61] Jittapalapong S, Nimsupan B, Pinyopanuwat N., Chimnoi W, Kabeya H, Maruyama S (2006) Seroprevalence of *Toxoplasma gondii* antibodies in stray cats and dogs in the

[62] Pena HJF, Soares RM, Amaku M, Dubey JP, Gennari SM (2006) *Toxoplasma gondii* infection in cats from Saõ Paulo state, Brazil: seroprevalence, oocyst shedding, isolation

[63] Gauss CBL, Almeria S, Ortuno A, Garcia F, Dubey JP (2003) Seroprevalence of *Toxoplasma gondii* Antibodies in Domestic Cats from Barcelona, Spain. J. parasitol. 89:

[64] Dorny P, Speybroeck N, Verstraete S, Baeke M, De Becker A, Berkvens D, Vercruysse J (2002) Serological survey of *Toxoplasma gondii*, feline immunodeficiency virus and feline

[65] Corbett LK (1979) Feeding ecology and social organization of wildcats (*Felis silvetris*)

[66] Uga S, Minami T, Nagata K (1996) Defecation habits of cats and dogs and contamination by *Toxocara* eggs in public park sandpits. Am. j. trop. med. hyg. 54: 122-

[67] Lass A, Pietkiewicz H, Modzelewska E, Dumètre A, Szostakowska B, Myjak P (2009) Detection of *Toxoplasma gondii* oocysts in environmental soil samples using molecular

[68] Du F, Feng HL, Nie H, Tu P, Zhang QL, Hu M, Zhou YQ, Zhao JL (2012) Survey on the contamination of *Toxoplasma gondii* oocysts in the soil of public parks of Wuhan, China.

[69] Bobić B, Jevremović I, Marinković J, Šibalić D, Djurković-Djaković O (1998) Risk factors for *Toxoplasma* infection in a reproductive age female population in the area of Belgrade,

[70] Kortbeek LM, De Melker HE, Veldhuijzen IK, Conyn-Van Spaendonck MAE (2004) Population-based *Toxoplasma* seroprevalence study in the Netherlands. Epidemiol.

[71] Frenkel JK, Hassanein KL, Hassanein RS, Brown E, Thulliez P, Quintero-Nunez R (1995) Transmission of *Toxoplasma gondii* in Panama city, Panama: A five-year prospective

and domestic cats (*Felis catus*) in Scotland. Scotland: University of Aberdeen.

in mice, and biologic and molecular characterization. Res. vet. sci. 81: 58-67.

leukaemia virus in urban stray cats in Belgium. Vet. rec. 151: 626-629.

methods. Eur. j. clin. microbiol. Infect. dis. 28: 599-605.

ecol. evol. 22: 95-102.

447-450.

1067-1068.

126.

Vet. parasitol. 184: 141-146.

infect. 132: 839-845.

Yugoslavia. Eur. j. epidemiol. 14: 605-610.

programs. J. am. vet. med. assoc. 203: 449-452.

in Jerusalem cats. Vet. parasitol. 124: 167-177.

New York. Can. j. zool. 67: 178-181.

historic centre of Rome. Appl. anim. behav. sci. 14: 289-304.

Bangkok metropolitan area, Thailand. Vet. parasitol. 145: 138-141.


[55] Bradley CA, Altizer S (2007) Urbanization and the ecology of wildlife diseases. Trends ecol. evol. 22: 95-102.

26 Toxoplasmosis – Recent Advances

Acta vet. hung. 56: 81-88.

Parasitology 134: 1963-1971.

parasitol. 38: 1017-1023.

890.

153-155.

[39] Kays RW, DeWan AA (2004) Ecological impact of inside/outside house cats around a

[40] Rosenthal BM (2009) How has agriculture influenced the geography and genetics of

[41] Afonso E, Thulliez P, Gilot-Fromont E (2006) Transmission of *Toxoplasma gondii* in an

[42] Hornok S, Edelhofer R, Joachim A, Farkas R, Berta K, Repasi A, Lakatos B (2008) Seroprevalence of *Toxoplasma gondii* and *Neospora caninum* infection of cats in Hungary.

[43] Lee SE, Kim NH, Chae HS, Cho SH, Nam HW, Lee WJ, Kim SH, Lee JH (2011) Prevalence of *Toxoplasma gondii* infection in feral cats in Seoul, Korea. J. parasitol. 97,

[44] Dubey JP, Weigel RM, Siegel AM, Thuilliez P, Kitron UD, Mitchell MA, Mannelli A, Mateus-Pinilla NE, Shen SK, Kwok OCH, Todd KS (1995) Sources and reservoirs of

[46] Afonso E, Thulliez P, Pontier D, Gilot-Fromont E (2007) Toxoplasmosis in prey species and consequences for prevalence in feral cats: not all prey species are equal.

[47] Dabritz HA, Miller MA, Gadner IA, Packham AE, Atwill ER, Conrad PA (2008) Risk factors for *Toxoplasma gondii* infection in wild rodents from central coastal California

[48] Genchi G, Polidori GA, Zaghini L, Lanfranchi P (1991) Aspetti epidemiologici della toxoplasmosi nell'allevamento intensivo del suino. Arch. vet. ital. 42, 105–111. [49] Marshall PA, Hughes JM, Williams RH, Smith JE, Murphy RG, Hide G (2004) Detection of high levels of congenital transmission of *Toxoplasma gondii* in natural urban

[50] Afonso E, Lemoine M, Poulle ML, Ravat MC, Romand S, Thulliez P, Villena I, Aubert D, Riche B, Rabilloud M, Gilot-Fromont E (2008) Spatial distribution of soil contamination by *Toxoplasma gondii* in relation to cat defecation behaviour in an urban area. Int. j.

[51] Dabritz HA, Miller, Atwill ER, Gardner IA, Leutenegger CM, Melli AC, Conrad PA (2007) Detection of *Toxoplasma gondii*-like oocysts in cat feces and estimates of the

[52] Dabritz HA, Atwill ER, Gardner IA, Miller MA, Conrad PA (2006) Outdoor fecal deposition by free-roaming cats and attitudes of cat owners and nonowners toward

environmental oocyst burden. J. am. vet. med. assoc. 231: 1676–1684.

stray pets, wildlife, and water pollution. J. am. vet. med. assoc. 229: 74-81. [53] Forman RTT, Godron M (1986) Landscape ecology. Wiley and Sons, New York, USA. [54] McKinney ML (2002) Urbanization, biodiversity, and conservation. BioScience 52: 883-

*Toxoplasma gondii* infection on 47 swine farms in Illinois. J. parasitol. 81: 723-729. [45] Cavalcante GT, Aguiar DM, Chiebao D, Dubey JP, Ruiz VLA, Dias RA, Camargo LMA, Labruna MB, Gennari SM (2006) Seroprevalence of *Toxoplasma gondii* antibodies in cats

and pigs from rural Western Amazon, Brazil. J. parasitol. 92: 863-864.

and a review of *T. gondii* prevalence in rodents. J. parasitol. 94: 675–683.

populations of *Mus domesticus*. Parasitology 128: 39-42.

urban population of domestic cats (*Felis catus*). Int. j. parasitol. 36: 1373-1382.

suburban nature reserve. Anim. conserv. 7: 273-283.

animal parasites? Trends parasitol. 25: 67-70.


cohort study of children, cats, rodents, birds, and soil. Am. j. trop. med. hyg. 53: 458- 468.

The Life Cycle of *Toxoplasma gondii* in the Natural Environment 29

[87] Herrmann DC, Pantchev N, Vrhovec MG, Barutzki D, Wilking H, Fröhlich A, Lüder CG, Conraths FJ, Schares G (2010) Atypical *Toxoplasma gondii* genotypes identified in

[88] Klun I, Djurkovic-Djakovic O, Katic-Radivojevic S, Nikolic A (2006) Cross-sectional survey on *Toxoplasma gondii* infection in cattle, sheep and pigs in Serbia: seroprevalence

[89] Panadero R, Painceira A, López C, Vázquez L, Paz A, Díaz P, Dacal V, Cienfuegos S, Fernández G, Lago N, Díez-Baños P, Morrondo P (2010) Seroprevalence of *Toxoplasma gondii* and *Neospora caninum* in wild and domestic ruminants sharing pastures in Galicia

[90] Beral M, Rossi S, Aubert D, Gasqui P, Terrier ME, Klein F, Geers R, Abrial D, Gilot-Fromont E, Richomme C, Hars J, Jourdain E (submitted) Environmental factors associated with the seroprevalence of *Toxoplasma gondii* in wild boar (*Sus scrofa*), France. [91] Richomme C, Afonso E, Tolon V, Ducrot C, Halos L, Alliot A, Perret C, Thomas M, Boireau P, Gilot-Fromont E (2010) Risk factors for *Toxoplasma gondii* infection in wild

boar from the Mediterranean island of Corsica. Epidemiol. infect. 138: 1257–1266. [92] Langlais M, Lélu M, Avenet C, Gilot-Fromont E (2012) A simplified model system for *Toxoplasma gondii* spread within a heterogeneous environment. Nonlinear Dynamics,

[93] Lopes WDZ, dos Santos TR, dos Santos da Silva R, Rossanese WM, de Souza FA, D'Ark de Faria Rodrigues J, Paranhos de Mendonça R, Soares VE, da Costa AJ (2010) Seroprevalence of and risk factors for *Toxoplasma gondii* in sheep raised in the

[94] Mainar RC, de la Cruz C, Asencio A, Dominguez L, Vazquez-Boland IA (1996) Prevalence of agglutination antibodies to *Toxolasma gondii* in small ruminants of the Madrid region, Spain, and identification of factors influencing seropositivity by

[95] Mateus-Pinilla NE, Hannon B, Weigel M (2002) A computer simulation of the prevention of the transmission of *Toxoplasma gondii* on swine farms using a feline *T-*

[96] Kijlstra A, Jongert E (2008) Control of the risk of human toxoplasmosis transmitted by

[97] Gilot-Fromont E, Aubert D, Belkilani S, Hermitte P, Gibout O, Geers R, Villena I (2009) Landscape, herd management and within-herd seroprevalence of *Toxoplasma gondii* in

[99] Cabezón O, García-Bocanegra I, Molina-López R, Marco I, Blanco JM, Höfle U, Margalida A, Bach-Raich E, Darwich L, Echeverría I, Obón E, Hernández M, Lavín S, Dubey JP, Almería S (2011) Seropositivity and risk factors associated with *Toxoplasma*

beef cattle herds from Champagne-Ardenne, France. Vet. parasitol. 161: 36-40. [98] Hill DE, Chirukandoth S, Dubey JP (2005) Biology and epidemiology of *Toxoplasma*

Jaboticabal, microregion, São Paulo State, Brazil. Res. vet. sci 161: 324-326.

oocysts shed by cats in Germany. Int. j. parasitol. 40: 285-292.

and risk factors. Vet. parasitol. 135: 121–131.

(Northwest Spain). Res. vet. sci 88: 111–115.

multivariate analysis. Vet. res. com. 20: 153-159.

*gondii* in man and animals. Anim. health res. rev. 6: 41-61.

*gondii* infection in wild birds from Spain. PloS ONE 6: 1–7.

*gondii* vaccine. Prev. vet. med. 55: 17–36.

meat. Int. j. parasitol. 38: 1359–1370.

doi: 10.1007/s11071-011-0255-4.


[87] Herrmann DC, Pantchev N, Vrhovec MG, Barutzki D, Wilking H, Fröhlich A, Lüder CG, Conraths FJ, Schares G (2010) Atypical *Toxoplasma gondii* genotypes identified in oocysts shed by cats in Germany. Int. j. parasitol. 40: 285-292.

28 Toxoplasmosis – Recent Advances

468.

7.

3: 135–141.

Sweden. Vet. parasitol. 177: 275-280.

in Scotland. Res. vet. sci. 29: 36–40.

toxoplasmosis in Canada. J. hyg. Camb. 77: 11–21.

environmental conditions. Vet. parasitol. 152: 186-193.

*gondii* in Ontario Mammals. Can. j. comp. med. 42: 177-183.

and Costa Rica. Am. j. trop. med. hyg. 24: 439–443.

prevalence and spatio-temporal variations.

samples? FEMS microbiol. rev. 27: 651-661.

microbiol. 98: 308-317.

10.1128/AEM.00246-12.

borne zoonotic dis. 10: 465–469.

cohort study of children, cats, rodents, birds, and soil. Am. j. trop. med. hyg. 53: 458-

[72] Lehmann T, Graham DH, Dahl E, Sreekumar C, Launer F, Corn JL, Gamble HR, Dubey JP (2003) Transmission dynamics of *Toxoplasma gondii* on a pig farm. Infect. genet. evol.

[73] Gauss CB, Dubey JP, Vidal D, Cabezon O, Ruiz-Fons F, Vicente J, Marco I, Lavin S, Gortazar C, Almería S (2006) Prevalence of *Toxoplasma gondii* antibodies in red deer (*Cervus elaphus*) and other wild ruminants from Spain. Vet. parasitol. 136: 193-200. [74] Malmsten J, Jakubek EB, Bjorkman C (2011) Prevalence of antibodies against *Toxoplasma gondii* and *Neospora caninum* in moose (*Alces alces*) and roe deer (*Capreolus capreolus*) in

[75] Williamson JMW, Williams H (1980) Toxoplasmosis in farmed red deer (*Cervus elaphus*)

[76] Simon A, Chambellant M, Ward BJ, Simard M, Proulx JF, Levesque B, Bigras-Poulin M, Rousseau AN, Ogden NH (2011) Spatio-temporal variations and age effect on *Toxoplasma gondii* seroprevalence in seals from the Canadian Arctic. Parasitology 138: 1-

[77] Tizard IR, Fish NA, Quinn JP (1976) Some observations on the epidemiology of

[78] Gotteland C, Aubert A, Gilbert P, Moinet M, Klein F, Villena I, Game Y, Gilot-Fromont E (submitted) *Toxoplasma gondii* in natural populations of Ungulates in France :

[79] Dumètre A, Dardé ML (2003) How to detect *Toxoplasma gondii* oocysts in environmental

[80] Davies, CM, Altavilla N, Krogh M, Ferguson CM, Deere DA, Ashbolt NJ (2005) Environmental inactivation of *Cryptosporidium* oocysts in catchment soils. J. appl.

[81] Langkjær M, Roepstorff A (2008) Survival of *Isospora suis* oocysts under controlled

[82] Frenkel JK, Ruiz A, Chinchilla M (1975) Soil survival of *Toxoplasma* oocysts in Kansas

[83] Lélu M, Villena I, Dardé ML, Aubert D, Geers R, Dupuis E, Marnef F, Poulle ML, Gotteland C, Dumètre A, Gilot-Fromont E (2012) Quantitative assessment of the survival of *Toxoplasma gondii* oocysts in soil. Appl. env. microbiol. doi:

[84] Stenseth NC,Viljugrein H, Jedrejewski W, Mysterud A, Pucek Z (2002) Population dynamics of *Clethrionomys glareolus* and *Apodemus flavicollis* : seasonal components of density dependence and density independence. Acta theriol. 47 (Suppl. 1): 39–67. [85] Tizard R, Harmeson J, Lai CH (1978) The Prevalence of Serum Antibodies to *Toxoplasma* 

[86] Bobić B, Klun I, Nikolić A, Nikolić A, Vujanić M, Zivković T, Ivović V, Djurković-Djakovic O (2010) Seasonal variations in human *Toxoplasma* infection in Serbia. Vector


[100] Sobrino R, Cabezón O, Millán J, Pabón M, Arnal MC, Luco DF, Gortázar C, Dubey JP, Almería S (2007) Seroprevalence of *Toxoplasma gondii* antibodies in wild carnivores from Spain. Vet. parasitol. 148: 187-192.

The Life Cycle of *Toxoplasma gondii* in the Natural Environment 31

[116] Sepúlveda MA, Muñoz-Zanzi C, Rosenfeld C, Jara R, Pelican KM, Hill D (2011) *Toxoplasma gondii* in feral American minks at the Maullín river, Chile. Vet. parasitol. 175:

[117] Stieve E, Beckmen K, Kania SA, Widner A, Patton S (2010) *Neospora caninum* and *Toxoplasma gondii* antibody prevalence in Alaska wildlife. J. wildl. dis. 46: 348-355. [118] Vikoren T, Tharaldsen J, Fredriksen B, Handeland K (2004) Prevalence of *Toxoplasma gondii* antibodies in wild red deer, roe deer, moose, and reindeer from Norway. Vet.

[119] Zarnke RL, Dubey JP, Kwok OC, Ver Hoef JM (2000) Serologic survey for *Toxoplasma*

[120] Miller MA, Miller WA, Conrad PA, James ER, Melli AC, Leutenegger CM, Dabritz HA, Packham AE, Paradies D, Harris M, Ames J, Jessup DA, Worcester K, Grigg ME (2008) Type X *Toxoplasma gondii* in a wild mussel and terrestrial carnivores from coastal California: New linkages between terrestrial mammals, runoff and toxoplasmosis of sea

[121] Massie GN, Ware MW, Villegas EN, Black MW (2010) Uptake and transmission of *Toxoplasma gondii* oocysts by migratory, filter-feeding fish. Vet. parasitol. 169: 296-303. [122] Prestrud KW, Asbakk K, Fuglei E, Mørk T, Stien A, Ropstad E, Tryland M, Gabrielsen GW, Lydersen C, Kovacs KM, Loonen MJ, Sagerup K, Oksanen A (2007) Serosurvey for *Toxoplasma gondii* in arctic foxes and possible sources of infection in the high Arctic of

[123] Prestrud KW, Asbakk K, Mørk T, Fuglei E, Tryland M, Su C (2008) Direct highresolution genotyping of *Toxoplasma gondii* in arctic foxes (*Vulpes lagopus*) in the remote arctic Svalbard archipelago reveals widespread clonal Type II lineage. Vet parasitol.

[124] Maubon D, Ajzenberg D, Brenier-Pinchart MP, Dardé ML, Pelloux H (2008) What are the respective host and parasite contributions to toxoplasmosis? Trends parasitol. 24:

[125] Innes EA (1997) Toxoplasmosis: comparative species susceptibility and host immune

[126] Shimizu K (1958) Studies on toxoplasmosis I: An outbreak of toxoplasmosis among

[127] Gustafsson K, Järplid B (1997) *Toxoplasma gondii* infection in the mountain hare (*Lepus timidus*) and domestic rabbit (*Oryctolagus cuniculus*). I. Pathology. J. comp. pathol. 117:

[128] Gustafsson K, Wattrang E, Fossum C, Heegaard PMH, Lind P, Uggla A (1997) *Toxoplasma gondii* infection in the mountain hare (*Lepus timidus*) and domestic rabbit

(*Oryctolagus cuniculus*). II. Early immune reactions. J. comp. pathol. 117: 361–369. [129] Sedlak K, Literak I, Faldyna M, Toman M, Benak J (2000) Fatal toxoplasmosis in brown hares (*Lepus europaeus*): Possible reasons of their high susceptibility to the infection. Vet.

response. Comp. immunol. microbiol. infect. dis. 20: 131-138.

hares (*Lepus timidus ainu*) in Sapporo. Jap. j. vet. res. 6: 157–166.

*gondii* in selected wildlife species from Alaska. J. wildl. dis. 36: 219-224.

60–65.

parasitol. 120: 159-169.

otters. Int. j. parasitol. 38: 1319–1328.

Svalbard. Vet parasitol. 150: 6-12.

158: 121-128.

299–303.

351–360.

parasitol. 93: 13–28.


[116] Sepúlveda MA, Muñoz-Zanzi C, Rosenfeld C, Jara R, Pelican KM, Hill D (2011) *Toxoplasma gondii* in feral American minks at the Maullín river, Chile. Vet. parasitol. 175: 60–65.

30 Toxoplasmosis – Recent Advances

1730.

Spain. Vet. parasitol. 148: 187-192.

eukariot. microbiol. 48: S197-S198.

from Spain. Vet. parasitol. 123: 265–270.

Spain. Vet. parasitol. 153: 152–156.

Vet. parasitol. 171: 331-336.

vet. sci. 75: 21-25.

23: 240-243.

163.

[100] Sobrino R, Cabezón O, Millán J, Pabón M, Arnal MC, Luco DF, Gortázar C, Dubey JP, Almería S (2007) Seroprevalence of *Toxoplasma gondii* antibodies in wild carnivores from

[101] García-Bocanegra I, Dubey JP, Martínez F, Vargas A, Cabezón O, Zorrilla I, Arenas A, Almería S (2010). Factors affecting seroprevalence of *Toxoplasma gondii* in the

[102] Degen AA, Kam M, Khokhlova IS, Krasnov BR, Barraclough TG (1998) Average daily metabolic rate of rodents: habitat and dietary comparisons. Funct. ecol. 12: 63–73. [103] Speakman JR (2005) Body size, energy metabolism and lifespan. J. exp. biol. 208: 1717–

[104] Ottaviani D, Cairns SC, Oliverio M, Boitani L (2006) Body mass as a predictive variable

[105] Ruiz A, Frenkel JK (1980) Intermediate and transport hosts of *Toxoplasma gondii* in

[106] Lindsay DS, Phelps KK, Smith SA, Flick G, Sumner SS, Dubey JP (2001) Removal of *Toxoplasma gondii* oocysts from seawater by Eastern oysters (*Crassostrea virginica*). J

[107] Putignani L, Mancinelli L, Del Chierico F, Menichella D, Adlerstein D, Angelici MC, Marangi N, Berrilli F, Caffara M, di Regalbono A (2011) Investigation of *Toxoplasma gondii* presence in farmed shellfish by nested PCR and real-time PCR fluorescent

[108] Dubey JP (2004) Toxoplasmosis - a waterborne zoonosis. Vet. parasitol. 126: 57-72. [109] Bollo E, Pregel P, Gennero S, Pizzoni E, Rosati S, Nebbia P, Biolatti B (2003) Health status of a population of nutria (*Myocastor coypus*) living in a protected area in Italy. Res.

[110] Nardoni S, Angelici MC, Mugnaini L, Mancianti F (2011). Prevalence of *Toxoplasma gondii* infection in *Myocastor coypus* in a protected Italian wetland. Parasite and Vectors

[111] Almeria S, Calvete C, Pages A, Gauss CBL, Dubey, JP (2004). Factors affecting the seroprevalence of *Toxoplasma gondii* infection in wild rabbits (*Oryctolagus cuniculus*)

[112] Gamarra JA, Cabezón O, Pabón M, Arnal MC, Luco DF, Dubey JP, Gortázar C, Almería S (2008) Prevalence of antibodies against *Toxoplasma gondii* in roe deer from

[113] Germain E, Benhamou S, Poulle ML (2008) Spatio-temporal sharing between the

[114] Jokelainen P, Isomursu M, Näreaho A, Oksanen A (2011) Natural *Toxoplasma gondii* infections in European brown hares and mountain hares in Finland: proportional mortality rate, antibody prevalence, and genetic characterization. J. wildl. dis. 47: 154–

[115] Jokelainen P, Näreaho A, Knaapi S, Oksanen A, Rikula U, Sukura A (2010) *Toxoplasma gondii* in wild cervids and sheep in Finland: north-south gradient in seroprevalence.

European wildcat, the domestic cat and their hybrids. J. zool. 276: 195–203.

of homerange size among Italian mammals and birds. J. zool. 269: 317–330.

endangered Iberian lynx (*Lynx pardinus*). Vet. parasitol. 167: 36-42.

amplicon generation assay (FLAG). Exp. parasitol 127: 409-417.

Costa Rica. Am. J. trop. med. hyg. 29: 1161–1166.


[130] Kenny DE, Lappin MR, Knightly F, Baler J, Brewer M, Getzy DM (2002) Toxoplasmosis in Pallas' cats (*Otocolobus felis manul*) at the Denver Zoological Gardens. J. zoo wildl. med. 33: 131-138.

The Life Cycle of *Toxoplasma gondii* in the Natural Environment 33

[143] Aspinall TV, Marlee D, Hyde JE, Sims PF (2002) Prevalence of *Toxoplasma gondii* in commercial meat products as monitored by polymerase chain reaction - food for

[144] Dumètre A, Ajzenberg D, Rozette L, Mercier A, Dardé ML (2006) *Toxoplasma gondii* infection in sheep from Haute-Vienne, France: seroprevalence and isolate genotyping

[145] Halos L, Thébault A, Aubert D, Thomas M, Perret C, Geers R, Alliot A, Escotte-Binet S, Ajzenberg D, Dardé ML, Durand B, Boireau P, Villena I (2010) An innovative survey underlining the significant level of contamination by *Toxoplasma gondii* of ovine meat

[146] Owen MR, Trees AJ (1999) Genotyping of *Toxoplasma gondii* associated with abortion in

[147] Jungersen G, Jensen L, Rask MR, Lind P (2002) Non-lethal infection parameters in mice separate sheep Type II *Toxoplasma gondii* isolates by virulence. Comp. immunol.

[148] Dubey JP, Velmurugan GV, Rajendran C, Yabsley MJ, Thomas NJ, Beckmen KB, Sinnett D, Ruid D, Hart J, Fair PA, McFee WE, Shearn-Bochsler V, Kwok OC, Ferreira LR, Choudhary S, Faria EB, Zhou H, Felix TA, Su C (2011) Genetic characterisation of *Toxoplasma gondii* in wildlife from North America revealed widespread and high

[149] Wendte JM, Gibson AK, Grigg ME (2011) Population genetics of *Toxoplasma gondii*: new perspectives from parasite genotypes in wildlife. Vet. parasitol. 182: 96-111. [150] Smith DD, Frenkel JK (1995) Prevalence of antibodies to *Toxoplasma gondii* in wild mammals of Missouri and east central Kansas: biologic and ecologic considerations of

[151] de Thoisy B, Demar M, Aznar C, Carme B (2003) Ecologic correlates of *Toxoplasma gondii* exposure in free-ranging neotropical mammals. J. wildl. dis. 39: 456-459. [152] Santos PS, Albuquerque GR, da Silva VM, Martin AR, Marvulo MF, Souza SL, Ragozo AM, Nascimento CC, Gennari SM, Dubey JP, Silva JC. (2011) Seroprevalence of *Toxoplasma gondii* in free-living Amazon River dolphins (*Inia geoffrensis*) from central

[153] Sobral CA, Amendoeira MR, Teva A, Patel BN, Klein CH (2005) Seroprevalence of infection with *Toxoplasma gondii* in indigenous Brazilian populations. Am. j. trop. med.

[154] Berengo A, Pampiglione S, De Lalla F (1974) Serological studies on toxoplasmosis in

[155] Johnson WE, Eizirik E, Pecon-Slattery J, Murphy WJ, Antunes A, Teeling E, O'Brien SJ (2006) The late Miocene radiation of modern Felidae: a genetic assessment. Science 311:

[156] IUCN SSC Group CS (1996) Cat Website. Available: http://lynx.uio.no/lynx/catsgportal/cat-website/20\_cat-website/home/index\_en.htm.

some groups of Babiga Pygmies in Central Africa. Riv. parassitol. 35: 81-86.

prevalence of the fourth clonal type. Int. j. parasitol. 41: 1139-1147.

thought? Int. j. parasitol. 32: 1193-1199.

sheep. J parasitol. 85: 382-384.

microbiol. infect. dis. 25: 187-195.

transmission. J. wildl. dis. 31: 15-21.

Amazon, Brazil. Vet. parasitol. 29: 171-173.

hyg. 72: 37-41.

73-77.

02.04.2012.

by microsatellite analysis. Vet. parasitol. 142: 376-379.

consumed in France. Int. j. parasitol. 40: 193-200.


[143] Aspinall TV, Marlee D, Hyde JE, Sims PF (2002) Prevalence of *Toxoplasma gondii* in commercial meat products as monitored by polymerase chain reaction - food for thought? Int. j. parasitol. 32: 1193-1199.

32 Toxoplasmosis – Recent Advances

med. 33: 131-138.

293-299. Erratum in: Parasitology 130: 586.

toxoplasmosis. J wildl dis. 41: 691-700.

groups. Int j. parasitol. 32:27-38.

assay. J. clin. microbiol. 48: 4641-4645.

346-349.

parasitol. 36: 841-848.

(*Sus scrofa*) in France. Vet. parasitol. 164: 296–300.

*gondii* in raptors from France. J. wildl. dis. 44: 172-173.

[130] Kenny DE, Lappin MR, Knightly F, Baler J, Brewer M, Getzy DM (2002) Toxoplasmosis in Pallas' cats (*Otocolobus felis manul*) at the Denver Zoological Gardens. J. zoo wildl.

[131] Basso W, Edelhofer R, Zenker W, Möstl K, Kübber-Heiss A, Prosl H (2005) Toxoplasmosis in Pallas' cats (*Otocolobus manul*) raised in captivity. Parasitology 130:

[132] Brown M, Lappin MR, Brown JL, Munkhtsog B, Swanson WF (2005) Exploring the ecologic basis for extreme susceptibility of Pallas' cats (*Otocolobus manul*) to fatal

[133] Convention on international trade in threatened species (2001) Checklist of CITES species Convention on International Trade in Threatened Species, Geneva, Switzerland,

[134] Ajzenberg D, Bañuls AL, Tibayrenc M, Dardé ML (2002) Microsatellite analysis of *Toxoplasma gondii* shows considerable polymorphism structured into two main clonal

[135] Su C, Khan A, Zhou P, Majumdar D, Ajzenberg D, Dardé ML, Xing-Quan Zhu XQ, Ajioka JW, Rosenthal BM, Dubey JP, Sibley LD (2012) Globally diverse *Toxoplasma gondii* isolates comprise six major clades originating from a small number of distinct

[136] Ajzenberg D, Collinet F, Mercier A, Vignoles P, Dardé ML (2010) Genotyping of *Toxoplasma gondii* isolates with 15 microsatellite markers in a single multiplex PCR

[137] De Craeye S, Speybroeck N, Ajzenberg D, Dardé ML, Collinet F, Tavernier P, Van Gucht S, Dorny P, Dierick K (2010) *Toxoplasma gondii* and *Neospora caninum* in wildlife:

[138] Richomme C, Aubert D, Gilot-Fromont E, Ajzenberg D, Mercier A, Ducrot C, Ferté H, Delorme D, Villena I (2009) Genetic characterization of *Toxoplasma gondii* from wild boar

[139] Aubert D, Ajzenberg D, Richomme C, Gilot-Fromont E, Terrier ME, de Gevigney C, Game Y, Maillard D, Gibert P, Dardé ML, Villena I (2010) Molecular and biological characteristics of *Toxoplasma gondii* isolates from wildlife in France. Vet. parasitol. 171:

[140] Aubert D, Terrier ME, Dumètre A, Barrat J, Villena I (2008) Prevalence of *Toxoplasma*

[141] Jokelainen P, Isomursu M, Nareaho A, Oksanen A (2010) *Toxoplasma gondii* killing European brown hares and mountain hares in Finland: Proportional mortality rate, seroprevalence, and genetic characterization. 9th Biennial Conference of the European

[142] Su C, Zhang X, Dubey JP (2006) Genotyping of *Toxoplasma gondii* by multilocus PCR-RFLP markers: a high resolution and simple method for identification of parasites. Int. j.

Wildlife, Association, Vlieland, The Netherlands, 13 to 16 September 2010.

common parasites in Belgian foxes and Cervidae? Vet. parasitol. 178: 64-69.

http://www.felineconservation.org/feline\_species/pallas\_\_cat.htm.

ancestral lineages. Proc. nat. acad. sci. doi: 10.1073/pnas.1203190109.


[157] Elmore SA, Jones JL, Conrad PA, Patton S, Lindsay DS, Dubey JP (2010) *Toxoplasma gondii*: epidemiology, feline clinical aspects, and prevention. Trends parasitol. 26: 190- 196.

The Life Cycle of *Toxoplasma gondii* in the Natural Environment 35

Kauffmann-Lacroix C, Pujol S, Rabodonirina M, Bougnoux ME, Cuisenier B, Duhamel C, Duong TH, Filisetti D, Flori P, Gay-Andrieu F, Pratlong F, Nevez G, Totet A, Carme B, Bonnabau H, Dardé ML, Villena I (2009) Genotype of 88 *Toxoplasma gondii* isolates associated with toxoplasmosis in immunocompromised patients and correlation with

[171] Lehmann T, Marcet PL, Graham DH, Dahl ER, Dubey JP (2006) Globalization and the population structure of *Toxoplasma gondii*. Proc. natl. acad. sci. U S A 103: 11423-11428. [172] Dubey JP, Quirk T, Pitt JA, Sundar N, Velmurugan GV, Kwok OC, Leclair D, Hill R, Su C (2008) Isolation and genetic characterization of *Toxoplasma gondii* from raccoons (*Procyon lotor*), cats (*Felis domesticus*), striped skunk (*Mephitis mephitis*), black bear (*Ursus*

[173] Kapperud G, Jenum PA, Stray-Pedersen B, Melby KK, Eskild A, Eng J (1996) Risk factors for *Toxoplasma gondii* infection in pregnancy. Results of a prospective case-

[174] Bowie WR, King AS, Werker DH, IsaacRenton JL, Bell A, Eng SB, Marion SA (1997) Outbreak of toxoplasmosis associated with municipal drinking water. Lancet 350: 173-

[175] Aubert D, Villena I (2009) Detection of *Toxoplasma gondii* oocysts in water: proposition of a strategy and evaluation in Champagne-Ardenne Region, France. Memorias Do

[176] Baril L, Ancelle T, Goulet V, Thulliez P, Tirard-Fleury V, Carme B (1999) Risk factors for *Toxoplasma* infection in pregnancy: A case-control study in France. Scand. j. infect.

[177] AFSSA (2005) Toxoplasmose : état des connaissances et évaluation du risque lié à l'alimentation Rapport du groupe de travail « *Toxoplasma gondii* » de l'Afssa. http://www.anses.fr/cgi-bin/countdocs.cgi?Documents/MIC-Ra-Toxoplasmose.pdf. [178] Opsteegh M, Prickaerts S, Frankena K, Evers EG (2011) A quantitative microbial risk assessment for meatborne *Toxoplasma gondii* infection in The Netherlands. Int. j. food

[179] Kijlstra A, Meerburg BG, Mul MF (2004) Animal-friendly production systems may cause re-emergence of *Toxoplasma gondii*. Njas-Wageningen Journal of Life Sciences 52:

[180] van der Giessen J, Fonville M, Bouwknegt M, Langelaar M, Vollema A (2007) Seroprevalence of *Trichinella spiralis* and *Toxoplasma gondii* in pigs from different

[181] Meerburg BG, Van Riel JW, Cornelissen JB, Kijlstra A, Mul MF (2006) Cats and goat whey associated with *Toxoplasma gondii* infection in pigs. Vector-Borne and Zoonotic

[182] Kijlstra A, Meerburg B, Cornelissen J, De Craeye S, Vereijken P, Jongert E (2008) The role of rodents and shrews in the transmission of *Toxoplasma gondii* to pigs. Vet.

housing systems in The Netherlands. Vet. parasitol. 148: 371-374.

clinical findings. J. infect. dis. 199: 1155-1167.

Instituto Oswaldo Cruz 104: 290-295.

177.

dis. 31: 305-309.

119-132.

microbiol. 150: 103-114.

Diseases 6: 266-274.

parasitol. 156: 183-190.

*americanus*), and cougar (*Puma concolor*). J. parasitol. 94: 42-45.

control study in Norway. Am. j. epidemiol. 144: 405-412.


Kauffmann-Lacroix C, Pujol S, Rabodonirina M, Bougnoux ME, Cuisenier B, Duhamel C, Duong TH, Filisetti D, Flori P, Gay-Andrieu F, Pratlong F, Nevez G, Totet A, Carme B, Bonnabau H, Dardé ML, Villena I (2009) Genotype of 88 *Toxoplasma gondii* isolates associated with toxoplasmosis in immunocompromised patients and correlation with clinical findings. J. infect. dis. 199: 1155-1167.

[171] Lehmann T, Marcet PL, Graham DH, Dahl ER, Dubey JP (2006) Globalization and the population structure of *Toxoplasma gondii*. Proc. natl. acad. sci. U S A 103: 11423-11428.

34 Toxoplasmosis – Recent Advances

parasitol. 172: 144-146.

196.

658.

1196.

microbiol. 36: 324.

[157] Elmore SA, Jones JL, Conrad PA, Patton S, Lindsay DS, Dubey JP (2010) *Toxoplasma gondii*: epidemiology, feline clinical aspects, and prevention. Trends parasitol. 26: 190-

[158] Ullmann LS, da Silva RC, de Moraes W, Cubas ZS, dos Santos LC, Hoffmann JL, Moreira N, Guimaraes AM, Montaño P, Langoni H, Biondo AW (2010) Serological survey of *Toxoplasma gondii* in captive Neotropical felids from Southern Brazil. Vet.

[159] Bevins SN, Carver S, Boydston EE, Lyren LM, Alldredge M, Logan KA, Riley SP, Fisher RN, Vickers TW, Boyce W, Salman M, Lappin MR, Crooks KR, VandeWoude S (2012) Three pathogens in sympatric populations of pumas, bobcats, and domestic cats:

[160] Boothroyd JC (2009) Expansion of host range as a driving force in the evolution of

[161] Carme B, Demar-Pierre M (2006) [Toxoplasmosis in French Guiana. Atypical neotropical features of a cosmopolitan parasitosis]. Med. trop. (Mars) 66: 495-503. [162] Carme B, Demar M, Ajzenberg D, Dardé ML (2009) Severe acquired toxoplasmosis caused by wild cycle of *Toxoplasma gondii*, French Guiana. Emerg. infect. dis. 15: 656-

[163] Dardé ML, Villena I, Pinon JM, Beguinot I (1998) Severe toxoplasmosis caused by a *Toxoplasma gondii* strain with a new isoenzyme type acquired in French Guyana. J. clin.

[164] Ajzenberg D, Banuls AL, Su C, Dumetre A, Demar M, Carme B, Dardé ML (2004) Genetic diversity, clonality and sexuality in *Toxoplasma gondii*. Int. j. parasitol. 34: 1185-

[165] Mercier A, Ajzenberg D, Devillard S, Demar MP, de Thoisy B, Bonnabau H, Collinet F, Boukhari R, Blanchet D, Simon S, Carme B, Dardé ML (2011) Human impact on genetic diversity of *Toxoplasma gondii*: Example of the anthropized environment from French

[166] Elbez-Rubinstein A, Ajzenberg D, Dardé ML, Cohen R, Dumètre A, Yera H, Gondon E, Janaud JC, Thulliez P (2009) Congenital toxoplasmosis and reinfection during pregnancy: case report, strain characterization, experimental model of reinfection, and

[167] Grigg ME, Bonnefoy S, Hehl AB, Suzuki Y, Boothroyd JC (2001) Success and virulence in *Toxoplasma* as the result of sexual recombination between two distinct ancestries.

[168] Su C, Evans D, Cole RH, Kissinger JC, Ajioka JW, Sibley LD (2003) Recent expansion of

[169] Mercier A, Devillard S, Ngoubangoye B, Bonnabau H, Bañuls AL, Durand P, Salle B, Ajzenberg D, Dardé ML (2010) Additional Haplogroups of *Toxoplasma gondii* out of Africa: Population structure and mouse-virulence of strains from Gabon. PLoS negl.

[170] Ajzenberg D, Yera H, Marty P, Paris L, Dalle F, Menotti J, Aubert D, Franck J, Bessieres MH, Quinio D, Pelloux H, Delhaes L, Desbois N, Thulliez P, Robert-Gangneux F,

*Toxoplasma* through enhanced oral transmission. Science 299: 414-416.

implications for infectious disease transmission. PLoS ONE 7: e31403.

*Toxoplasma*. Mem. inst. Oswaldo Cruz 104: 179-184.

Guiana. Infect. genet. evol. 11: 1378-1387.

review. J. infect. dis. 199: 280-285.

Science 294: 161-165.

trop. dis 4: e876.


[183] Dubey JP, Hill DE, Rozeboom DW, Rajendran C, Choudhary S, Ferreira LR, Kwok OC, Su C (2012) High prevalence and genotypes of *Toxoplasma gondii* isolated from organic pigs in northern USA. Vet parasitol. in press.

**Chapter 2** 

© 2012 Bobić et al., licensee InTech. This is an open access chapter distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

© 2012 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution,

*Toxoplasma gondii* **Infection in South-East** 

Branko Bobić, Ivana Klun, Aleksandra Nikolić and Olgica Djurković-Djaković

Toxoplasmosis is a globally distributed parasitic zoonosis. Cats and other *Felidae* are the definitive host of the *Toxoplasma gondii* parasite, in whose intestines the sexual cycle takes place, and they are the primary source of infection to all animals including humans, by way of contaminating the environment with oocysts excreted in the feces (Dubey, 2010). Herbivores are infected by ingestion of food and water contaminated by oocysts, and carnivores by eating tissue cysts present in the flesh of infected animals. Omnivores including humans are infected by both routes – by oocysts via improperly washed vegetables or fruits, contaminated water or hands, and by tissue cysts via improperly processed or raw meat. Vertical transmission, from mother to offspring, may occur, and is the cause of congenital toxoplasmosis (CT). CT is actually the major *T. gondii*-induced clinical entity, which, along with opportunistic infection in immunocompromised patients, defines the clinical significance of toxoplasmosis. The gravity of the potential consequences of CT on the one hand, and the preventability of the disease on the other, call for implementation of prevention programs. A prerequisite for an adequate choice of

prevention strategy is continuous monitoring of the local epidemiological situation.

This chapter reviews the epidemiology and epizootiology of *T. gondii* infection in South-East Europe (SEE). SEE is here considered as the territory comprising the Balkan Peninsula (35° – 46°53' N latitude, 13°23' – 30° E longitude), bordered by the Adriatic Sea to the west, the Mediterranean Sea to the south, the Black Sea to the east and the rivers Sava and Danube to the north, encompassing the countries descending from ex-Yugoslavia including Slovenia, Croatia, Bosnia & Herzegovina, Serbia, Montenegro and FYR of Macedonia (FYRoM), as well as Albania, Bulgaria and mainland Greece. Most of the area is mountainous, while the climate varies from Mediterranean to moderate. The whole region has a combined area of

and reproduction in any medium, provided the original work is properly cited.

**Europe: Epidemiology and Epizootiology** 

Additional information is available at the end of the chapter

550,000 km2 and a population of 55 million.

http://dx.doi.org/10.5772/50831

**1. Introduction** 

