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International Journal for Parasitology 38 (2008) 1257–1278 www.elsevier.com/locate/ijpara

Invited Review

Toxoplasma gondii infection in humans and animals in the United States
J.P. Dubey a,*, J.L. Jones b
United States Department of Agriculture, Agricultural Research Service, Animal and Natural Resources Institute, Animal Parasitic Diseases Laboratory, Building 1001, Beltsville, MD 20705-2350, USA b Division of Parasitic Diseases, National Center for Zoonotic, Vectorborne and Enteric Diseases, Coordinating Center for Infectious Diseases, Centers for Disease Control and Prevention, 4770 Buford Highway, MS: F22, Chamblee, GA 30341, USA Received 14 January 2008; received in revised form 11 March 2008; accepted 11 March 2008

Abstract This paper reviews clinical and asymptomatic Toxoplasma gondii infection in humans and other animals in the USA. Seroprevalence of T. gondii in humans and pigs is declining. Modes of transmission, epidemiology and environmental contamination with oocysts on land and sea are discussed. Published by Elsevier Ltd on behalf of Australian Society for Parasitology Inc.
Keywords: Toxoplasma gondii; Humans; Animals; Oocysts; Tissue cysts; USA

1. Introduction Toxoplasma gondii infections are prevalent in humans and animals worldwide (Dubey and Beattie, 1988). Felids are the key animal species in the life cycle of this parasite because they are the hosts that can excrete the environmentally-resistant stage, the oocyst. Humans become infected post-natally by ingesting tissue cysts from undercooked meat, consuming food or drink contaminated with oocysts, or by accidentally ingesting oocysts from the environment. However, only a small percentage of exposed adult humans or other animals develop clinical signs of disease. It is unknown whether the severity of toxoplasmosis in immunocompetent hosts is due to the parasite strain, host variability or other factors. Recently, attention has been focused on genetic variability among T. gondii isolates from apparently healthy and sick hosts. It has been 100 years since the discovery and naming of T. gondii. The parasite was first found in laboratory animals (for history see Dubey, 2007). Its medical importance remained unknown until 1939 when T. gondii was identified

conclusively in tissues of a congenitally-infected infant in New York City, USA (Wolf et al., 1939), and its veterinary importance became known when it was found to cause abortion storms in sheep in 1957 in Australia (Hartley and Marshall, 1957). In the present paper, we summarize information on clinical and sub-clinical T. gondii infections in humans and animals in the USA, including transmission, epidemiology and control. 2. Clinical and asymptomatic Toxoplasma gondii infection in humans and animals 2.1. Infection in humans 2.1.1. Asymptomatic infection Infection with T. gondii can occur pre- or post-natally. After birth, humans are usually infected with T. gondii by ingestion of oocysts in soil or water that have been contaminated with cat feces, or by ingestion of tissue cysts in undercooked meat (Dubey and Beattie, 1988; Bowie et al., 1997; Bahia-Oliveira et al., 2003; Dubey, 2004; Jones et al., 2005; de Moura et al., 2006). Transfusion or organ transplantation from an infected person can also transmit the organism (Shulman and Appleman, 1991; Schaffner,


Corresponding author. Tel.: +1 301 504 8128; fax: +1 301 504 9222. E-mail address: jitender.dubey@ars.usda.gov (J.P. Dubey).

0020-7519/$34.00 Published by Elsevier Ltd on behalf of Australian Society for Parasitology Inc. doi:10.1016/j.ijpara.2008.03.007


J.P. Dubey, J.L. Jones / International Journal for Parasitology 38 (2008) 1257–1278

2001). Most persons infected after birth are asymptomatic (Montoya and Liesenfeld, 2004; Remington et al., 2006), however, some develop a mild disease or in rare cases, a more severe systemic illness (see Section 2.1.2). Once infected, humans are believed to remain infected for life. Unless immunosuppression occurs and the organism reactivates, people usually remain asymptomatic. However, there is ongoing research on whether chronic T. gondii ˇ ek et al., infection has an effect on reaction time (Havlı ´c 2001), tendency for accidents (Flegr et al., 2002), behavior (Flegr et al., 1996, 2000; Lafferty, 2005, 2006) and mental illness (Yolken et al., 2001; Flegr et al., 2003; Brachmann et al., 2005; Brown et al., 2005). Selected serological surveys in humans in the USA are summarized in Table 1. Previous serological surveys were summarized by Dubey and Beattie (1988). A recent serosurvey using samples from the population-based National Health and Examination Nutrition Study (NHANES) found a decrease in the age-adjusted T. gondii prevalence in USA-born persons 12–49 years old from 14.1% in 1988–1994 to 9% in 1999–2004, a seroprevalence of 11% in USA-born women 15–44 years old in 1999–2004, and a seroprevalence of 28.1% in foreign-born women 1999– 2004 (Jones et al., 2007). The overall seroprevalence (USA and foreign-born combined) increased with age and was higher among non-Hispanic black persons and Mexican Americans than among non-Hispanic white persons; however, among USA-born persons Mexican Americans had a lower seroprevalence than non-Hispanic white or black persons (age 12–49, 5.1% versus 8.8% and 11.5%, respectively). An earlier study evaluating NHANES III (1988–1994) sera for all persons 612 years of age (USA and foreign-born combined) showed an overall ageadjusted seroprevalence of 22.5%, a relatively linear increase in T. gondii infection with age, and a higher ageadjusted seroprevalence in the northeastern USA (29.2%) compared with the South (22.8%), Midwest (20.5%) or West (17.5%) (Jones et al., 2001). In this study, the risk for T. gondii infection was higher among persons who were foreign-born, had a lower education level, lived in crowded conditions and worked in soil-related occupations. In a separate study using NHANES III sera, the rate of T. gondii seropositivity among persons seropositive for the soil-

transmitted helminth Toxocara spp. was nearly double the rate of T. gondii seropositivity among persons who were not seropositive for Toxocara spp., suggesting that sufficient soil exposure to lead to Toxocara spp. infection doubles the risk of T. gondii infection (Jones et al., 2008). Prior studies have also shown a decrease in T. gondii seroprevalence in the USA over time. For example, in 1962 and 1989 T. gondii seroprevalence was examined among military recruits, showing rates of 14.4% and 9.5%, respectively (Feldman, 1965; Smith et al., 1996). Although not a complete sampling of the USA regional populations, the studies in military recruits (Feldman, 1965; Smith et al., 1996) and an earlier study (Feldman and Miller, 1956) found lower rates of T. gondii infection in the West. The western region of the USA is generally drier and oocysts may not survive as well in the soil in this climate. However, due to variations in weather, cat populations and human behavior, there is likely to be a wide variation in T. gondii prevalence within regions of the USA. 2.1.2. Symptomatic infection A minority of healthy persons infected with T. gondii after birth develop symptoms, which are usually mild and include manifestations such as fever, malaise and lymphadenopathy (Montoya and Liesenfeld, 2004; Remington et al., 2006). However, in rare cases, humans who were previously healthy have developed severe and even fatal disease, including pulmonary and multivisceral involvement, possibly from more virulent types of the organism (Carme et al., 2002; Demar et al., 2007). In addition, up to 2% of healthy persons in the USA infected with T. gondii develop ocular disease (Holland, 2003), usually retinochoroiditis. A higher percentage of infected persons have been documented to develop ocular disease in other parts of the world, for example, one region of Southern Brazil (17.7% with ocular lesions) (Glasner et al., 1992). Retinochoroiditis can be due to congenital or post-natally acquired disease and can be associated with acute infection or reactivation (Montoya and Remington, 1996; Holland, 1999). Current thinking is that the majority of ocular toxoplasmosis comes from post-natally acquired disease (Holland, 1999, 2003). Acute toxoplasmic retinochoroiditis results in pain, photophobia, tearing and loss of vision.

Table 1 Selected USA human Toxoplasma gondii antibody prevalence studies Year sampled 1962 1987 1989 1992–1993 1988–1994 1999–2000 1999–2004

Age group U.S. young adult P18 years old U.S. young adult P18 years old U.S. age-adjusted P12 years old U.S. age-adjusted 12–49 years Women 12–49 years U.S. age-adjusted 12–49 years Women 15–44 years

Source of sera Military recruits Maryland community Military recruits Illinois swine farm workers NHANESa NHANES NHANES

No. tested 2680 251 2862 174 17,658 4234 15,960

% Positive 14 31 9.5 31 22.5 15.8 14.9 10.8 11.0

Reference Feldman (1965) Roghmann et al. (1999) Smith et al. (1996) Weigel et al. (1999) Jones et al. (2001) Jones et al. (2003) Jones et al. (2007)

NHANES, National Health and Nutrition Examination Study.

1984.. 2001). Wong et al. 1991. gondii infection and the estimate that up to 2% of persons with T. 1999.. The risk of congenital infection is lowest when maternal infection is in the first trimester (10–15%) and highest when infection occurs during the third trimester (60–90%) (Dunn et al. 1984. 2000.000 and seven per 10. especially when the lesions are near the central structures of the eye (Holland. if not treated. pp. 1999.2. 1993).. Jones et al..... 2.. Remington et al. Holland. Kaplan et al. 2003)..2. Congenital infection can lead to a wide variety of manifestations in the fetus and infant including spontaneous abortion. In another study. stupor and coma (Luft et al. confusion. a relatively large burden on the medical system (Lum et al. 1992.. In addition.P.9 cases . which serves Massachusetts and New Hampshire. 2007). 2004. McLeod et al.. particularly of the thalamus (Renold et al. In the USA. Speech abnormalities and hemiparesis are the most common focal neurological findings (Luft et al. Foulon et al. Only two (4%) of these infants were recognized to have congenital toxoplasmosis before the screening results were known. 1994. 2006). accessed 1/10/08 http://aidsinfo. although rare exceptions have been reported in which women were infected just before pregnancy (Vogel et al.. J. 1994). cerebral calcifications and retinochorioditis. 2001).. Wong et al. 1992). 1984). in immunosuppressed women reactivation of an infection acquired before pregnancy can lead to congenital toxoplasmosis (Mitchell et al. In the era before highly active antiretroviral therapy (prior to the mid-1990s). 1997).. A French study found that the rate of toxoplasmic encephalitis declined from 3. gondii seropositive HIV-infected persons with advanced immunosuppression who were not receiving prophylaxis with drugs active against T. Minkoff et al. 2006). 1984. However. Porter and Sande.. Dunn et al. 1974).. gondii (Benson et al.1. The clinical presentation often includes a focal encephalitis with headache. can progress to seizures. 1984. However.. Leport et al.. follow-up examinations identified signs of disease in 40% of the infants (Guerina et al. the annual incidence was 38% among T. The primary lesion is cerebral necrosis.pdf. Immunosuppression. Remington et al. All infants born in the catchment area are screened for a variety of conditions including T.1.000 births. 1986. 1990. Guerina et al. respectively (Kimball et al. 1994). A national survey of ophthalmologists resulted in estimates that there were over 250. resulting in an infection rate of approximately 1 per 10.gov/contentfiles/TreatmentofOI_AA.. still-birth.0 cases per 100 person-years among HIV-infected persons with a CD4+ T-lymphocyte count less than 100 cells per microlitre (Jones et al. 1996). 2005). Department of Health and Human Services. Systemic Review on Congenital Toxoplasmosis Study Group. 1999. Wong et al. 1971.26 million persons in the USA may have ocular toxoplasmosis (based on the 2000 census. motor weakness and fever and.1. an infant who fails to thrive or has CNS involvement or retinochoroiditis. 1983. Congenitally-infected children.. the annual incidence of toxoplasmic encephalitis was up to 33% among T. as many as 1. 1993) when persons become severely immunosuppressed. A subsequent study on the epidemiology of congenital toxoplasmosis identified in the New England Regional Newborn Screening Program 1988–1999 reported a rate of approximately 1 per 12.. 1999. 2003). 1974.L. congenital infection usually leads to more severe disease when it occurs in the first trimester (Desmonts and Couvreur. Toxoplasmic encephalitis is the most common clinical presentation of toxoplasmosis among persons with AIDS... a live infant with classic signs of congenital toxoplasmosis such as hydrocephalus or microcephalus. 2006). Schaffner. 1996). Of the 635. gondii seropositive persons with AIDS who were studied over 2 years (Israelski et al. or an apparently normal infant who develops retinochoroiditis or symptoms of CNS involvement later in life (McAuley et al. the incidence and deaths associated with toxoplasmic encephalitis declined markedly (Jones et al. 2006... but many will develop ocular or neurological manifestations (including learning disabilities) later in life (Wilson et al. Holliman. Jones / International Journal for Parasitology 38 (2008) 1257–1278 1259 Lesions tend to recur with progressive loss of vision over time.. gondii infection have ocular lesions in the USA (Holland.2. AidsInfo. 2003). 9– 12). 2007). however. 1996. other disseminated systemic disease or retinochoroiditis can be seen but are not as common as toxoplasmic encephalitis in HIVinfected persons.. with the highest risk occurring when the CD4+ T-lymphocyte count drops below 50 cells per microliter (Luft et al.nih.J. 1995... gondii infection. 1983.000 births. 1984. 1971. 1980. gondii during pregnancy (Remington et al. 52 were infected. 1996). Hooshyar et al. Treating opportunistic infections among HIV-infected adults and adolescents..000 live births (Jara et al.S. Most children are asymptomatic at birth (Guerina et al. Shulman and Appleman. 2006). An analysis of 90 inpatient and outpatient facilities in nine USA cities from January 1990 through August 1995 found an incidence rate of toxoplasmic encephalitis of 4. It is usually the result of reactivation of latent tissue cysts (Luft et al.... Congenital toxoplasmosis generally occurs when a woman is newly infected with T. 2.. Older prospective studies in Alabama and New York reported in the 1970s found rates of congenital toxoplasmosis of 13 per 10. 2002.000 visits to ophthalmologists for active or inactive ocular toxoplasmosis in a 2-year period. Remington et al. U.. 1994. Alford et al.000 infants who underwent serological testing between 1986 and 1992.. during the years that prophylaxis and highly active antiretroviral therapy became widely used (mid-1990s in most developed countries). Israelski et al. 1983. Pneumonia.. Koppe et al. Considering the prevalence of T. Dubey. Clinically severe toxoplasmosis can also occur in immunosuppressed persons with malignancies and after transfusions (rarely) or transplants with immunosuppressive therapy (Siegel et al. the most complete data on the rate of congenital infection comes from the New England Regional Newborn Screening Program... 1993).

with a 50% mortality rate. premature births and deaths soon after birth. were associated with T. tremors and diarrhea.. pigs and cattle are the main species of livestock in USA.9c 6 87.1c 20. The prevalence of T. gondii was isolated from 51 of 55 pigs when hearts and tongues were used for testing (Table 3). (1995) Weigel et al. 1996 and 1998 showed a steady decline (Table 2). There were still-births. The prevalence of T. USA (Farrell et al.3% to 92. 1995c). 2000)... J. tested 11. (2008b) Modified agglutination test. In poorly managed non-confinement systems.1 0.2.5 20 36 3.P. gondii infection in pigs (Weigel et al. Patton et al. These findings have not been confirmed and it seems likely that the outbreak was complicated by other causes (Dubey and Beattie. Risk assessment studies indicated that exposure to cats (oocysts) and mice.2 68. Sanger and Cole. Declining seroprevalence in market pigs. clinical toxoplasmosis in pigs is rare. Dubey. (1996) Assadi-Rad et al. As mentioned earlier. In one study of 1000 sows. (1999) e Dubey et al.229 623 1000 509 3479 3841 1885 5080 4252 2617 2238 4712 3236 1897 8086 5720 55 48 % Positive 23 42 22. market pigs) are mostly raised indoors in well managed facilities to prevent access to rodents and cats.. (1995a) Dubey et al. gondii varies dramatically among the classes of pigs surveyed (market pigs versus sows. seroprevalence was 23% in market pigs and 42% in breeder pigs (sows) (Dubey et al. 1:20 or higher dilution. (1979) described acute toxoplasmosis in a pig- Table 2 Prevalence of Toxoplasma gondii antibodiesa in sera of pigs from the USA Year sampled 1983–1984 1989–1992 1989–1990 1990 1991–1992 1992 1992–1993 1994–1995 1994–1995 1998 2000 2002 2006 a b c d e Type Market hogs Sows Sows All ages Sows Sows Market hogs Sows Market hogs Sows Market hogs Market hogs Sows Various ages Market hogs Sows Market hogs Market hogs Source of sera Nationwide Abattoir-Iowa 31 Farms-Hawaii NAHMSb Tennessee-343 herds Illinois-179 herds Illinois-47 herds North Carolina-14 herds NAHMS Connecticut. In these well managed facilities. Dubey et al.7% of pigs surveyed. depending on the source of pigs. 1988). 1991). 2001). class of pigs and the type of pork tested (Table 3). Jones / International Journal for Parasitology 38 (2008) 1257–1278 per 100 person-years in the time before the availability of highly active antiretroviral therapy (1992–1995) to 1 case per 100 person-years after it was available (1996–1998) (Abgrall et al. Massachusetts.1. The pigs used for unprocessed pork consumption (feeder pigs. prevalence had dropped to 20. gondii was isolated from 17% of these using heart tissue for bioassays. gondii were found in colostrums and sows’ milk.1260 J.7 c Reference Dubey et al. 2. (1995c) Davies et al. Livestock Poultry. Toxoplasmosis was first reported in pigs on a farm in Ohio..2. and not outdoor housing. Pigs.. Seropositivity in general is a good indicator of the presence of viable parasites. (2002a) Dubey et al. NewHampshire. Toxoplasma gondii infections in other animals 2. prevalence runs as high as 68% (Table 2). Prevalence of T. Patton et al. National Animal Health Monitoring System. lack of coordination. gondii in three NAHMS swine surveys in 1990. Pigs on this farm were reported to have cough. 22% were seropositive and viable T. When pigs from these same areas were tested in 1992.2 48. the isolation rate would likely have been higher if additional tissues were sampled. gondii was isolated from 0.1. viable T.5 0. Seroprevalence data are summarized in Table 2. 1995b).8% in breeders and 3. gondii has greatly declined in the last decade (Table 2). The institution of a National Animal Health Monitoring System (NAHMS) for swine now allows periodic surveillance of pigs for microbial infections.1% in finisher pigs (Dubey et al. Vermont NAHMS Massachusetts-1 herd Maryland-1 herd No. indoor pigs with a biosecurity system versus free-range).2c 15 47. 1952. (1999) d Gamble et al. (1991) Dubey et al. In a statistically valid population-based nationwide survey conducted in 1983– 1984. A decline to a similar incidence level was shown during the same time period across 11 USA cities (Kaplan et al. (1998). 1955). In another study.3c 15. Rode Island.L.8 2. . Clinical toxoplasmosis is not a major problem in these animals. gondii in pigs is also influenced by management systems. 2. (2000). Viable T. prevalence of T. (1995a) Dubey et al. (1992c) Patton et al.1.2. Viable T.5c 3.

and it is difficult to verify specificity using naturally-infected cattle. 1985b. 1985). (2008) Dubey et al. This piglet and 15 littermates were apparently normal at birth. gondii was isolated from an aborted fetus from a cow in Washington State. These authors reviewed other attempts to isolate T. The fourth steer became seronegative 15 months p. gondii oocysts after birth (Dubey et al.i.1. (1995b) Dubey et al. 2. This 500 kg cow was killed and 100– 500 g portions of its tissues were bioassayed in cats (500 g of each tissue) and mice (100 g of each tissue). 3.. lymphadenitis. (2004b) let on a farm in Indiana. Minnesotta Pennsylvania Pennsylvania No. (2004b) Dubey et al. and viable T. 2005b). gondii infection in cattle because several tests that are used to diagnose toxoplasmosis in humans give erratic results with cattle sera (Dubey et al.. 1993). nor were viable organisms detected (Dubey et al. (Dubey and Thulliez. This piglet had necrosis of intestine. gondii from beef are given in Table 3. Viable T. viable T. Little is known about the specificity and sensitivity of serological diagnosis of T. In a recent survey. but developed diarrhea within 1–2 weeks. gondii was not isolated from chicken breast meat samples obtained from . None of the 12 cats fed approximately 6 kg of beef shed oocysts. (2005b) Dubey et al.7 0.. (2008b) Jacobs et al. (1960) Remington (1968) Dubey et al. 539.2 4 77. four steers weighing 100–150 kg were each fed 10.1.L. Iowa Massachusetts-1 herd Retail meat. Viable T. gondii infection in cattle. many tissues from each animal were bioassayed (100 g each for bioassay in mice and 500 g each for bioassay in cats) for viable T. 2. 1990a). gondii. perhaps due to innate resistance. gondii were isolated from steers killed 350 to 1191 days p. pneumonia and encephalitis. 1979). gondii infection in cattle.. Results of these experiments highlight the difficulties in detecting T. the parasite is eliminated or reduced to undetectable levels within a few weeks (Dubey. Other attempts to isolate T. (2008c) Jacobs et al. 1983. Dubey and Thulliez. There are no confirmed reports of clinical toxoplasmosis in cattle (Dubey.2.i.9 0 0 0 0 21 28. 2002).P. Although cattle can be successfully infected with T. nationwide Maryland-1 herd Abattoir-Maryland Retail meat Abattoir-Maryland Abattoir-Maryland Abattoir-Ohio Retail meat. gondii in cattle from Montana.2.8 9. Poultry. nationwide Alabama Mississppi Iowa. In one experiment...3. gondii was not isolated from any tissue either in cats or mice. 1992). nationwide Retail meat. (2005b) Lindsay et al. (1991) Dubey et al. (1995a) Dubey et al. 1985b). The dye test.2% of 2539 cattle had MAT titers of 1:128–1:512 (Dubey.J. (1960) Dubey et al. a titer of 1:100 or higher in the modified agglutination test (MAT) appears to be indicative of T.. gondii oocysts. gondii from a naturally-exposed beef cow (Dubey. gondi.. Viable T. by bioassays in cats but not by bioassays in mice. At necropsy. Dubey. Jones / International Journal for Parasitology 38 (2008) 1257–1278 Table 3 Isolation of Toxoplasma gondii from various food animals in the USA Species Pigs Source Abattoir-Maryland Abattoir. gondii and N. tachyzoites were demonstrable in lesions. gondii. Epidemiological data indicated that the piglets probably became infected with T. In another study. Among all serological tests evaluated. (2005b) Dubey et al. antibodies to T.3 36. 1993).7 17 35. Before the discovery of Neospora caninum as a cause of abortion in cattle (Thilsted and Dubey. USA. (1960) Dubey and Streitel (1976) Dubey et al. 1988c).000 oocysts and killed 350. (2002a) Dubey et al. In a study of beef from retail grocers. 1191 and 1201 days p. caninum are morphologically similar parasites (Dubey et al. gondii from bovine fetuses in the USA (Canada et al. which is the most specific test for humans. 1986). T. Although congenital toxoplasmosis can be induced in pigs (Dubey et al. gondii was not isolated from any of the edible tissues of the cow by bioassays in mice but was isolated from a homogenate of intestine of the cow. gives false or erratic results with cattle sera (Dubey et al. Whether the cow had aborted due to toxoplasmosis could not be determined because a histological examination of the fetus was not made. USA (Canada et al.2. Cattle. bioassayed and tissue 50 diaphragms 1000 sow hearts 55 market hogs Hearts and tongues 2094 pork 36 hearts 86 50 lamb chops 68 lamb hearts 60 diaphragms 350 mixed tissues 2094 beef 2094 breast meat 19 73 88 28 10 % Positive 24 17 92. gondii. an attempt was made to isolate T. Cattle are considered a poor host for T. 1989) it is likely that this parasite in cattle was misdiagnosed as T.i. J. 1986). we are not aware of a documented case of toxoplasmic abortion in pigs in the USA or elsewhere. and eight of those died within 3–4 weeks. 2002).7 70 Reference 1261 Sheep Cattle Chickens Deer Black bear Jacobs et al. gondii were not found by ELISA in meat juice from any of the 2049 samples of beef bioassayed during the National Retail Meat Survey for T. In one serological survey of T.

Dubey and Welcome. Ostendorf and Henderson [Ostendorf and Henderson. USA.2.. XII World Poultry Congress. Jacobs and Melton (1966) isolated viable T. Two recent surveys indicate a low prevalence of T. dermatitis and encephalitis. Unlike chickens raised in confinement. 36 pulmonary. Proc. 1988). 1989b). 1981a. There has been no nationwide survey of the prevalence of T. an inability to stand. Dubey and Kirkbride (1989b) isolated T. Abortion and neonatal mortality was reported in dairy goats in Montana. two hepatic. and lateral recumbancy were the only clinical signs..4% of chickens had antibodies to T. Lambs that survived the first week after birth remained asymptomatic and were bled at 3–4 months of age. Affected cats may appear depressed and anorexic and die suddenly with no obvious clinical signs. Dubey et al. two cutaneous and seven neurologic...628 cats (Vollaire et al. Dubey and Carpenter. Dubey and Kirkbride. Toxoplasma gondii was found histologically in 11 of 30 lambs that were born dead. Cats can suffer from clinical toxoplasmosis (Meier et al. antibodies (MAT 1:1024 or higher) were found in 67 of 112 lambs. 2. Australia. myositis. Marked lesions were limited to the brain which had multiple areas of necrosis. however herpes virus infection (Marek’s disease) could not be ruled out.3.. Illinois. gondii. The seroprevalence of T. Jones / International Journal for Parasitology 38 (2008) 1257–1278 retail meat stores (Table 3). Horses. gondii was higher in feral than pet or owned cats (Table 5). 2. Isolated occurrences and epidemics of abortion and neonatal mortality have been reported in sheep in the USA (Dubey et al. gondii in cats in the USA and the available data were summarized by Conrad et al. An unusual finding was the presence of numerous tissue cysts and tachyzoites in lesions. in press. and Maryland (Dubey. Clinical toxoplasmosis is most severe in congenitally infected kittens.1. one pancreatic. Toxoplasma gondii was isolated from the hearts of three. Pneumonia is the most important clinical manifestation of feline toxoplasmosis. backyard raised chickens are commonly infected with T.. J. tongues of seven. gondii in horses (Table 4). USA by Olafson and Monlux (1942) is now considered to have been due to a Sarcocystis-like parasite (Dubey and Beattie. gondii (Dubey et al. gondii varies with the age and lifestyle of the cat (Dubey. The prevalence of T.. Horses are resistant to T. Eight of these lambs with MAT titers of 1:4096 or higher were slaughtered when they were 7 months old. (2005).6.1262 J.4. Texas. Georgia and Louisiana (Gibson and Eyles. 2005b). however. pancreatic necrosis. Foster et al. We are not aware of any confirmed report of clinical toxoplasmosis in horses anywhere in the world. 2003a. gondii in chickens raised in large-scale confinement operations. Dubey. antibodies to T.P. Goodwin et al. leg of lamb in eight. The case of encephalomyelitis in an adult sheep in New York..L. 16 abdominal. 2.. Among 100 histologically confirmed cases of toxoplasmosis in cats.b. 2007b). 385–387] reported that chickens with toxoplasmic encephalitis and chorioretinitis died on chicken farms in Indiana. There is no other serological survey for T. 1988).5. 1986a) in the USA. myocarditis. 1988). 2006).1. 1981. gondii from eight of eight naturally-infected lambs from a flock in South Dakota. More recently. Sheep. gondii (Dubey et al. 1990b.2. 1989a. Torticollis. these chickens were primarily kept for egg production. Connecticut.. 1957. Viable T. The prevalence of T. These data are comparable among different categories of cats because results are based on 1:25 serum dilution tested by the MAT and most of those were performed in one laboratory. 36 were considered to have generalized toxoplasmosis. in a group of 14 backyard chickens in Illinois. Pets and other animals 2. In the largest survey using convenience samples from sick cats submitted for diagnosis to a clinical laboratory. Congenitally-infected lambs that survive the first week after birth usually grow normally and can be a source of infection for humans. 2. Ohio. 1990. three birds died suddenly (Dubey et al. gondii.2. gondii was isolated from 27% to 100% of chickens from backyard operations on small farms in Mississippi. 1957. Dubey and Lappin. 1973). 1981a. Most of the goats raised in the USA are dairy goats and females are used to produce milk and cheese. Section paper 385. Eyles et al. perivascular lymphocytic cuffs and gliosis. Domestic cats (Felis domesticus) Toxoplasma gondii infection in cats is both of epidemiological and clinical significance. gondii in adult sheep and lambs in the USA is high (Tables 3 and 4). Toxoplasma gondii causes abortion and neonatal mortality in sheep worldwide. Montana. adult goats can develop clinical toxoplasmosis involving liver. Although abortion and neonatal mortality are the main clinical signs. 1959. 1969. 1993a. pp. Goats. one cardiac.3. Dubey et al. the source of these chickens was not known. 2007b). The remaining 11 chickens remained asymptomatic and all contained viable T.1. in 14 cats . The lambs were from a flock that had aborted due to toxoplasmosis. The number of goats in the USA is small and they are often raised in small operations. and lamb chops of seven by bioassays of 100 g of each tissue in mice (Dubey and Kirkbride. uveitis. An example is given here. kidneys and brain (Dubey and Beattie. One of these birds was necropsied. Sydney. gondii from ovaries and oviducts of three and leg muscle of one of 108 chickens from a slaughter house in Maryland. 2007b). meat juice from 1. There is no documented case of clinical toxoplasmosis in adult sheep in the USA or elsewhere. Other common clinical manifestations are hepatitis. There are three reports of clinical toxoplasmosis in chickens from the USA. 1962. Dubey. (1994) reported peripheral neuritis in three chickens from Georgia and the diagnosis was made immunohistochemically. 1981.1. gondii were found in 31% of 12. Toxoplasmosis in chickens. 1986b. 2005).

4:55. concurrent microbial or other conditions were identified (Dubey and Carpenter. (2007a) a b Abattoir. Although FIV infection can experimentally predispose cats to generalized toxoplasmosis (Davidson et al.1 20 10 6.3 Serologic test ELISA IHAT IHAT MAT MAT MAT MAT IHAT MAT IHAT DT MAT MAT LAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT Cut-off value ? 1:64 1:64 1:16 1:16 1:64 1:25 1:64 1:40 1:64 1:2 1:20 1:25 1:32 1:25 1:25 1:32 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 Reference 1263 Malik et al. Minnesotta. . New England. South Dakota. New Jersey.4 59.5 64. this phenomenon appears to be rare in natu- rally-infected cats as there are only a few reports of fulminating toxoplasmosis in cats with FIV (Heidel et al. 1989). suffer from an immunodeficiency virus (FIV).2 17 18. (1992b) Goats Wild pigs White-tailed deer Black bear (Ursus americanus) Raccoon (Procyon lotor) 1989–1993 1992–1993 1984–1988 1989–1993 1993–1996 20 188 885 379 99 70 67 15 49 46 MAT MAT MAT MAT MAT 1:25 1:25 1:32 1:25 1:50 Brillhart et al. New York.8 27.2 0. 1990. (1996) Humphreys et al.2 80 78. Jones / International Journal for Parasitology 38 (2008) 1257–1278 Table 4 Serological prevalence of Toxoplasma gondii in selected species of animals in the USA Species Sheep Locality and source New Jersey. (1981) Dubey et al.5 80. New Jersey. like humans. (1998) Dubey et al.2 MAT MAT 1:25 1:25 Hancock et al. unpublished data Nutter et al. gondii and FIV is intriguing because both the seroprevalence and the magnitude of titers were higher in dually infected cats (Witt et al.0 24:8 20. (1998) Mitchell et al.. (2005) Dubey et al. (2004b) Dubey. (1983) Dubey et al. 2006).J. (2008c) Ruppanner et al. (1986b) Dubey and Welcome (1988) Dubey and Kirkbride (1989a) Dubey et al.8 65. (1977) Huffman et al. (1994) Dubey et al. (1993) Dubey et al.064 180 583 1367 73 84 665 28 80 37 143 427 % Positive 62.6 84 50.9 0. (1975c) Al-Khalidi and Dubey (1979) Dubey et al. Cats. 1993). Idaho. Oregon Idaho Maryland New York Iowa. (2004b) Dubey et al. (2008d) Briscoe et al. Dubey. Dubey and Lappin.. (1999) Dubey et al. (1990) Riemann et al. Ohio. Pennsylvania California. The interaction between T. Farms.. (1999) Lindsay et al.. South Carolina. (1978) Dubey and Adams (1990) Riemann et al. Virginia California Washington Horses Nationwide Midwest 24 states Wyoming California Georgia (mainland) (Ossawa island) South Carolina Pennsylvania Minnesotta Mississippi Iowa Pennsylvania Pennsylvania Pennsylvania Pennsylvania North Carolina Iowa. (1997a) Diderrich et al. (2004b) Dubey et al. Pennsylvania Kansas Illinois Iowa Illinois Florida. (1995c) Hill et al. (1997a) Dubey et al.9 31 60 30 46.P.5 75. (2001) 2000–2001 2006 256 54 84. Kansas Maryland. (1996) Dubey et al. J. Pennsylvania.L.P. (1995) Vanek et al. J. Massachusetts Virginia Wisconsin Year Not stated 1974–1976 Not stated 1984 1987 1983–1987 2006 1974–1977 1982–1984 1973 1976–1977 1998 2002 1982–1983 1979–1980 1992–1994 1990 1991 1990–1993 2002–2003 2007 1989–1992 1993 1998 2007 1996–1997 Not stated Number tested Adults-309a Lambs-345a Ewes-2164a Lambs-1056a Ewes-250b Sheep-78b Ewes-592b Ewes-1564b Lambs-383a Dairy goats-1054b Dairy goats-1000 1294 500a 1054a 276b 135 170 1. 1993a).8 41 73. (2003c) Clark et al.1 23 22.

squirrels (van Pelt and Dieterich. Miller et al. tested 4 16 74 391 20 94 80 78 % Positive 100 100b 41. 2004a). Common clinical manifestations of toxoplasmosis in dogs are pneumonia. diarrhea. T. Indiana.3. 2. are highly susceptible to toxoplasmosis (Ratcliffe and Worth. Viable T. 1988b. These animals can die suddenly. especially New World primates. myocarditis. Rhyan and Dubey.3. 1972. Enteritis characterized by necrosis of the cells of lamina propria and mesenteric lymphadenitis suggest oral infection with food and water contaminated with oocysts. antibodies to T. gondii in deer suggests widespread contamination of the environment with oocysts. Dubey and Beattie. Dubey et al. Wild animal species. hepatitis and blindness. 1966.. 1988. Jones / International Journal for Parasitology 38 (2008) 1257–1278 Table 5 Seroprevalence of Toxoplasma gondii in domestic cats (Felis domesticus) from the USA according to the type and habitat of the cat Cat type Goat farm Sheep farm Pig farms Pig farms Feral Rural Barn Outside Feral Shelter Owned North Carolina Feral Owned a b Locality Marland Maryland Iowa Illinois Iowa Ohio No. including from the USA (Boorman et al. 1960). Old World primates are generally resistant to clinical toxoplasmosis but there is a report of toxoplasmosis in macaques. Among marsupials. wallabies and kangaroos. Acute toxoplasmosis has been described in ring-tailed lemurs. Dubey et al. Other pets Sporadic cases of clinical toxoplasmosis occur in rabbits (Leland et al.. 1992a). 1985a. especially in canaries and finches (Dubey.. 1992. (1992) Dubey et al. Dubey.9% of red foxes (Vulpes vulpes) and gray foxes (Urocyon cinereoargenteus) in Kentucky. Bears and raccoons are omnivores and infections in those indicate cumulative contamination with oocysts and intermediate hosts in the environment. gondii were found in 85..3. gondii infections in deer.2.5. Michigan and Ohio. Dogs Primary toxoplasmosis in dogs is rare. (Hessler et al.. The most severe disease occurs in pups but to our knowledge there is no documented case of congenital toxoplasmosis in dogs. gondii isolated from tissues of nine of 16 cats. Dubey et al. viciousness. Lagothrix sp. before the lesions develop in the brain. Reported clinical signs in squirrels were anorexia. 2008). (1986a) Dubey et al. J. often with visceral toxoplasmosis. Spencer et al. Bangari et al. An unusual clinical presentation of toxoplasmosis in canaries is blindness with almost complete destruction of the eyes (Dubey.. 1992. including game Among wild animals. Dubey and Beattie. Dubey et al. 2002. (1995c) Hill et al. 1959. 2004). marmoset. Lemur catta (Dubey et al. 2. 1974). Dubey et al.. Anderson and McClure. 2006. (2002b) Rhode Island 84 116 100 76 50 36 DeFeo et al. 1968. Toxoplasmosis in the endangered Hawaiian crow (Corvus hawaiiansis) is of particular interest because there are few (<25) animals left and four animals in the wild died of clinical toxoplasmosis (Work et al.P. 2007). Macacca mulata (Wong and Kozek.. and viable T. There are many reports of severe toxoplasmosis in wallabies and kangaroos in zoos worldwide. (2002) Nutter et al.. (1986b) Smith et al. and they can die even after treatment with sulfonamides and pyrimethamine (Dubey and Crutchley. 2. 2004b).. bears and raccoons are of epidemiological significance and prevalence data are summarized in Table 4. gondii was isolated from both of these hosts (Walton and Walls. 1982). Dubey et al.. Saimiri sciureus (McKissick et al.L..4. 1988). 1989. 2001) and pet birds. Adkesson et al. lethargy.1264 J. 1977.3 80 50 38 62 Reference Dubey et al. Most cases of acute toxoplasmosis in the USA were observed in dogs not vaccinated against the immunosuppressive canine distemper virus (CDV) (Capen and Cole. and wooley monkey. 1951. 2002). hepatitis and encephalitis (Dubey and Lappin. labored breathing and in two cases squirrels had bitten children. 2.. Oedipomidus oedipus (Benirschke and Richart. 2007). Toxoplasmosis in squirrels can simulate signs of rabies (Soave and Lennette. (1998) Dubey et al. (2004) 63 34 Data based on modified agglutination test. 1992). Deer are strictly herbivores and the high prevalence of T. Clinical signs include pneumonia.. 1971). Soave and Lennette. Captive zoo animals and endangered species Certain species of zoo animals.. 2006). mink (Frank. macropodids are highly susceptible to acute toxoplasmosis. 1959). 2000). In one survey.9 68.. 1964. . squirrel monkeys.

Dubey and Beattie. gondii-associated encephalitis in other marine mammals including seals. tested 80 77 100 25 21 10 53 27 18 380 311 32 8 9 117 94 47 146 49 % Positive 36 61 82 52 38 100 5. (2003b) Dubey et al. (2002a) Miller et al. 1997. 2003). in press c).. Although T. (2003b) Lambourn et al. 2005. gondii were found in a variety of marine mammals including sea otters. Sundar et al. (2003b) Dubey et al. and the serological test used. irrespective of the dose (Dubey and Frenkel.5% of sea otters. protozoal encephalitis was considered the cause of death in 39 of 334 (11. Viable T. 2005). 2006). 2003b. there were isolated reports of T. (2008) Miller et al. Pathogenesis of toxoplasmosis including T. 1979..6 50 11. Dubey. Recently. Jones / International Journal for Parasitology 38 (2008) 1257–1278 1265 2.1 1. 2000. J. The high seroprevalence of T. 2.. gondii was isolated from bottlenose dolphins (Dubey et al.1 7.. de Moura et al. 2007). South Carolina South Carolina Hawaiian monk seal (Monachus schauinslandi). gondii was not a major cause of mortality in these sea otters. Transplacental toxoplasmosis is considered rare in marine mammals (Dubey et al.. T.. 2003). (2003b) Aguirre et al.. Conrad et al. gondii were not found in 65 sea otters from Alaska (Miller et al. gondii (Table 6). source (California versus Washington). 2005. antibodies were found in other marine mammals from Alaska (Table 6).6 29.8 100 100 53 Test IFAT IFAT MAT MAT IFAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT MAT Titer 1:320 1:25 1:25 1:320 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 1:25 Reference Miller et al. dolphins. oocyst-induced infections are more severe than those induced by tissue cysts and bradyzoites by the natural oral route. 1973. depending on the category (live versus dead). North Carolina and South Carolina.7 96.. seals and walruses (Table 6). 2008). (2003b) Dubey et al. gondii infection in marine mammals in the USA are most intriguing. Thus. (2008e) Walruses (Obobenus rosmarus) Sea lions (Zalophus californianus) Harbor seals (Phoca vitulina) Ringed seal (Phoca hispida) Bearded seals (Erignathus barbatus) Spotted seals (Phoca largha) Hawaiian monk seal (Monachus schauislandi) Bottlenosed dolphin (Tursiops truncatus) . 2007). Prevalence in sea otters from California was higher than in Washington otters.4 15.. T. Experimentally. Antibodies to T.L. (2003b) Sundar et al. Miller et al. Aguirre et al. (2005a) Dubey et al. It is unknown whether the severity of toxoplasmosis in immunocompetent individuals is due to the parasite strain.. Before this. gondii genotype Only a small percentage of exposed adult humans or animals develop clinical toxoplasmosis.6 16. 2002b. (2003b).. 2008).. Sarcocystis neurona in seven otters (Kreuder et al. (2007) Dubey et al. A very high seroprevalence of T. gondii and 12 had dual infections (Thomas et al. (2003) Dubey et al. (2002a) Dubey et al. (2003b) Dubey et al. Severe clinical toxoplasmosis was reported in humans and was linked epidemiologically to ingestion of T. indicating water in the Pacific Ocean is infected (Honnold et al.. (2003b) Dubey et al. Bowie et al.. viable T. (2003b) Dubey et al. gondii was considered to be the primary cause of death in 17 and a related parasite.. gondii in sea otters was high but varied between 47% and 100%. 1997b). Otters with preexisting T.P. (2001) Dubey et al.6 61. Florida.. Both antibodies and clinical toxoplasmosis were found in the Table 6 Prevalence of Toxoplasma gondii antibodies in marine mammals in the USA Species Sea otters (Enhydra lutris) Source California – live Dead Live Dead Washington live Dead Alaska Alaska California Washington Alaska Alaska Alaska Alaska Hawaii California Florida Florida. host variability or other factors. except that there is a high prevalence of eye dis- No. Dubey. Benenson et al.. neurona. 2002a). Miller et al.7.6. (2002a) Sundar et al. In another investigation involving otters that died in California and Washington State but necropsied at the National Wildlife Health Center in Wisconsin. 1982. gondii was isolated from 85 sea otters from California and Washington (Cole et al. Marine mammals Recent findings of T. indicating that marine mammals on both coasts of the USA are exposed to T.J. 1988. gondii antibodies in sea otters has been verified parasitologically. gondii encephalitis were considered prone to shark attack (Kreuder et al. Dubey et al. The sea otter is listed as an endangered species (Conrad et al. five with T. Two groups of researchers made detailed studies of causes of mortality in sea otters. gondii oocysts in food or water (Teutsch et al.. 1997. Of the 105 otters necropsied at the Californian laboratories from 1998 to 2001. (2008) Dubey et al. Little is known about the effect of race or geography on clinical toxoplasmosis in humans.. sea lions and dolphins.3%) sea otters. and these were summarized by Dubey et al. of these 22 were infected with S. In a preliminary report Thomas and Cole (1996) reported protozoal encephalitis in 8. gondii was found in the bottlenose dolphin (Tursiops truncatus) from California. Prevalence of T.

We are not aware of a risk assessment study in the USA but in a retrospective study of 131 mothers who had given birth to children infected with T. J. similar to strains from clinical cases of humans in Brazil (see Khan et al. gondii isolates from apparently healthy and sick hosts. gondii have been found in sea otters from California and Washington (Miller et al. Therefore... 2008). Ajzenberg et al. most of the retail chicken sold in the USA is frozen. Lehmann et al. gondii in retail meat. Role of infected meat In the USA poultry. which also kills T. T. 1992. Circumstantial evidence suggests that certain genetic types of T. Most of the evidence is based on epidemiological investigations and prevalence studies in animals. gondii tissue cysts (Hill et al. Therefore. gondii-infected meat..3%) were Type III.L.5 billion chickens are killed annually for human consumption in the USA.1. Whether these new T.. 2003. Pena et al. gondii was considered to be clonal with very little genetic variability. Jones / International Journal for Parasitology 38 (2008) 1257–1278 ease in southern Brazil (Glasner et al. 1995) and until recently. seven (8. these three otters also had other potentially fatal conditions (Sundar et al. in another study. beef or game) was identified as the main risk (Cook et al.. Toxoplasma gondii is one of three pathogens (together with Salmonella and Listeria) which account for >75% of all deaths due to foodborne disease in the USA and economic cost to care for congenitally-infected children are high (Roberts et al. of the 86 T.. pork and beef are the main meat types consumed. 2005).. 1997.1266 J. there is no national identification system for individual pigs destined for human consumption and pigs are not tested for T. Approximately 100 million pigs. gondii for humans. Grigg et al. There is very little information regarding the genetic diversity of T. ingestion of infected tissues and ingestion of oocysts are the three main modes of transmission of T.. 2002. Silveira et al. 2002. Toxoplasma gondii isolates have been classified into three genetic Types (I. gondii infection transplacentally..... gondii infection was considered incidental (Sundar et al. 2006). 50% recalled having eaten uncooked meat (Boyer et al. The proportion of the human population that becomes infected by ingesting T. gondii as the primary cause of meningoencephalitis in nine of 12 otters. 3. Khan et al. 2005.1. In the remaining 34 otters T... gondii.1%) were Type I and four (4. gondii Congenital infection. gondii infection in market pigs alone cannot account for the 10–40% seroprevalence in humans in the USA (Jones et al.. Boothroyd and Grigg. 2007). Overall. Epidemiology and transmission of T..or region-specific. 2005).. but genetic characterization has essentially been limited to isolates from patients with toxoplasmosis. (2004) observed localized clustering of genotype Type X near Morro Bay and reported Type X T. food or water contaminated with oocysts is unknown and currently there are no tests to distinguish meat. Recently. Miller et al. 2008).8%) were Type II. 2005b). In France. Mead et al. Holland. gondii strains with atypical genotypes (Ajzenberg et al. 2005.. III) based on restriction fragment length polymorphism (RFLP) (Howe and Sibley. Two new genotypes (Types A and X) of T. less than 1% of humans and livestock acquire T. gondii-infected pigs from . gondii infection at slaughter. 3. Dubey et al. 2004. the routes by which T.. 2007). 6%) were atypical.. In a recent nationwide study of the prevalence of T. 2006). In humans in French Guiana and Suriname. gondii. two (2. Dubey. attention has been focused on genetic variability among T. 2008). 2001.P.versus oocyst-acquired infections. In a multicentre European study of pregnant women.. Some of the salt treatments (labelled as ‘‘enhanced” meat) kill T. 2003). we must be cautious in claiming a linkage between parasite genotypes and disease presentations without more complete knowledge of the T. 73 (84. while the scope of human infection resulting from meat sources remains undetermined. 2006) have been found in asymptomatic chickens and cats from Brazil (Dubey et al. Khan et al. However. 2006 and references contained therein). gondii genotypes in human populations and the environment. needs further investigation. Serological or parasitological surveys based on slaughterhouse samples do not provide a true assessment of risk to humans because nearly half of the pork and a substantial amount of chicken in retail meat is injected with salts and water (Dubey et al. gondii.. gondii genotypes (Types A and X) are host. gondii isolates circulating in the general human population...1. Sundar et al. II... In the following section we will discuss meat sources of T. gondii isolates obtained from patients with clinical toxoplasmosis. there was no apparent association between severity of disease and genotype (Ajzenberg et al. gondii may be associated with clinical toxoplasmosis in humans in the USA (Howe et al. 2001. Pigs Currently.. Mouse-virulent strains with atypical genotypes. 1999). 3. 1994.. and their association with mortality. 2004. Based on newer markers for genetic characterization and using recently isolated strains.. gondii was considered a contributing cause of death in only three of 37 (25 from California and 12 from Washington) sea otters. It has been suggested that Type I isolates or recombinants of Types I and III are more likely to result in clinical toxoplasmosis (see Khan et al. 2006. Conrad et al. 2005. viable organisms were isolated from only seven of 2094 pork samples and none of 2094 beef or 2094 chicken meat samples (Dubey et al. severe cases of toxoplasmosis in immunocompetent patients have been related to T. a higher genetic variability has been revealed than was previously reported (Ajzenberg et al. 2007.. 2002. Further. 2000). T. 2002). Demar et al. the low prevalence of T. 2008). 2004. 2004). The surge of infections in teenage and low prevalence in young children suggests that transmission by meat is important in the USA. Thus. ingestion of inadequately cooked meat (lamb. 30 million cattle and 8.

2005b). 1982).P. Chickens In the USA.1.2 kg and approximately 8. three persons developed symptomatic toxoplasmosis linked to eating Kibee Nayee (a meat dish made with raw beef) at a Syrian restaurant (Lord et al. Riemann et al. There are several reasons why the results of this study do not negate the possibility that infected chickens may be important sources of infection for humans. and the per capita consumption of lamb meat in the USA is approximately 0.L.3. 1997). Cattle The ingestion of beef or dairy products is not considered important in the epidemiology of T. These data suggest that T. gondii in chickens consumed by humans and it will be necessary to cook the meat properly to protect consumers from infection. even a 1% infection rate would amount to 1 million infected pigs going to market for human consumption. 2005b). We are not aware of any report of toxoplasmosis in humans directly linked to eating infected pork in the USA but clinical toxoplasmosis and blindness were linked to the ingestion of undercooked pork in Korea (Choi et al. (1975a) reported on an infant that developed toxoplasmosis after drinking raw milk from goats. In this study.6% of infected chickens (Dubey. gondii.. chicken breasts were selected for sampling because of the experimental design that required testing of 1 kg of boneless meat for each sample. breeders are probably not important with respect to transmission of T. 1975). 1969). gondii was not isolated from any of the 2094 chicken meat samples obtained from retail meat stores in the USA (Dubey et al.4. gondii were found in 1.1. while older animals are classified as sheep and their meat (mutton) is sold for pet food and export. However. 1986c)... A 50 kg market pig would account for over 300 servings of meat.. 1982).5 million lambs are slaughtered in the USA for food each year. fever. Jones / International Journal for Parasitology 38 (2008) 1257–1278 1267 highly endemic areas enter the market and the role these pigs have in the overall epidemiology of T. antibodies toT. Between 3 and 3. The recent trend of consumers demanding meat from organically grown free-range poultry will increase the prevalence of T. In another instance. 3. 3. 1975. T. gondii survives the processing procedures. gondii does occur in commercially marketed chickens in the USA but processing and handling procedures inactivate the organisms prior to sale to consumers.2. he never ground lamb in the same grinder and when pork was ground it was at the end of the day and the grinder was washed thoroughly after use. with values six times higher than in control chicken sera (Dubey et al. Milk from goats is more easily digested than cow’s milk by children. in press). Any part of infected pork can be a source of infection because T. gondii is declining. 3.. milk from goats has been implicated in human toxoplasmosis (Riemann et al. However.1. Finally. sheep <1 year old (without permanent teeth) are classified as lambs and slaughtered for human consumption. gondii in humans remain unknown. Department of Agriculture regulations. Sacks et al. In contrast to the bioassay results. four of the 10 goats had antibodies to T. gondii in chicken eggs is extremely low and the ingestion of uncooked eggs is not considered an important risk for toxoplasmosis (Dubey.usda. lymphadenopathy. Goats The amount of goat meat consumed in the USA is unknown but goat meat is popular with several ethnic groups. gondii was isolated from breast meat of only 18. Thus. Symptomatic toxoplasmosis in a family in New York City was circumstantially linked to eating rare lamb (Masur et al. gondii parasites.. gondii in chicken breast is lower than in other tissues. The restaurant had bought the meat from a local butcher who insisted that the meat was unadulterated beef. gondii is highly susceptible to freezing. Further. although the labels indicated otherwise. myalgia and splenomegaly during the second week after eating rare hamburgers at a university snack bar (Kean et al.1. 1978).and naturallyinfected pigs (Dubey et al. For example. the per capita yearly consumption of poultry is estimated to be 37. Sheep According to U. eggs should be cooked thoroughly before human consumption.. one 39-year-old woman had chorioretinitis (Sacks et al. two small outbreaks of toxoplasmosis were linked to ingestion of infected beef. 2005 www. especially from Asia. The prevalence of T. .5 kg per year (NASS Agricultural Statistics. In addition to infected meat. J. gondii in lambs can be high (Tables 3 and 4) but the role of ingestion of infected lamb in the epidemiology of toxoplasmosis in humans remains to be determined.gov/nass/pubs/agr05/agstats2005. In the USA lambs and sheep are slaughtered in separate commercial slaughter facilities. Results of a recent study and previous surveys indicate the prevalence of T.J. Epidemiologically. T. gondii to humans.. although the authors were aware that that the prevalence of T.pdf). Dubey. both in experimentally. in press).. 3. In a recent survey. A small outbreak of toxoplasmosis in humans was attributed to drinking goat milk. Five medical students developed symptoms of acute toxoplasmosis characterized by headache. some of the samples collected might have been frozen or hard chilled. gondii has been found in most edible tissues or cuts of meat. gondii transmission as only relatively small amounts of beef have been tested for viable T.5.3% of the juice extracted from the breast meat using an ELISA. gondii because cattle are not a good host for this parasite.S..5 billion chickens are killed annually for human consumption. gondii. many of the chicken breasts had been injected with enhancing solutions that have a deleterious effect on T. we cannot be sure that beef does not play a role in T. Standards of hard chill are vague and T. Although the prevalence of T. Meat from breeder pigs is generally processed for sausages and it is highly unlikely that T.

1983).L. based on the observation that most cats only shed oocysts for about 1 week in their life (Dubey. have been documented in humans who had consumed undercooked venison. gondii could not be determined by PCR and bioassays were not performed. Hundreds of bear. tested 510 1604 1000 16 274 No. whether these oocysts were T. There are approximately 78 million domestic cats and 73 million feral cats (reviewed by Conrad et al. gondii were found in 30–60% of deer (Table 3) and viable T.000 viable oocysts were found in rectal contents of an asymptomatic cat on a sheep farm in Maryland (Dubey et al. gondii oocysts morphologically from oocysts of related parasites in cat feces. gondii-like oocysts in feces of 48 of 24. We are aware of only a few surveys for T.b) estimated an annual burden of 94 to 4671 oocysts/m2 in California. Toxoplasma gondii oocysts were detected in cat feces.. 150. Jones / International Journal for Parasitology 38 (2008) 1257–1278 3. 1995d). Antibodies to T.. 3.6. gondii oocysts in cat feces because half of the cats were shedding H.106 cats from Germany and other European countries. However. In two large surveys (Wallace. In the most extensive study performed to date. 1995. Schares et al. gondii are prevalent in white-tailed deer in the USA. Dabritz et al.pdf).5 cats per farm. gondii and 22 (0. because bioassays are needed to distinguish T. 2006). gondii oocysts in feces. by ingesting either infected tissues or sporulated oocysts. Recently. At any given time. (2007a. there is strong potential for T. a mean of 8. 2001. 2001).dnr. gondii. gondii-infected mouse (Dubey and Frenkel. antibodies to T.8) Reference Dubey (1973) Wallace (1971) Dubey et al. seroprevalence data are more meaningful than determining the prevalence of T. 2001). 1993b... In one reported case. gondii oocysts in cats in the USA (Table 7). 1972). 1972). gondii post-natally. Millions of oocysts were shed by cats fed even a few bradyzoites (Dubey.1268 J. Most cats become infected with T. Under laboratory conditions.. including ocular manifestations (Ross et al. During the 2006 deer hunting seasons in Iowa and Minnesota. 10.552 and 270.us/outdoor_activities/hunting/ deer/2006_harvestreport. Thus. gondii can occur but is rare in domestic cats and congenitally-infected kittens can shed oocysts (Dubey and Carpenter.state. (1977) Dubey et al..7) 1 (6.. cats can shed as many as 500 million oocysts after ingesting one T. Dubey et al. This study also demonstrates the importance of proper identification of T. eviscerated tissues of these animals left in the field are a source of infection for carnivores. Using a titer of 1:25 in MATs as a positive cut-off. gondii oocysts (Miller et al. (1986b) Dubey et al. gondii oocysts were present per gram of cat feces (Schares et al. positive (%) 0 12 (0. gondii oocysts were found in of only a few (<1%) cats. 1995c. (2007b) detected T. The prevalence of T. Both domestic cats (F. gondii was isolated from 17% to 28% (Table 4). Weigel et al.P. gondii in black bears is very high (Tables 3 and 4). It is probable that every farm in the USA has cats. Dubey.. 1999). wild pig and other game are harvested in the USA each year. Dabritz et al.. Toxoplasma gondii transmission is more efficient through ingestion of infected tissues than ingestion of oocysts (Dubey. (2008) found T.7) 7 (0.. moose.11%) were identified as T. Most cats seroconvert after they have shed oocysts (Dubey and Frenkel. soil or water samples on six farms (Dubey et al. 1979). approximately 1% of cats are expected to shed oocysts. T. In one study of pig farms in Illinois. more data are needed on the actual numbers of oocysts/g of feces in naturally-infected cats in the USA. 2004). hammondi oocysts which have no zoonotic significance. elk. Up to 13 million T. 1999). Role of oocysts Environmentally-resistant oocysts are essential in the life cycle of T. These surveys are time consuming and expensive. 1986b).7.2. with a mean of six seropositive cats on each farm (Weigel et al. Congenital transmission of T. 2008). including cats. 1977). 1971.. feed.1. gondii has been isolated from horses slaughtered for export.2) 5 (1. 2005).mn. gondii transmission in rural settings. In addition to the possibility of transmission to humans. Dubey et al. Venison and other game Deer are popular game animals in the USA and the deer population is estimated in millions. Approximately one-third of households in the USA own a cat and this number is steadily increasing. Thus. horse meat is not used for human consumption in the USA (Al-Khalidi and Dubey. of these 26 (0. The number of oocysts shed by naturally-infected domestic cats is largely unknown. 1972). (1995c) . 3. it is a reasonable assumption that most seropositive cats have already shed oocysts. 366 cats were trapped on 43 farms. For epidemiological studies. If one assumes a 30% seropositivity of 151 (78 domestic and 73 feral) million cats and a conservative shedding of 1 million oocysts per cat then there will be enormous numbers of oocysts (50 million  1 million) in the environment. Horses Although viable T.778 deer were harvested in the respective states (http://files. J. Only felids are known to excrete T. gondii-like oocysts in feces of three of 326 cats from the Morrow Bay area of California.09%) as Hammondia hammondi. Table 7 Isolation of Toxoplasma gondii oocysts from feces of naturally-exposed cats in the USA Cat type Homes Shelter Shelter Sheep farm Pig farms Locality Kansas Hawaii Ohio Maryland Illinois No. Cases of clinical toxoplasmosis (Sacks et al.1. domesticus) and other felids may shed oocysts.

1975). (2001) Roelke et al. Dubey. A large waterborne outbreak of toxoplasmosis in humans was epidemiologically linked to oocyst contamination of a water reservoir in British Columbia.. Contamination of sea water with T. a sylvatic cycle of T. % Positive 73 72 69 44 18 66 83. The number of bobcats in the USA is thought to be millions and one study estimated thousands of cougars (Conrad et al.. It is noteworthy that 12. viable oocysts were detected in rectal contents (samTable 8 Serological prevalence of Toxoplams gondii in large wild cats in the USA No.. for 334 days when covered (Yilmaz and Hopkins. eviscerated tissues (gut piles) from hunted deer and black bears would be a source of infection for wild cats. Dubey. MAT. latex agglutination test. Dume Direct detection of viable oocysts in drinking water in the USA has not been achieved. uncovered. 1998). gondii (Table 8). Toxoplasma gondii oocysts are highly resistant to disinfectants but are killed by temperatures above 60 °C (Dubey et al. b Viable T. Isospora felis. 1995).. Young and weak white-tailed deer and small mammals are common prey for bobcats and 60% of white-tailed deer in Pennsylvania. indirect hemagglutination test. gondii has not been established in the USA.P. 2004b). 1970. coinfection with a related coccidian. Although oocysts were not detected in drinking water taken from the reservoir after the outbreak (Isaac-Renton et al.. (2004) Kikuchi et al. (2006) Riley et al. gondii oocysts is technically difficult. (2004) Paul-Murphy et al. can cause reshedding of large numbers of T. gondii oocysts from chronically infected cats (Chessum. gondii isolates from these cougars now suggest that both faecal samples might be from the same cougar (Dubey et al. modified agglutination test. (1993) Cougars (Felis concolar) Lynx (Felis lynx) Panther (Felis concolor coryi) DT. 83% of 131 bobcats were found to have T. Although cats can be reinfected with T. but oocysts have been isolated from animal feed and soil from pig farms (Table 7). Wainwright et al. gondii in rural USA is feasible and appears to be efficient. 1972. Ultraviolet rays also have a deleterious effect on oocysts. (1975b) Franti et al. 2007a). It is anticipated that the heat generated and lack of oxygen will kill some or all oocysts. Dubey. Bobcats fed with tissue cysts shed T.. 3. dye test. (2004b) Mucker et al. gondii antibodies (Mucker et al. The fate of cat feces disposed of in the toilet or in domestic trash destined for landfills is unknown. The role of cougars in the sylvatic cycle of T. 1995). gondii was isolated from five of six bobcats from Georgia (Dubey et al. 1998). depending on the conditions. for 46 days. grain or hay.2. kinetic enzyme linked immunosorbent assay.5 million oocysts were present in sample A and 250. 1995).. However.L. 2004. The bobcat (Lynx rufus) and cougar (Felis concolor) are the two main wild cats in continental USA. 2007b. depending on the dose (Wainwright et al. 2006) and viable T. Little is known of the sporulation or survival rate of oocysts openly exposed to sun and other environmental conditions. grass. In addition to live prey. LAT. (1979) b Dubey et al. (1975) Marchiondo et al. gondii they are considered to shed oocysts only once in their life based on short-term experiments in the laboratory (Davis and Dubey. In one long-term experiment four of nine cats reshed oocysts after challenge (Dubey. Canada (Bowie et al. IHAT. It is likely that oocysts are carried into our homes on shoes contaminated with oocysts on street pavements.3 83 88 50 58 19. gondii oocysts (Miller et al. Ideally..1. In a recent survey in Pennsylvania. domestic cats like to bury their feces in soft and moist soil but one can find cat feces on street pavements... This washing is apparently very effective in removing dirt and feces from their body hair (Dubey. Assessing environmental contamination with T. 1998). 2008)... gondii infection in California sea otters (Miller et al. 1972). 1972) and outdoors in soil buried at the depth of 3–9 cm in Kansas for 18 months (Frenkel et al. 1992.. gondii oocysts Freshwater runoff has been suggested as a risk factor for T. Oocysts survived outdoors in Texas (6–36 °C) in native cat feces. Coinfection with FIV does not cause reshedding of T. West Virginia Florida Georgia Pennsylvania California USA California USA Alaska Florida Reference a Walton and Walls (1964) Riemann et al. gondii isolated from hearts of the five seropositive bobcats. USA were seropositive for T. Dubey. ` tre et al.. . gondii oocysts (Lappin et al.J. (2004) Zarnke et al.1 15 9 Locality Georgia California California New Mexico Georgia. 1995). gondii isolated from brain of one of 16 bobcats. 2005).. 1976). Thus. (1976) Oertley and Walls (1980) Burridge et al. (1994) Kikuchi et al. ELISA.. they clean their feet and body by licking. examined Bob cats (Lynx rufus) 15 12 86 27 150 3 6 131 25 52 36 320 255 56 Test DT IHAT IHAT DT IHAT IHAT MAT MAT MAT LAT LAT LAT MAT ELISA Titer 1:4 1:64 1:64 1:8 1:16 1:64 1:25 1:25 1:25 1:64 1:64 1:64 1:25 1:48 ple A) of a wild trapped cougar (Felis concolor vancouverensis) and in a fecal pile (sample B) in the vicinity of the reservoir (Aramini et al. J. 1997).000 in sample B (Aramini et al. a Viable T. Genotyping data on the T. Jones / International Journal for Parasitology 38 (2008) 1257–1278 1269 Cats bury or hide their feces and unless they are ill. 2008a).

How much marine water is cycled through the gut of sea otters in a day is unknown but it is likely to be large volumes. gondii in zoos is of special importance because many species of captive animals are highly susceptible to clinical toxoplasmosis and there is the possibility of transmission to zoo visitors. 2005). 1988a. mostly descendents of cats imported from Russia in 1994 (Kenny et al. 2007). USA (Teutsch et al. gondii as illustrated by the following two studies. Dubey. are not a host for T. fever. gondii tachyzoites and although the parasite survived for various amounts of time. lymphadenopathy and abortion in one of three pregnant patrons. there was no evidence that it multiplied in cold blooded animals (Dubey and Beattie.2.. it is not known if the sporozoite excysts after ingestion by cold blooded animals. an isolate from one cat was as mouse-virulent as the isolate from the human fetus. The natural habitat of Pallas cats is the high mountains of Tibet... Toxoplasma gondii infection of other marine mammals that mainly eat fish is even more intriguing. however. Dubey et al. canaries and finches.. an outbreak of acute toxoplasmosis occurred in patrons of a riding stable in Atlanta.3.g. Dubey et al. An epidemiological investigation suggested that the patrons acquired T. 1997a). gondii oocysts. Lindsay et al. sea otters might ingest oocysts directly from marine water. gondii before pregnancy can repeatedly transmit T. including fish. Historically. and the first documented outbreak linked to inhalation or ingestion of oocysts.. With respect to pathogenicity of the T. 1979).. 1988. gondii were isolated had no detectable antibodies to T. gondii oocysts (Arkush et al. squirrel monkeys) die of acute toxoplasmosis (Dubey and Beattie. Basso et al. Contamination of zoos with oocysts shed by Pallas cats and other felids Transmission of T. She aborted in December.2.L. Jones / International Journal for Parasitology 38 (2008) 1257–1278 2002b.. cougars and panthers (Table 7) which are native to the USA. Unlike humans. 2002)... The prevalence of T. 1988. and this difference was attributed to the absence of cats on Ossabaw island (Dubey et al. Unusual aspects of the outbreak were: a very high rate (95%) of clinical toxoplasmosis in exposed persons. Attempts were made to isolate T.1270 J. Bulldozing of the arena between collection of samples 1 and 2 might have contributed to differences in results obtained with these two samples. 2005). although attempts to isolate oocysts from 29 samples of soil. J. That woman was in her first trimester at the time of the outbreak. Molluscs. Seroprevalence of T. With respect to oocysts. gondii in bottlenose dolphins from both coasts of the USA is very high (Dubey et al. gondii (Omata et al. 2002). there were no genetic markers available and those isolates were not cryopreserved.9% of 1264 pigs) compared with 18. Before the discovery of the oocyst stage of T. gondii to their kittens (Basso et al.P. 2005). All four mice and the five cats from whose tissues viable T. 1977 (sample 1) but not from 12 mice.. and four of four mice trapped in the stable in November. a burden of 36 oocysts/m2 over the region was calculated by Dabritz et al.. gondii isolates. 2005) and acute toxoplasmosis is a leading cause of mortality in Pallas cats (Riemann et al. Kenny et al. gondii from oocysts aerosolized during the riding activity. 1977 and viable T. Conrad et al. three of three adult cats. 1980. 2003b. Certain species of macropodids (e. this was the first large outbreak in humans linked to ingestion of oocysts. Turkestan and Mongolia. Control. gondii in pigs and mice on pig farms in Illinois. wallabies).. can act as transport hosts for T. 2008). Pallas cats can shed T.. (2007a). In addition to bobcats. Atlanta stable outbreak of human toxoplasmosis In October 1977.. 2003.. Thirty-five of 37 patrons of the stable had clinical toxoplasmosis characterized by headache. 3. 2004). USA was greatly reduced when cats on these farms were vaccinated orally with a strain of T... Sea otters eat approximately 25% of their body weight in invertebrate prey each day (Conrad et al. Dubey. Cold blooded animals.g. in press c). At that time. 1977 (sample 2). three rats and four cotton rats trapped in the stable in December. There are a few Pallas cats in USA zoos. gondii at a 1:2 serum dilution (prozone was excluded) of serum tested using a dye test. 1988). 2005).. Based on an estimate of 10 million oocysts shed by each cat and a 36% defecation rate outdoors.. gondii from animals trapped in and around stable. and all those are potential shedders of T..2. 1981b). gondii might be related to insensitivity of the dye test for cat sera. Of particular interest is the housing of Pallas cats (Felis manul) in zoos. numerous experiments were done to infect various invertebrates including arthropods with T. prevention and future developments Cats are key to the transmission of T. doses of one tachyzoite. sand and sawdust from different parts of the stable were unsuccessful (Dubey et al. Georgia. gondii. gondii were isolated from the fetus’s amniotic fluid (Teutsch et al. These results indicated that asymptomatic animals can harbor mouse-virulent T. gondii in pigs from a remote island (Ossabaw Island. Several aspects of this episode are epidemiologically and biologically interesting and therefore recalled here. Viable T. gondii were isolated from tissues of two of four kittens. In addition. one bradyzoite and one oocyst were lethal to mice (Dubey et al. 2002). western Siberia. gondii oocysts (Dubey et al. gondii. The seroprevalence of T. many other felids are kept captive in zoos throughout the USA (de Camps et al.. These authors considered this level of land contamination is likely to be high enough for oocysts to reach marine waters (Dabritz et al. 1981b). and New World monkeys (e.2% of 170 feral pigs from mainland Georgia. Results also indicated that timing for sampling is important in an epidemiological investigation because different results were obtained with samples 1 and 2. Georgia) was very low (0. 1974. 1981b). gondii that does not produce . Pallas cats infected with T. Seronegativity of the five cats and the four mice with demonstrable T. 3. 4.

Pelloux. Antonelis. a recently reported large European multicenter cohort study found no evidence that pre-natal treatment with either spiramycin or sulphonamide combined with pyrimethamine had an effect on maternal transmission (Gilbert and Gras. 97–100. C. Rabaud. Filisetti. gondii-positive infants are identified.. ´ . To screen for congenital toxoplasmosis is controversial (Kim. References Abgrall.A. M. However. Clin. 229–241. gondii in wild game and venison in the USA is very high and hunters need to be aware of the risk of transmission of infection to humans and. gondii infection in women before they reach childbearing age (and therefore fewer women susceptible to acute infection during pregnancy)...... Infect. spread of infection in the environment. Infect. 34. J. Gorman... D. However.J. then medication is switched to pyrimethamine and sulfadiazine (Montoya and Liesenfeld.. Alford Jr.. Because T. more importantly. Dis. Reif.P. 36.. However. Yochem.. gondii to humans. M. Acknowledgements The findings and conclusions in this report are those of the author(s) and do not necessarily represent the views of the Department of Health and Human Services or the Centers for Disease Control and Prevention or the U. 2001.L. controlled trials demonstrating the effectiveness of neonatal screening and treatment (Petersen. 2007. with a limited pre-natal screening program there would be 18. H. Paris. Int. M. Bessie Ajzenberg.S. 2006). clonality and sexuality in Darde Toxoplasma gondii. 684–689. gondii transmission risk because feral cats can spread oocysts in any suitable location. A meta-analysis of 22 European cohorts found weak evidence that treatment started within 3 weeks of seroconversion reduced mother-to-child transmission compared with treatment started after 8 weeks or more (Thiebaut. Although there have been neonatal screening programs for toxoplasmosis in Denmark and Brazil.T. Marty. gondii infection and treatment of the mother and infant has been conducted in countries with a higher prevalence of T. Bull. One current practice is to start a woman on spiramycin if she becomes newly infected during pregnancy in the first trimester. with confirmatory testing in the mother when T.E. C. Keefe. Dubey.. Darde Toxoplasma gondii isolates associated with human congenital toxoplasmosis. 2007). M.J. Dubey.... 2002. Kashinsky. ` res. (1997) determined that in countries with low T. Infectious disease monitoring of the endangered Hawaiian monk seal.S.. 1974. 1999). Parasitol. Pre-natal screening for T. Saliki. NY Acad.. practicing good hygienic measures appears to be the best option to minimize transmission of T. 1992). but follow-up results for more than 6 years (as in Guerina et al.. D. with special reference to subclinical disease. J. ` tre.. Wildlife Dis. Bader et al.. L.. 2003).W. In another series of infants referred for care. V. J. even with a relatively low amniocentesis complication rate (0. 2004)... Reynolds. 1990c). gondii infection in animals and humans. Prevalence of T. 186. A.K. 43.J. Costagliola. Department of Agriculture. France (monthly if seronegative) (Thulliez. 1185–1196.. and therapeutic considerations.H.L. Braun.L. gondii infection.N. for example.. The viscera of hunted animals need to be buried to prevent scavenging by animals. Thulliez.. Incidence and risk factors for toxoplasmic encephalitis in human immunodeficiency virusinfected patients before and during the highly active antiretroviral therapy era. resulting in a relatively high cost compared with costs in countries with a higher prevalence of T.. A. and correlation with clinical findings.. Ajzenberg. S. with none on the horizon. There is currently no non-viable. because most women of childbearing age in the USA (89%. A. N. tissue cysts in meat are easily killed by freezing meat in a household freezer (Kotula et al. effective vaccine to prevent T. P. 1994). Langan. Although oocysts are almost indestructible. 2006).. J. J..Clinical Epidemiology Group of the French Hospital Database on HIV.L. Carme. Hsiao. it is difficult to establish the definite modes of transmission on an individual basis. J. randomized studies of newborn screening and treatment would be costly because of the large population needed for evaluation. 2004.. Educational programs directed at reducing environmental contamination with T.. S. Vet. Jones et al. Genotype of 86 D. Jones / International Journal for Parasitology 38 (2008) 1257–1278 1271 oocysts in cats but immunizes them against shedding of oocysts (Mateus-Pinilla et al. P. 2007) are susceptible to T.5 fetal losses for each congenital toxoplasmosis case avoided. If fetal infection is detected. Early results suggested that treatment of infants in this program may be beneficial.. Genetic diversity. 2007). B. Stagno. G.36% fetal death). J. Pathol. Adkesson. most would need to be screened repeatedly during pregnancy. In addition..A. J. M. Therefore. laboratory. R. it is important to note that maternal screening and treatment are not without risk.. T. C. Stott. 1747–1755.. L. 2007. there are no randomized. Dume ´ . Ban ˜ uls. . gondii would eventually curtail the cost of treating humans for clinical toxoplasmosis. and then perform amniocentesis to detect fetal infection. D. D. 1992) and Austria (each trimester if seronegative) (Aspock and Pollak. Goldstein. Aguirre. initiating treatment for congenital toxoplasmosis after birth produced results which suggest a benefit from infant treatment compared with historical controls (McLeod et al. gondii is transmitted by multiple modes and sources. 50. Demar.. P.. 33. Med. Cogne ´ .. 1994) are needed to determine outcomes. gondii prevalence such as the USA. Toxoplasma gondii inclusions in peripheral blood leukocytes to a red-necked wallaby (Macropus rufogriseus). Whittington. Infants are treated for 1 year.L. gondii is conducted in Massachusetts (started in1986) and New Hampshire (started in 1988) (Guerina et al. Infant heel stick filter paper blood spots are tested. especially cats... gondii infection than the USA. 160–181.K.. Clin. J. M. 1991) and by cooking until the internal temperature reaches 66 °C (Dubey et al. Pyrimethamine and sulfadiazine may be used initially in the late second and third trimesters when acute infection is detected.P. T. Risk assessment researchers should bear in mind that cats are everywhere in the USA and owning a cat does not directly relate to T. Su.. Dis.. Congenital toxoplasmosis: clinical. J. Newborn screening for T. There is still a need for cost–benefit studies to evaluate pre-natal screening for toxoplasmosis in the USA.

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