The Primate Malarias

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Front Matter -- 1. Evolution of the Primate Malarias -- 2. Historical Review -- 3. Ecology of the Hosts in Relation to the Transmission of Malaria -- 4.

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Life Cycle and the Phenomenon of Relapse -- 5. Plasmodium vivax -- 6. Plasmodium cynomolgi -- 7. Plasmodium eylesi -- 8. Plasmodium gonderi -- 9.

Plasmodium hylobati -- Plasmodium jefferyi -- Plasmodium pitheci -- Plasmodium schwetzi -- Plasmodium simium -- Plasmodium youngi -- Plasmodium ovale -- Plasmodium fieldi -- Plasmodium simiovale -- Plasmodium malariae -- Plasmodium brasilianum -- Plasmodium inui -- Plasmodium rodhaini -- Plasmodium falciparum -- Plasmodium coatneyi -- Plasmodium fragile -- Plasmodium reichenowi -- Plasmodium knowlesi -- These molecules fulfill a variety of cellular roles, including participation in key processes such as N-glycosylation, electron transport ubiquinone , and protein prenylation.

Ioprenoid biosynthesis in apicomplexan parasites occurs via a metabolic pathway housed in the apicoplast, known as the methylerythritol phosphate MEP pathway. Because this organelle is cyanobacterial in origin, the MEP pathway is shared by the majority of eubacteria and other plastid-containing eukaryotes, such as plants and algae.

Merozoites, gametes and sporozoites are haploid. The only diploid stage is the 'young' zygote, just after fertilisation. The dividing stages, such as schizonts and oocysts are 'polyploid', because DNA replication and nuclear division is not immediately followed by cell division, resulting in a 'syncytial' cell with many nuclei. The ookinete stage has a nucleus containing the 'tetraploid' amount of DNA resulting from fertilisation and meiosis without immediate nuclear division.

Nuclear division only starts in the oocyst stage.

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During asexual blood stage development DNA synthesis starts around 16 hour after invasion in the old trophozoite, just before the first nuclear division in the schizont. Throughout schizogony DNA replication and genome segregation are alternating events. During male gametogenesis, three rounds of genome replication take place within 10 minutes after activation of the male gametocytes.

The content of resulting 'octoploid' nucleus is divided over the eight gametes, resulting in the haploid male gametes. It has been calculated that the entire haploid genome is replicated in, on average, 3. This suggests that this DNA synthesis represents the genome replication during the first meiotic division like in other eukaryotes.

See Figs. Due to the lack of condensation and the small size chromosomes cannot be visualised by light-microscopy or by standard electron microscopy. The total length of a telomere is about These repeats and variation in copy number have been characterised in detail. The genomes of RMP contain a number of multigene families located in the subtelomeric chromosomal regions.

Animal handling by the CDP veterinarians complied with the criteria for animal care defined in the guidelines of the American Society of Mammalogists www. Note that five of these sequences the ones from Asian rodent Hepatocystis , although published by Genbank, have not been associated to any publication so far.

For phylogenetic Bayesian inferences, the software MrBayes 3. Bayesian posterior probabilities were computed under the same ML model by running two runs, four chains for 1,, MCMC generations using the program default priors on model parameters. Mixing and convergence to stationary distributions were evaluated by inspecting graphically the trace of the log likelihood or sampled parameter values against the generation numbers. A total of samples were screened: from four species of bats and 76 from five different species of NHPs For more details see Table 1.

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Individuals from the two groups of vertebrates were found infected by haemosporidian parasites and belonging to different genera Hepatocystis and Plasmodium. The infection rates are reported in Table 1. We found The infections rates varied from one species to another: E. Phylogenetic analyses revealed that haemosporidian parasites identified among the bats species clustered with several lineages of the Hepatocystis genus identified previously in other African areas Figure 2 and Supporting Information Figure S1.

Thus, all lineages obtained in this study infecting African bats belonged to a monophyletic group including all lineages recently described in several species of West Central and East African bats Figure 2. In our phylogeny, we can discriminate at least four distinct clades infecting the African bats clade 1—4 in Figure 3.

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As previously noted by Schaer et al. Phylogenetic analyses revealed that all sequences of parasites identified during our study and all different lineages from diverse geographical zones countries are mixed throughout the tree for African chiropterans. Thus, analyses indicate no strict geographic distribution within Hepatocystis isolates identified Figure 4. Beside the chiropteran hosts, five monkey species were included in the study: Mandrillus sphinx, Cercocebus torquatus, Cercophithecus cephus, Cercopithecus nictitans, and Cercophithecus solatus.

Three of them M. Regarding the Hepatocystis , they form a monophyletic group. This group comprises parasites from both African and Asian monkeys and is closely related to the Hepatocystis of Asian rodents and bats Figure 2 and Supporting Information Figure S1.

Within the tree, we observe no grouping of sequences by geographical areas, rather a mixed of sequences from different countries and no grouping by host species or genus Figure 4. No host switch was observed between bats and primates i.

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For the Plasmodium species infected monkeys, we identified two parasites: Plasmodium sp. These species were only found among mandrills. The discovery of new species in groups of parasites yet historically well studied because of their proximity to human malaria agents e. In this study, we investigated the diversity of extant malaria parasites circulating among bats and monkeys living in the same area, using molecular tools to analyze the parasite content of each host species collected and determine whether transfers of parasites may occur among host groups.

Molecular analyses revealed that the individuals of different groups of mammals primates and bats are infected by haemosporidian parasites belonging to at least two distinct genera: Plasmodium and Hepatocystis the focus of this study Figure 2 and Table 1. No parasites of Polychromophilus and Nycteria genera which were previously reported among bats Duval, et al. Regarding the genus Hepatocystis , they were found in bats and primates in our study. In the Gabonese bats, we observed a large diversity of lineages belonging to distinct clades.

Most of these lineages were genetically close or similar to lineages recently described in West and East African bats Schaer, et al.

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Such diversity of sequences may reflect the presence of different species of Hepatocystis or a single species with a large geographical distribution and a complex evolutionary history that led to the maintenance of divergent lineages within this species. They generally display a large geographical range but also a large host range, infecting different species and even different genera Figure 4.

Thus, certain lineages observed in Epomops franqueti from our study area in Gabon were also described from the same host species in Uganda or South Sudan or from a different bat genus e. This lack of clustering according to host and geography may be of multiple origins and future studies exploring this in more details should be done.

This will necessitate more systematic data on the diversity of Hepatocystis in the African bats all over the continent and in more host species. This variability could, for instance, be related to the historical expansion and contraction of bat distribution ranges and their secondary or primary contacts. The diversity of lineages observed in bats varies from one species to another. In our study, most lineages were for instance observed from only one species of bats, E. This could obviously be linked to sample size that was far higher for this species than for the others but it has been demonstrated for other pathogen groups like viruses that ecological factors such as the home range, the size of the animals, the roosting patterns, or the physiological characteristics may also explain these variations in the diversity of pathogen communities among species Maganga, et al.

Again more systematic data on the diversity of Hepatocystis in bats will be needed to explore such aspects in details. To our knowledge, our study is the first to report infections with haemosporidian parasites in Eidolon helvum.

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This result needs nevertheless to be considered with caution. PCR methods are known to be very sensitive and may, in certain cases, detect DNA of parasites in the blood of a wrong host after the parasites have died Vafa Homann et al.

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  6. The only valid proof of an infection would be the observation of the parasites in a blood smear or, at least, if only molecular analyses are available, confirmation of infection in multiple individuals of the same species. Regarding the Hepatocystis found in the African monkeys, they form a monophyletic clade with the Hepatocystis from the Asian monkeys, distinct from that of the African bats Putaporntip, et al. As in bats, the Hepatocystis from African monkeys form a diverse clade and it is yet unclear whether these Hepatocystis lineages are the image of multiple taxa or a single widely dispersed species Ayouba, et al.

    So it is likely that these different lineages correspond to different species. Our study and previous ones Ayouba, et al.