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It has been suggested that for these host species, the endobacteria had been present but subsequently were lost during further evolution [5]. As a corollary, the loss of Wolbachia led to a further hypothesis that the bacterial genes essential to the host fitness might have been successfully transferred and expressed into the host genome.

Although still subject of discussion [21] , some support for this hypothesis derives from studies on lateral gene transfer, as shown with several insect and filarial hosts [37]. Remnants of Wolbachia -like gene sequences have been identified in the filarial host genomes of the endobacteria-free L. The elimination of the bacteria might be an adaptive advantage because their antigens are inflammation inducers and contribute to filarial pathologies and immunological responses [39]. However, a recent study suggests that the bacteria might act as a decoy target for polynuclear neutrophils, preventing harmful effect of eosinophils on filariae [40].

Furthermore, a strain of Wolbachia that over-replicates in Aedes aegypti inhibits the development of Brugia malayi larvae and switches on a few important immune system genes [41] — [43]. In our previous study [5] , it appeared that the number of endobacteria-free filarial species had been underestimated and that several species without Wolbachia detected were parasitic in lizards and frogs.

Thus it was suggested that these filariae from reptiles and anurans diversified before the first bacterial invasion on the onchocercid lineage which had been tentatively dated to mya [3] , [6]. Indeed the origin of the Oswaldofilariinae, parasitic in crocodiles and squamates, was hypothetically dated from the late Jurassic, at the beginning of Gondwanian dislocation, mya [44] , [45].

However representatives of this subfamily had not yet been screened. This is not surprising since the recovery of filariae from connective tissues, their main localization, is not easy. Our investigation was resolutely rooted in biodiversity, expecting that the exploration of a broader range of filarial species would contribute to decipher the history of the Wolbachia -filaria symbiosis.

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The recovery of materials from wild animals from several biomes was undertaken. The first oswaldofilarine, the first splendidofilarine a parasite of birds , and several onchocercid genera from mammals have now been screened through PCR, as well as classic immunohistochemical staining and whole mount fluorescent analysis [21].

This study confirms that Wolbachia are not detected until now in the filarioid species parasitic in amphibians and reptiles. Several other features have emerged from this study: The screening for Wolbachia was performed on 35 species Table 1 ; specimens detailed in Figure S1 and Tables S1 , S2 , of which 28 are here examined for the first time and one recently by us [48]. These were the first representatives of Oswaldofilariinae, Piratuba scaffi from a lizard, and of Splendidofilariinae, Aproctella sp.

The histoimmunostainings are presented according to the following genera: Whole mount fluorescent analysis is presented on Figure 5. Transverse sections of gravid females of three species. Litomosoides sigmodontis , laboratory strain. Litomosoides yutajensis: Litomosa chiropterorum: Detail of uteri with microfilarioid eggs Wb—. Scale bars: Onchocerca dewittei japonica: Onchocerca eberhardi: Onchocerca skrjabini: Loxodontofilaria caprini: For staining, see Fig.

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Transverse sections of females of four species Wb—, at two magnifications. Cercopithifilaria crassa: Cercopithifilaria longa: Cercopithifilaria minuta: Cercopithifilaria shohoi: Cercopithifilaria japonica: Mansonella Cutifilaria perforata: Hypodermal lateral chord of a specimen of Onchocerca dewittei japonica without Wolbachia. Idem, a second specimen with a few Wolbachia arrow.

Epithelial somatic gonad with Wolbachia in Mansonella Cutifilaria perforata. In the same specimen, intestinal wall cells in with Wolbachia and above, lateral chord ch without Wolbachia. Wolbachia in epithelial somatic gonad of Cercopithifilaria japonica. Metaphase in a Cercopithifilaria crassa zygote, showing the absence of Wolbachia in the embryo. Polar bodies PB on the left. Names at the terminal nodes are those of the host species with the exceptions of the outgroup species. A—J are the supergroups names according to [1] , [3] , [8] , [14].

The tree has been obtained by Bayesian inference of phylogeny, using MrBayes 2. Species in bold are those newly found positive in the screening for Wolbachia performed in this study. Accession numbers are given for the sequences present in the databases. Dotted lines have been added to emphasize major discrepancies between filarial and Wolbachia trees. For further details, see paragraph 7 in the Results section. The presence or absence of Wolbachia were confirmed in species previously screened Tables 2 and S2. Species previously studied and as expected confirmed positive are: Onchocerca volvulus female worm from a human nodule, Cameroon , Dirofilaria repens from an Italian patient , Litomosoides sigmodontis 2 females recovered from a wild Sigmodon hispidus , Venezuela and a less commonly studied species, Dipetalonema gracile a female recovered from Cebus olivaceus , collected in Yutaje, Venezuela, like in [5] Tables 2 and S2.

The species confirmed negative were Loa loa two batches of infective larvae recovered from Chrysops vectors, Cameroon and the single Litomosoides species that did not harbour Wolbachia , L. Eight newly screened filarial species harbored Wolbachia but, for some of them, not all specimens are positive Tables 2 and S2. The species in which Wolbachia was detected are the following: Tetrapetalonema atelensis amazonae , from Cebus olivaceus Tables 2 and S2. However, in four infected filarial species, Wolbachia were not detected in each specimen.

These species were Lo. In Wolbachia positive filarial species lateral hypodermal chords might be infected, might not be infected, or weakly infected. The presence of the bacteria in the female germline was not constantly associated to their presence in the lateral hypodermal chords. Among the Wolbachia positive species in which the tissue distribution of Wolbachia was studied, the lateral chords harboured Wolbachia in L. On the contrary, Wolbachia were not observed in the lateral chords of Lo.

Rare or novel tissue Wolbachia localizations were observed in Mansonella Cutifilaria perforata and Cercopithifilaria japonica. Moreover, bacteria were constantly found in the cells of the intestine wall, in both sectioned and whole mount material Figures 3C—F; 5D. Wolbachia were also detected in the somatic gonad of C. Four types of Wolbachia were identified in the whole study. According to the Wolbachia supergroups, four types of Wolbachia were identified in this study: Wolbachia from representatives of supergroups A, B and E formed separated and supported clusters; Wolbachia from Ctenocephalides and Dipetalonema formed two separate lineages, that were recently assigned to supergroups I and J respectively [14].

Wolbachia was not detected in more than half of the newly screened species Tables 2 and S2. The Wolbachia negative species are: Inconsistency between phylogenies of Wolbachia and filarial hosts was evidenced in the cocladogenetic analysis. Thus the null hypothesis that the host and parasite have strictly co-speciated was rejected. The major inconsistency was due to the newly screened Cercopithifilaria japonica and Mansonella Cutifilaria perforata Figure 7.

Another point of inconsistency in the phylogenies of hosts and symbionts regards the positioning of L. Indeed, based on morphological adult and larval criteria, Litomosoides is closer to lymphatic filariae than to Mansonella [45]. As expected, the screening of a broader and more diversified set of species samples, by a combination of PCR and gene sequencing, immunohistological staining and whole mount fluorescent analysis, revealed new information about Wolbachia biology and evolution: In addition, the occurrence of Wolbachia in some members of a species and its absence in others raises questions about the evolution of its obligate requirement [51].


It is clear that tissues other than the female germline and the hypodermal lateral chords may be infected with Wolbachia. One of these infected tissues is the somatic gonad epithelial layer , once briefly reported previously [52]. Interestingly, they are both members of the supergroup F of Wolbachia. The real novelty is the Wolbachia tissue localization in the intestinal wall; it was only observed in M. These divergent localizations suggest a more complex and diversified relationship between the bacteria and filariae. They also raise the question of how and when Wolbachia bacteria reach the appropriate filarial host tissues.

It is likely that it is an early event, since it was shown in Brugia malayi an asymmetric distribution of bacteria in the egg followed by a preferential segregation in defined blastomeres [21]. The species screened in this study generally confirm the previously identified types of Wolbachia in the Onchocercidae Figure 6 ; see for instance [9].

The newly screened Loxodontofilaria is placed among the species of Onchocerca [53]. In addition, endosymbionts from this filaria belong to Wolbachia supergroup C Figure 6. There is a major phylogenetic congruence discrepancy between Wolbachia and their hosts and it occurs in the genus Cercopithifilaria and the supergroup F of Wolbachia. One African and seven Japanese species of Cercopithifilaria have no Wolbachia, while one species in Japan is Wolbachia positive.

The filarial hosts belong to a well-supported genus, Cercopithifilaria as evidenced by adult morphology [45] , [54] , larval morphology [55] , 12S rDNA gene sequences [53] , [56] , the transmission by hard ticks and the skin-dwelling microfilariae [45]. A parsimonious interpretation of the Wolbachia screening is that a single acquisition event took place in C. This hypothesis is supported by the co-cladogenetic analyses Figure 7. The supergroup F is intriguing as it is presently the only Wolbachia type infecting both insects and onchocercid nematodes [7].

The Wolbachia supergroup F contains the species of Mansonella studied so far: Esslingeria perstans [13] , [15] , and in this study, M. Tetrapetalonema atelensis amazonae, M.

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  8. Cutifilaria perforata. Cercopithifilaria japonica in supergroup F suggests a transversal transmission event, likely recent due to limited occurrence among the species of this genus. This Mansonella species, M. The bacterial host switching might have occurred between the two filarial parasites of the bear, perhaps via an oral infection route. Indeed filariae, despite their apparent small mouth, can ingest particles from their environment, such as red blood cells [59] and larger bodies, such as microfilariae released in the coelomic cavities of the filarial host [60]. The filarial species in which Wolbachia were not detected appeared more numerous than it was thought, based upon previous observations.

    In [34] , the percentage of negative species was It has to be emphasized that the negative results were not due to DNA degradations or bad extractions because all of the Wolbachia PCR negative samples gave positive amplifications using filarial nematode specific primers. However, we must take into account the fact that, in a few species, Wolbachia were not detected in all the specimens.

    This can partly be explained in species which do not harbour Wolbachia in the lateral chords, or at very low density, as Onchocerca dewittei japonica Figures 5A,B and Loxodontofilaria caprini Figures 2H,I. Thus, in M. Esslingeria perstans from humans. It is interesting to note that in both of these cases, the Wolbachia supergroup is F. Further, research performed on deeply studied filariae, such as Brugia malayi , have also shown that the amount of Wolbachia carried by a worm may vary greatly over time and be stage-dependent [21].

    This dynamic probably has little impact when considering developing larval stages, because these are transient and the chance of recovering them in the wild is extremely low. More interesting is the observation that the female worms recovered in the wild are not all fully gravid in some species Figure 2F. This was not the case of filariae from frogs, lizards, bats, birds, but was the case from some parasites of Japanese ungulates.

    It is worth to note that we paid great attention to the part of worm sampled to ensure that the germline was screened in all species. It is clear that there is a need to increase the number of screened specimens for more solid results. However, in several cases, the global results are impressive and the distribution of the Wolbachia negative species does not appear random.

    As a matter of fact, at present, the species parasitic in frogs and lizards are negative for Wolbachia if they are Waltonellinae three species of Ochoterenella , Oswaldofilariinae one species of Piratuba , or Dirofilariinae two species of Foleyella. It was also shown here that the first screened species parasitic in birds, an Aproctella in the Splendidofilariinae, was Wolbachia negative 7 females screened with PCR, from two passeriform species, totalising 7 specimens hosts.

    Wolbachia were not detected also in several species of Onchocercinae from mammals. The first is Litomosa chiropterorum, from an African bat 8 specimens screened from 8 Miniopterus schreibersi , an unexpected observation since the single Litomosa species screened previously, Li. The second species is Monanema martini 8 females screened from 8 murid specimens.

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    The absence of the bacteria is an important feature, considering that this filaria was used, in the past, as a model of onchocerciasis, a true limit due to the important role played by the bacteria in the pathology of the disease [39] , [61] — [63]. The absence of Wolbachia could explain partly the weak ocular pathology induced in the murid host [49]. Severe or mild ocular onchocerciases have been related to a strain-dependent variation of density of Wolbachia per filarial genome [64].

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    Two hypotheses might be taken into account to explain the absence of Wolbachia. In the first case, Wolbachia could be considered to be present and then were subsequently lost [38]. This could have occurred before the mutually dependent symbiosis between symbiont and host developed. The second hypothesis is that the bacteria were not yet acquired within that filarial group. The first hypothesis seems to possibly explain some of the observed cases, such as Litomosoides yutajensis 5 samples: In the cases of Acantocheilonema viteae and Onchocerca flexuosa [38] , the absence of Wolbachia appears to be a secondary loss, because some genes of the bacteria were incorporated in the filarial genomes.

    However, it is not clear whether this loss occurred before symbiosis was established or not. This is of interest because if loss was after symbiotic establishment, perhaps some of the genes incorporated into the host genome were those required by the host now provided by Wolbachia , but the extent of this event is still to be understood. Estimation of dates of divergence has been proposed for some groups of nematodes based on molecular phylogenetic analyses [66]. In filariae, data from morphology, biology, geographic distribution, host range and palaeontology led to the proposal that the Oswaldofilariinae emerged during the late Jurassic, at the beginning of the Gondwanian break up, mya [44] , [45].

    This is before the hypothesized ancestral acquisition of a Wolbachia by an onchocercid [3] ; it follows that the absence of Wolbachia in Oswaldofilariinae could be primitive. Foleyella is another parasite of saurians, which appears to have no Wolbachia ; it is presently placed in the Dirofilariinae, which includes the Wolbachia positive Dirofilaria , but this systematic position needs to be revised because the characters of the infective larvae are very distinct [55]. A solid phylogeny of Onchocercidae linking traditional and molecular data is needed and warrants further investigations.

    The subfamily Setariinae, parasitic in ungulates and in which Wolbachia were not detected, as first shown by [67] , also deserves a comment. Based on larval morphology, it has been hypothesized that it evolved separately from the other onchocercids and derived from a group of spirurid Habronematinae [65]. Until now no spirurids have been found infected with Wolbachia [5] , [34].

    Did the bacterial infection never occur, or was the useful part of Wolbachia genome incorporated in the host genome and subsequently the bacteria eliminated, to reply to some adverse constraint? Further genomic analyses will solve this question concerning the absence of Wolbachia in ancestral filariae. At present, the features observed on tissue or specimen distribution, and a very probable recent lateral transfer, suggest complex evolutionary dynamics of interactions between the symbionts, their host filariae and the nematode hosts.

    In the sampling studied, which will be further enlarged, it may become possible to use some defined species to decipher these questions. Specimens of filariae were recovered during dissections of the vertebrate hosts captured in the wild from different geographic areas [50] , [54] , [68] — [72]. All experiments, procedures and ethical issues were conformed to the competent national ethical bodies: Japanese serows, sika deer, bears, and wild boars were killed by hunters who have an individual permit to kill wild animals in accordance with the conservation and control policies of the Ministry of the Environment of Japan.

    Italian samples were collected by veterinarians and physicians and no permits were necessary. African rodents and agama do not belong to protected species and were obtained from local hunters. African mountain reedbuck, plain zebra, gemsbok and porcupine were also obtained from local hunters. Bats from South African bats have been collected for previous studies [48] in which no permits were necessary. Many of the filariae from large mammals were extracted from the subconnective tissue, dermis, or tendons of limbs.

    Afterwards the parts of the animals were dissected to collect living filarioids for Wolbachia study. From frogs, lizards, birds, bats and rodents filariae were generally recovered immediately after host death. Samples used for positive and negative PCR controls were laboratory strains. Some species are not yet named, but all are under study, and morphological analysis and sequencing of coxI gene in this study and in [53] showed that they represent distinct molecular entities. Co-infection of a host specimen by several congeneric or non-congeneric filarial species was rather frequent in large mammals, but in a few cases, the same filarial species was recovered from two host species.

    The supraspecific levels of taxonomy followed the systematic works of [46] , [73] , [74] and more recent studies for some taxa: Abbreviations used in the text are: In many cases, filarial specimens were cut into anterior a , median m and posterior p parts, which were fixed for the different analysis approaches. Since Wolbachia are transmitted by female filariae, almost all of the studies were based on female worms; male specimens were rarely examined, and we analysed this sex alone in a single case, Di. The thermal profile used was: PCR conditions for this amplification were as described in [79].

    MgCl 2 concentrations at 2. DNA preparations from filarial species harbouring Wolbachia D. DNA preparations from a filarial species not harbouring Wolbachia A. The thermal profiles we used were: In all cases, in order to ascertain the DNA conditions before Wolbachia screenings and to confirm morphological identification, coxI amplification was performed as described in [12]. A list of the sequences including accession numbers is available in Table S2. Immunohistochemical staining was performed according to [24] , [48].

    Negative controls were carried out by omitting the primary antibody. Fixed female worms were divided in three main parts, posterior p , median c and anterior a in order to observe the different regions of the genital tract. Transverse sections were made at different levels of each part, and few of them were stained with hemalun-eosin for anatomical identification.

    Lateral cuticular internal crests were identified to orient the worm section; hypodermal chords extend above and on the side of the crests. The filarial species used for histology are opisthodelphic and the initial part of the ovaries is in the posterior part of the worm; the distal region of the ovary is composed of a cytoplasmic axis, the rachis [82] , [83] , and an outer cytoplasmic layer with the nuclei of oogonia and oocytes in the text we will refer to these states as oocytes, and germline to describe the whole production of the gonad.

    An epithelial layer and an outer muscle layer surround the gonad, both referred to as somatic gonad in the text [25]. Uteri occupy almost the whole body and are found in median and anterior part of worms. The different uterine contents are ovulae, spermatozoa, divided eggs and microfilariae. Eggs were identified as aborted by hemalun-eosin staining when divided eggs were eosinophilic and nuclei not discernible. The laboratory strain of Litomosoides sigmodontis, which has been shown in several studies to harbour Wolbachia [84] — [86] was used as a positive reference for Wolbachia immunostaining.

    Tissues were mounted in Vectashield Vector Laboratories [25]. The species analyzed were C. The appropriate model of sequence evolution for ML and BI was estimated via likelihood ratio test using Modeltest 3. Two independent runs were performed, each using 1 million steps with four chains sampling every steps. Four Wolbachia alignments relative to the genes 16S rDNA, ftsZ , groEL and dnaA were screened for presence of recombination events by using a set of nonparametric detection programs: The first six programs search for putative recombination breakpoints in a set of aligned DNA sequences and are implemented in RDP3 software package, whilst LARD checks signals detected by other methods [92].

    Sequences were auto-masked for optimal recombination detection. General recombination settings were as follows: Method specific options were as follow: MaxChi and Chimaera were run with a variable window size; Bootscan and SisScan were forced for exploratory screening. Successive phases of refining analysis and manual tests were performed where needed. The first tested whether there was a greater than random correspondence between reconstructed nodes for host and symbiont. This was performed in Component R. The second test examined the null hypothesis that the endosymbionts have undergone cocladogenesis with their hosts.

    ML trees for each dataset were first estimated using the successive approximation method [89]. Then, the scores for each of these ML trees based on the host dataset were compared using the [98] test. This was repeated using the parasite dataset. A single uninfected filarial T. Position of the genera screened in the present study indicated on a schematic representation of a key of the onchocercid subfamilies, based on morphological characters following [46].

    Total number of genera per subfamily listed. Results of Wolbachia screening based on PCR, immunostaining assays and whole mount fluorescent analysis in 35 filarial nematodes. Taxa are presented in alphabetical order. Wolbachia distribution in the tissues of 13 onchocercid species. We are also grateful to Prof. Takaoka and Dr. Fukuda, Oita University, Japan for their contribution in providing animal hosts and Mr. Competing Interests: The authors have declared that no competing interests exist.

    The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Europe PMC requires Javascript to function effectively. Recent Activity. The snippet could not be located in the article text. This may be because the snippet appears in a figure legend, contains special characters or spans different sections of the article.

    PLoS One. Published online Jun Sara G. Received Feb 22; Accepted May Copyright Ferri et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are properly credited. This article has been cited by other articles in PMC. Abstract Background Wolbachia are intriguing symbiotic endobacteria with a peculiar host range that includes arthropods and a single nematode family, the Onchocercidae encompassing agents of filariases.

    Introduction The alpha proteobacteria Wolbachia Rickettsiales are present in two distinct zoological groups: Results The screening for Wolbachia was performed on 35 species Table 1 ; specimens detailed in Figure S1 and Tables S1 , S2 , of which 28 are here examined for the first time and one recently by us [48]. Open in a separate window. The 35 species of filariae included in this study, their hosts and collection place. Species, genera, subgenera and subfamilies screened for the first time are in bold characters. Figure 1. Wolbachia immunostaining in the genera Litomosoides L. Figure 2.

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