RESUMEN
Holometaboly is a key evolutionary innovation that has facilitated the spectacular radiation of insects. Despite the undeniable advantage of complete metamorphosis, the female of some holometabolous species have lost the typical holometabolous development through neoteny. In Xenos vesparum Rossi (Strepsiptera: Stylopidae), a derived species of the holometabolous endoparasitic order Strepsiptera, neotenic females reach sexual maturity without the pupal and the imaginal stages, thus retaining their larval morphology (with the exception of the anterior part of the body or cephalothorax), while males undergo normal pupal-based metamorphosis. Expression of the "adult-specifier" E93 factor has been shown to be required for proper metamorphosis in holometabolous insects. Here, we investigated the involvement of E93 in female neoteny by cloning XvE93. Interestingly, while we detected a clear up-regulation of XvE93 expression in pupal and adult stages of males, persistent low levels of XvE93 were detected in X. vesparum females. However, a specific up-regulation of XvE93 was observed in the cephalothorax of late 4th female instar larva, which correlates with the occurrence of neotenic-specific features in the anterior part of the female body. Moreover, the same expression dynamic in the cephalothorax and abdomen was also observed for other two critical metamorphic regulators, the anti-metamorphic XvKr-h1 and the pupal specifier XvBr-C. The specific up-regulation of XvE93 and XvBr-C in the female cephalothorax seems to be the result of an increase in 20-hydroxyecdysone (20E) signaling in this region for we detected higher expression levels of the 20E-dependent nuclear receptors XvHR3 and XvE75 in the cephalothorax. Overall, our results detect a sex-specific expression pattern of critical metamorphic genes in X. vesparum, suggesting that neoteny in Strepsiptera results from the modification of the normal expression of E93, Br-C and Kr-h1 genes.
Asunto(s)
Proteínas de Insectos/metabolismo , Insectos/metabolismo , Animales , Ecdisterona/metabolismo , Femenino , Regulación del Desarrollo de la Expresión Génica/fisiología , Larva/metabolismo , Masculino , Metamorfosis Biológica/fisiología , Pupa/metabolismo , Transducción de Señal/fisiología , Factores de Transcripción/metabolismo , Regulación hacia Arriba/fisiologíaRESUMEN
Telomeres have a DNA component composed of repetitive sequences. In most eukaryotes these repeats are very similar in length and sequence and are maintained by a highly conserved specialized cellular enzyme, telomerase. Some exceptions of the telomerase mechanism exist in eukaryotes of which the most studied are concentrated in insects, and from these, Drosophila species stand out in particular. The alternative mechanism of telomere maintenance in Drosophila is based on targeted transposition of 3 very special non-LTR retrotransposons, HeT-A, TART and TAHRE. The fingerprint of the co-evolution between the Drosophila genome and the telomeric retrotransposons is visible in special features of both. In this chapter, we will review the main aspects of Drosophila telomeres and the telomere retrotransposons that explain how this alternative mechanism works, is regulated, and evolves. By going through the different aspects of this symbiotic relationship, we will try to unravel which have been the necessary changes at Drosophila telomeres in order to exert their telomeric function analogously to telomerase telomeres, and also which particularities have been maintained in order to preserve the retrotransposon personality of HeT-A, TART and TAHRE. Drosophila telomeres constitute a remarkable variant that reminds us how exceptions should be treasured in order to widen our knowledge in any particular biological mechanism.
Asunto(s)
Proteínas de Drosophila/genética , Drosophila/genética , Productos del Gen gag/genética , Genoma de los Insectos , Retroelementos/genética , Telómero/genética , Animales , Mapeo Cromosómico , Epigénesis Genética , Evolución Molecular , Modelos Genéticos , Especificidad de la Especie , Homeostasis del Telómero/genéticaRESUMEN
Telomeres across the genus Drosophila are maintained, not by telomerase, but by two non-LTR retrotransposons, HeT-A and TART, that transpose specifically to chromosome ends. Successive transpositions result in long head-to-tail arrays of these elements. Thus Drosophila telomeres, like those produced by telomerase, consist of repeated sequences reverse transcribed from RNA templates. The Drosophila repeats, complete and 5'-truncated copies of HeT-A and TART, are more complex than telomerase repeats; nevertheless, these evolutionary variants have functional similarities to the more common telomeres. Like other telomeres, the Drosophila arrays are dynamic, fluctuating around an average length that can be changed by changes in the genetic background. Several proteins that interact with telomeres in other species have been found to have homologues that interact with Drosophila telomeres. Although they have hallmarks of non-LTR retrotransposons, HeT-A and TART appear to have a special relationship to Drosophila. Their Gag proteins are efficiently transported into diploid nuclei where HeT-A Gag recruits TART Gag to chromosome ends. Gags of other non-LTR elements remain predominantly in the cytoplasm. These studies provide intriguing evolutionary links between telomeres and retrotransposable elements.
Asunto(s)
Drosophila/genética , Retroelementos/genética , Telómero/genética , Animales , Núcleo Celular/genética , Cromosomas/genética , Proteínas de Drosophila/genética , Proteínas de Drosophila/metabolismo , Evolución Molecular , Productos del Gen gag/genética , Productos del Gen gag/metabolismo , Proteínas Fluorescentes Verdes/genética , Proteínas Fluorescentes Verdes/metabolismo , Microscopía Fluorescente , Estructura Terciaria de Proteína/genética , Proteínas Recombinantes de Fusión/genética , Proteínas Recombinantes de Fusión/metabolismo , Retroelementos/fisiología , Telómero/metabolismoRESUMEN
Drosophila telomeres have been maintained by retrotransposition for at least 60 MY, which predates the separation of extant species of this genus. Studies of D. melanogaster, D. yakuba, and D. virilis show that, in Drosophila, telomeres are composed of two non-LTR retrotransposons, HeT-A and TART. Far from being static, HeT-A and TART evolve faster than Drosophila euchromatic genes. In spite of their high rate of sequence change, HeT-A and TART maintain their basic structures and unusual individual features. The maintenance of their separate identities suggests that HeT-A and TART cooperate either in the process of retrotransposition onto the chromosome end, or in the formation of telomere chromatin by transposed DNA copies. The telomeric retrotransposons and the Drosophila genome constitute an example of a robust symbiotic relationship between mobile elements and the genome.
Asunto(s)
Mapeo Cromosómico , Drosophila/genética , Evolución Molecular , Retroelementos , Animales , Secuencia de Bases , ADN/química , ADN/genética , Datos de Secuencia MolecularRESUMEN
The distribution of repetitive sequences, or microsatellites, formed by either one or two base pairs and longer than eight units, has been studied in almost 1 Mb of the sequenced Arabidopsis thaliana genome. Except for those formed by only G and C residues, the repetitions are more abundant in the Arabidopsis genome than can be calculated from its nucleotide composition. They are distributed in proportions higher than expected in introns, and in the intergenic regions both proximal and distal to the coding sequences. In exons, only the TC/GA microsatellite seems to be particularly abundant. The AT/TA microsatellites produce more length variation between Arabidopsis ecotypes than the A/T repeated sequences. These two classes are more abundant per kilobase than coding sequences in the Arabidopsis genome. The results indicate not only that the presence of microsatellites is not an effect of random distribution of nucleotides, but that their resolution as molecular markers may be equivalent to the number of genes and also that they do not seem to be systematically linked to specific regulatory sequences proximal to genes.
RESUMEN
Although the genome of Arabidopsis thaliana has a small amount of repetitive DNA, it contains representatives of most classes of mobile elements. However, to date, no miniature inverted-repeat transposable element (MITE) has been described in this plant. Here, we describe a new family of repeated sequences that we have named Emigrant, which are dispersed in the genome of Arabidopsis and fulfil all the requirements of MITEs. These sequences are short, AT-rich, have terminal inverted repeats (TIRs), and do not seem to have any coding capacity. Evidence for the mobility of Emigrant elements has been obtained from the absence of one of these elements in a specific Arabidopsis ecotype. Emigrant is also present in the genome of different Brassicae and its TIRs are 74% identical to those of Wujin elements, a recently described family of MITEs from the yellow fever mosquito Aedes aegypti.