RESUMO
Lutzomyia longipalpis s.l. is a complex of sibling species and is the principal vector of American visceral leishmaniasis. The present review summarises the diversity of efforts that have been undertaken to elucidate the number of unnamed species in this species complex and the phylogenetic relationships among them. A wide variety of evidence, including chemical, behavioral and molecular traits, suggests very recent speciation events and complex population structure in this group. Although significant advances have been achieved to date, differential vector capacity and the correlation between structure of parasite and vector populations have yet to be elucidated. Furthermore, increased knowledge about recent epidemiological changes, such as urbanisation, is essential for pursuing effective strategies for sandfly control in the New World.
Assuntos
Animais , Psychodidae/classificação , Psychodidae/genética , Especificidade da Espécie , Genes de Insetos , Biodiversidade , Insetos Vetores/classificação , Insetos Vetores/genética , Leishmaniose Visceral/transmissão , Filogenia , Polimorfismo Genético , Polimorfismo de Fragmento de Restrição , BrasilRESUMO
Lutzomyia longipalpis s.l. is a complex of sibling species and is the principal vector of American visceral leishmaniasis. The present review summarises the diversity of efforts that have been undertaken to elucidate the number of unnamed species in this species complex and the phylogenetic relationships among them. A wide variety of evidence, including chemical, behavioral and molecular traits, suggests very recent speciation events and complex population structure in this group. Although significant advances have been achieved to date, differential vector capacity and the correlation between structure of parasite and vector populations have yet to be elucidated. Furthermore, increased knowledge about recent epidemiological changes, such as urbanisation, is essential for pursuing effective strategies for sandfly control in the New World.
Assuntos
Biodiversidade , Insetos Vetores/classificação , Insetos Vetores/genética , Psychodidae/classificação , Psychodidae/genética , Animais , Brasil , Genes de Insetos , Leishmaniose Visceral/transmissão , Filogenia , Polimorfismo de Fragmento de Restrição , Especificidade da EspécieRESUMO
BACKGROUND: Brazilian populations of Lutzomyia longipalpis may constitute a complex of cryptic species, and this report investigates the distribution and number of potential sibling species. One of the main differences observed among Brazilian populations is the type of acoustic signal produced by males during copulation. These copulation song differences seem to be evolving faster than neutral molecular markers and have been suggested to contribute to insemination failure observed in crosses between these sibling species. In previous studies, two main types of copulation songs were found, burst-type and pulse-type. The latter type can, in turn, be further subdivided into five different patterns. METHODS: We recorded male song from 13 new populations of the L. longipalpis complex from Brazil and compared the songs with 12 already available. RESULTS: Out of these 25 populations, 16 produce burst-type and 9 produce pulse-type songs. We performed a principal component analysis in these two main groups separately and an additional discriminant analysis in the pulse-type group. The pulse-type populations showed a clear separation between the five known patterns with a high correspondence of individuals to their correct group, confirming the differentiation between them. The distinctiveness of the burst-type subgroups was much lower than that observed among the pulse-type groups and no clear population structure was observed. This suggests that the burst-type populations represent a single species. CONCLUSION: Overall, our results are consistent with the existence in Brazil of at least six species of the L. longipalpis complex, one with a wide distribution comprising all the populations with burst-type songs, and five more closely related allopatric siblings with different pulse-type song patterns and more restricted distribution ranges.
Assuntos
Psychodidae/fisiologia , Animais , Brasil , Copulação , Feminino , Masculino , Psychodidae/classificação , Vocalização AnimalRESUMO
BACKGROUND: Lutzomyia longipalpis, the main vector of visceral leishmaniasis in Latin America, is a complex of sibling species. In Brazil, a number of very closely related sibling species have been revealed by the analyses of copulation songs, sex pheromones and molecular markers. However, the level of divergence and gene flow between the sibling species remains unclear. Brazilian populations of this vector can be divided in two main groups: one producing Burst-type songs and the Cembrene-1 pheromone and a second more diverse group producing various Pulse song subtypes and different pheromones. METHODOLOGY/PRINCIPAL FINDINGS: We analyzed 21 nuclear loci in two pairs of Brazilian populations: two sympatric populations from the Sobral locality (1S and 2S) in northeastern Brazil and two allopatric populations from the Lapinha and Pancas localities in southeastern Brazil. Pancas and Sobral 2S are populations of the Burst/Cembrene-1 species while Lapinha and Sobral 1S are two putative incipient species producing the same pheromone and similar Pulse song subtypes. The multilocus analysis strongly suggests the occurrence of gene flow during the divergence between the sibling species, with different levels of introgression between loci. Moreover, this differential introgression is asymmetrical, with estimated gene flow being higher in the direction of the Burst/Cembrene-1 species. CONCLUSIONS/SIGNIFICANCE: The results indicate that introgressive hybridization has been a crucial phenomenon in shaping the genome of the L. longipalpis complex. This has possible epidemiological implications and is particularly interesting considering the potential for increased introgression caused by man-made environmental changes and the current trend of leishmaniasis urbanization in Brazil.
Assuntos
Variação Genética , Psychodidae/classificação , Psychodidae/genética , Animais , Brasil , Quimera , Genoma de Inseto , Masculino , Dados de Sequência Molecular , Tipagem de Sequências Multilocus , IrmãosRESUMO
The sand fly Lutzomyia longipalpis (Diptera: Psychodidae: Phlebotominae), the most important vector of American visceral leishmaniasis, is widely distributed in Latin America. There is currently a consensus that it represents a species complex, however, the number and distribution of the different siblings is still uncertain. Previous analyses have indicated that Brazilian populations of this vector can be divided into two main groups according to the type of courtship song (Burst vs. Pulse) males produce during copulation. Nevertheless, no diagnostic differences have been observed between these two groups with most molecular markers used to date. We analyzed the molecular divergence in a fragment of the paralytic (para) gene, a locus involved in the control of courtship songs in Drosophila, among a number of Lu. longipalpis populations from Brazil producing Burst and Pulse-type songs. Our results revealed a very high level of divergence and fixed differences between populations producing the two types of songs. We also compared Lu. longipalpis with a very closely related species, Lutzomyia cruzi, which produces Burst-type songs. The results indicated a higher number of fixed differences between Lu. cruzi and the Pulse-type populations of Lu. longipalpis than with those producing Burst-type songs. The data confirmed our previous assumptions that the presence of different sibling species of the Lu. longipalpis complex in Brazil can be divided into two main groups, one representing a single species and a second more heterogeneous group that probably represents a number of incipient species. We hypothesize that para might be one of the genes directly involved in the control of the courtship song differences between these two groups or that it is linked to other loci associated with reproductive isolation of the Brazilian species.
Assuntos
Corte , Genes de Insetos/genética , Proteínas de Insetos/genética , Filogenia , Psychodidae/genética , Canto , Animais , Brasil , Evolução Molecular , Feminino , Genética Populacional , Geografia , Haplótipos/genética , Masculino , Polimorfismo GenéticoRESUMO
BACKGROUND: It has been suggested that genes involved in the reproductive biology of insect disease vectors are potential targets for future alternative methods of control. Little is known about the molecular biology of reproduction in phlebotomine sand flies and there is no information available concerning genes that are expressed in male reproductive organs of Lutzomyia longipalpis, the main vector of American visceral leishmaniasis and a species complex. METHODS/PRINCIPAL FINDINGS: We generated 2678 high quality ESTs ("Expressed Sequence Tags") of L. longipalpis male reproductive organs that were grouped in 1391 non-redundant sequences (1136 singlets and 255 clusters). BLAST analysis revealed that only 57% of these sequences share similarity with a L. longipalpis female EST database. Although no more than 36% of the non-redundant sequences showed similarity to protein sequences deposited in databases, more than half of them presented the best-match hits with mosquito genes. Gene ontology analysis identified subsets of genes involved in biological processes such as protein biosynthesis and DNA replication, which are probably associated with spermatogenesis. A number of non-redundant sequences were also identified as putative male reproductive gland proteins (mRGPs), also known as male accessory gland protein genes (Acps). CONCLUSIONS: The transcriptome analysis of L. longipalpis male reproductive organs is one step further in the study of the molecular basis of the reproductive biology of this important species complex. It has allowed the identification of genes potentially involved in spermatogenesis as well as putative mRGPs sequences, which have been studied in many insect species because of their effects on female post-mating behavior and physiology and their potential role in sexual selection and speciation. These data open a number of new avenues for further research in the molecular and evolutionary reproductive biology of sand flies.
Assuntos
Proteínas de Insetos/genética , Insetos Vetores/genética , Psychodidae/genética , Testículo/fisiologia , Transcriptoma/genética , Sequência de Aminoácidos , Animais , Biomarcadores/metabolismo , Biologia Computacional , Etiquetas de Sequências Expressas , Feminino , Masculino , Dados de Sequência Molecular , Análise de Sequência com Séries de Oligonucleotídeos , Filogenia , RNA Mensageiro/genética , Reação em Cadeia da Polimerase em Tempo Real , Reação em Cadeia da Polimerase Via Transcriptase Reversa , Homologia de Sequência de AminoácidosRESUMO
BACKGROUND: Acoustic signals are part of the courtship of many insects and they often act as species-specific signals that are important in the reproductive isolation of closely related species. Here we report the courtship songs of the sand fly Lutzomyia (Nyssomyia) intermedia, one of the main vectors of cutaneous leishmaniasis in Brazil. FINDINGS: Recordings were performed using insects from three localities from Eastern Brazil: Posse and Jacarepaguá in Rio de Janeiro State and Corte de Pedra in Bahia State. The three areas have remnants of the Brazilian Atlantic forest, they are endemic for cutaneous leishmaniasis and L. intermedia is the predominant sand fly species. We observed that during courtship L. intermedia males from all populations produced pulse songs consisting of short trains. No significant differences in song parameters were observed between the males of the three localities. CONCLUSIONS: L. intermedia males produce acoustic signals as reported for some other sand flies such as the sibling species of the Lutzomyia longipalpis complex. The lack of differences between the males from the three localities is consistent with previous molecular studies of the period gene carried out in the same populations, reinforcing the idea that L. intermedia is not a species complex in the studied areas and that the three populations are likely to have similar vectorial capacities.
Assuntos
Psychodidae/fisiologia , Vocalização Animal , Animais , Brasil , Feminino , MasculinoRESUMO
Lutzomyia migonei is a vector of leishmaniasis with a wide distribution in South America, which could favour population differentiation and speciation. Cryptic species of the Lutzomyia longipalpis complex, the widely distributed sand fly vector of visceral leishmaniasis in Latin America, have previously been shown to display distinct copulation songs. We found that Lu. migonei males also produce a song during copulation. This "lovesong" presents short trains (6-8 pulses) with an inter-pulse interval around 26 ms and is potentially involved in cryptic female choice and insemination success.
Assuntos
Animais , Feminino , Masculino , Corte , Copulação/fisiologia , Insetos Vetores/fisiologia , Psychodidae/fisiologia , Vocalização Animal/fisiologia , Insetos Vetores , Leishmaniose Visceral/transmissão , Psychodidae , Fatores de TempoRESUMO
The sand fly Lutzomyia cruzi (Mangabeira, 1938) is implicated as a vector of American visceral leishmaniasis (AVL) in some areas of Brazil. Lutzomyia cruzi is closely related to Lutzomyia longipalpis s.l. (Lutz and Neiva, 1912) the main Latin American vector of AVL and a species complex. Although females of the two species are identical, the males can be distinguished by differences in the genitalia. Nevertheless, pheromone analysis shows that Lu. cruzi males produce 9-methyl-germacrene-B, which has also been found in a number of Latin American populations of Lu. longipalpis s.l. In addition, analysis of microsatellite loci shows that the level of divergence between Lu. cruzi and Lu. longipalpis s.l. is similar to that observed among the Lu. longipalpis s.l. sibling species. Here we present the lovesongs of Lu. cruzi males which are similar to the Burst-type songs produced by one of the Lu. longipalpis s.l. sibling species. We also present data on the molecular polymorphisms of the period gene of Lu. cruzi that indicate this species as another sibling within the Lu. longipalpis complex. The results highlight the importance of an integrative approach to understand the patterns of genetic and phenotypic divergence among very closely related vector species.
Assuntos
Especiação Genética , Proteínas Circadianas Period/genética , Polimorfismo Genético , Psychodidae/genética , Vocalização Animal/fisiologia , Animais , Brasil , Corte , Feminino , Genes de Insetos/genética , Masculino , Filogenia , Polimorfismo Genético/fisiologia , Psychodidae/classificaçãoRESUMO
Lutzomyia migonei is a vector of leishmaniasis with a wide distribution in South America, which could favour population differentiation and speciation. Cryptic species of the Lutzomyia longipalpis complex, the widely distributed sand fly vector of visceral leishmaniasis in Latin America, have previously been shown to display distinct copulation songs. We found that Lu. migonei males also produce a song during copulation. This "lovesong" presents short trains (6-8 pulses) with an inter-pulse interval around 26 ms and is potentially involved in cryptic female choice and insemination success.
Assuntos
Copulação/fisiologia , Corte , Insetos Vetores/fisiologia , Psychodidae/fisiologia , Vocalização Animal/fisiologia , Animais , Feminino , Insetos Vetores/classificação , Leishmaniose Visceral/transmissão , Masculino , Psychodidae/classificação , Fatores de TempoRESUMO
This no-choice, laboratory study focuses on the feeding of dried, ground, homogeneous powdered, unprocessed fruit and leaves of Azadirachta indica and Melia azedarach to Lutzomyia longipalpis larvae to determine the effects on their mortality and metamorphosis. A. indica and M. azedarach fruit and leaves significantly increased larval mortality in comparison to larvae fed the untreated, standard diet. A. indica fruit and leaves blocked the molting of the larvae to the fourth instar, resulting in them remaining as third instars until the end of the experiment. M. azedarach fruit also blocked the molting of larvae, which remained permanently in the fourth instar. Feeding M. azedarach leaves resulted in greater molt inhibition. All insects in this group stopped their development as second-instar larvae. No antifeedant effect was detected for any experimental treatment. The results indicate that nontoxic, unprocessed materials obtained from A. indica and M. azedarach are potent development inhibitors of L. longipalpis larvae.
Assuntos
Azadirachta , Melia azedarach , Metamorfose Biológica/efeitos dos fármacos , Extratos Vegetais/farmacologia , Psychodidae/efeitos dos fármacos , Animais , Frutas , Larva/efeitos dos fármacos , Larva/crescimento & desenvolvimento , Folhas de Planta , Psychodidae/crescimento & desenvolvimentoRESUMO
BACKGROUND: Lutzomyia longipalpis is the primary vector of American visceral leishmaniasis. There is strong evidence that L. longipalpis is a species complex, but until recently the existence of sibling species among Brazilian populations was considered a controversial issue. In addition, there is still no consensus regarding the number of species occurring in this complex. METHODOLOGY/PRINCIPAL FINDINGS: Using period, a gene that controls circadian rhythms and affects interpulse interval periodicity of the male courtship songs in Drosophila melanogaster and close relatives, we analyzed the molecular polymorphism in a number of L. longipalpis samples from different regions in Brazil and compared the results with our previously published data using the same marker. We also studied the male copulation songs and pheromones from some of these populations. The results obtained so far suggest the existence of two main groups of populations in Brazil, one group representing a single species with males producing Burst-type copulation songs and cembrene-1 pheromones; and a second group that is more heterogeneous and probably represents a number of incipient species producing different combinations of Pulse-type songs and pheromones. CONCLUSIONS/SIGNIFICANCE: Our results reveal a high level of complexity in the divergence and gene-flow among Brazilian populations of the L. longipalpis species complex. This raises important questions concerning the epidemiological consequences of this incipient speciation process.
Assuntos
Insetos Vetores/fisiologia , Leishmaniose Visceral/epidemiologia , Psychodidae/fisiologia , Animais , Brasil/epidemiologia , Ritmo Circadiano/genética , Genes de Insetos , Variação Genética , Insetos Vetores/genética , Insetos Vetores/metabolismo , Leishmaniose Visceral/transmissão , Masculino , Psychodidae/genética , Psychodidae/metabolismo , Atrativos Sexuais/análise , Comportamento Sexual AnimalRESUMO
Lutzomyia longipalpis s.l., the main vector of Leishmania chagasi in Latin America, is a species complex although the exact number of siblings is yet unknown. In Brazil, the siblings differ in male copulatory courtship songs and pheromones that most certainly act as pre-zygotic reproductive barriers. Here we analysed the reproductive isolation between three allopatric and two sympatric populations of Lu. longipalpis s.l. from Brazil. The results indicate a strong copulatory and pre-mating isolation between the three allopatric populations. In addition, the results also indicate a stronger pre-mating isolation between the two sympatric siblings than between the three allopatric ones, suggesting a role for reinforcement in the speciation of the Lu. longipalpis s.l. complex.
Assuntos
Copulação/fisiologia , Psychodidae/fisiologia , Animais , Brasil , Feminino , Masculino , Feromônios/fisiologia , Reprodução/fisiologia , Vocalização Animal/fisiologiaRESUMO
Lutzomyia longipalpis s.l., the main vector of Leishmania chagasi in Latin America, is a species complex although the exact number of siblings is yet unknown. In Brazil, the siblings differ in male copulatory courtship songs and pheromones that most certainly act as pre-zygotic reproductive barriers. Here we analysed the reproductive isolation between three allopatric and two sympatric populations of Lu. longipalpis s.l. from Brazil. The results indicate a strong copulatory and pre-mating isolation between the three allopatric populations. In addition, the results also indicate a stronger pre-mating isolation between the two sympatric siblings than between the three allopatric ones, suggesting a role for reinforcement in the speciation of the Lu. longipalpis s.l. complex.
Assuntos
Animais , Feminino , Masculino , Copulação/fisiologia , Psychodidae/fisiologia , Brasil , Feromônios/fisiologia , Reprodução/fisiologia , Vocalização Animal/fisiologiaRESUMO
Lutzomyia longipalpis is the main vector of Leishmania infantum chagasi, the causative agent of American visceral leishmaniasis (AVL). Although there is strong evidence that Lu. longipalpis is a species complex, not all data concerning populations from Brazil support this hypothesis. The issue is still somewhat controversial for this large part of Lu. longipalpis distribution range even though that it is the Latin American region contributing to most of the cases of AVL. In this mini-review we consider in detail the current data for the Brazilian populations and conclude that Lu. longipalpis is a complex of incipient vector species with a complexity similar to Anopheles gambiae s.s. in Africa.
Assuntos
Insetos Vetores/classificação , Psychodidae/classificação , Animais , Brasil , Insetos Vetores/genética , Leishmaniose Visceral/transmissão , Psychodidae/genéticaRESUMO
Lutzomyia longipalpis is the main vector of Leishmania infantum chagasi, the causative agent of American visceral leishmaniasis (AVL). Although there is strong evidence that Lu. longipalpis is a species complex, not all data concerning populations from Brazil support this hypothesis. The issue is still somewhat controversial for this large part of Lu. longipalpis distribution range even though that it is the Latin American region contributing to most of the cases of AVL. In this mini-review we consider in detail the current data for the Brazilian populations and conclude that Lu. longipalpis is a complex of incipient vector species with a complexity similar to Anopheles gambiae s.s. in Africa.
Assuntos
Animais , Insetos Vetores/classificação , Psychodidae/classificação , Brasil , Insetos Vetores/genética , Leishmaniose Visceral/transmissão , Psychodidae/genéticaRESUMO
BACKGROUND: Lutzomyia intermedia and Lutzomyia whitmani (Diptera: Psychodidae) are important and very closely related vector species of cutaneous leishmaniasis in Brazil, which are distinguishable by a few morphological differences. There is evidence of mitochondrial introgression between the two species but it is not clear whether gene flow also occurs in nuclear genes. RESULTS: We analyzed the molecular variation within the clock gene period (per) of these two species in five different localities in Eastern Brazil. AMOVA and Fst estimates showed no evidence for geographical differentiation within species. On the other hand, the values were highly significant for both analyses between species. The two species show no fixed differences and a higher number of shared polymorphisms compared to exclusive mutations. In addition, some haplotypes that are "typical" of one species were found in some individuals of the other species suggesting either the persistence of old polymorphisms or the occurrence of introgression. Two tests of gene flow, one based on linkage disequilibrium and a MCMC analysis based on coalescence, suggest that the two species might be exchanging alleles at the per locus. CONCLUSION: Introgression might be occurring between L. intermedia and L. whitmani in period, a gene controlling behavioral rhythms in Drosophila. This result raises the question of whether similar phenomena are occurring at other loci controlling important aspects of behavior and vectorial capacity.
Assuntos
Proteínas Nucleares/genética , Polimorfismo Genético , Psychodidae/genética , Animais , Sequência de Bases , Brasil , Proteínas de Drosophila , Dados de Sequência Molecular , Proteínas Circadianas Period , Filogenia , Especificidade da EspécieRESUMO
Despite the importance of circadian rhythms in vector-borne disease transmission, very little is known about its molecular control in hematophagous insect vectors. In Drosophila melanogaster, a negative feedback loop of gene expression has been shown to contribute to the clock mechanism. Here, we describe some features of the circadian clock of the sandfly Lutzomyia longipalpis, a vector of visceral leishmaniasis. Compared to D. melanogaster, sandfly period and timeless, two negative elements of the feedback loop, show similar peaks of mRNA abundance. On the other hand, the expression of Clock (a positive transcription factor) differs between the two species, raising the possibility that the different phases of Clock expression could be associated with the observed differences in circadian activity rhythms. In addition, we show a reduction in locomotor activity after a blood meal, which is correlated with downregulation of period and timeless expression levels. Our results suggest that the circadian pacemaker and its control over the activity rhythms in this hematophagous insect are modulated by blood intake.
Assuntos
Relógios Biológicos/genética , Ritmo Circadiano/genética , Regulação para Baixo/fisiologia , Proteínas de Insetos/genética , Insetos Vetores/fisiologia , Psychodidae/fisiologia , Animais , Drosophila melanogaster , Ingestão de Alimentos/fisiologia , Feminino , Leishmaniose Visceral/transmissão , MasculinoRESUMO
Lutzomyia longipalpis, a sibling complex, is the main vector of Leishmania chagasi/infantum. Discriminating between siblings is important as they may differ in vectorial capacity. Lutzomyia longipalpis populations display distinct male sex pheromone chemotypes. We investigated the phylogeographic pattern of variation at microsatellite loci from 11 populations from Brazil and Venezuela related to their male pheromone. Temporal genetic differentiation was mostly not significant at the same site. Spatial genetic differentiation was, however, strong, although there was only a weak relationship between genetic differentiation and the geographic distance separating the samples (r2 < 0.10); geographic separation explained a much greater (54-97%) percentage of the genetic differences among populations when samples with the same pheromone type were analyzed separately. A cluster analysis showed five groups: Lu. cruzi (Brazil) and Lu. pseudolongipalpis (Venezuela) as separate species, two (mostly 9-methyl-germacrene-B) Venezuelan and Brazilian groups, and a very distinct cluster of Brazilian cembrene populations.
Assuntos
Filogenia , Psychodidae/classificação , Psychodidae/genética , Atrativos Sexuais , Animais , Brasil , Análise por Conglomerados , Feminino , Variação Genética , Genética Populacional , Genótipo , Geografia , Masculino , Repetições de Microssatélites , Psychodidae/fisiologia , Atrativos Sexuais/química , Especificidade da Espécie , VenezuelaRESUMO
Lutzomyia intermedia (Lutz & Neiva, 1912) and L. whitmani (Antunes & Coutinho, 1939) (Diptera: Psychodidae: Phlebotominae), two important vectors of American cutaneous leishmaniasis in Brazil, occur in sympatry in the locality of Posse county, Petropolis municipality, state of Rio de Janeiro, Brazil. We investigated the influence of the lunar cycle on the frequency of specimens of the two species caught while attempting to bite the collectors and in CDC light traps. Analysis of the numbers of sand flies captured in different lunar phases for two consecutive years in the peridomestic site and forest shows that there is a significant positive correlation between moonlight intensity and the numbers of L. intermedia and L. whitmani females collected while blood-feeding, whereas the opposite was observed for the CDC traps.