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Viroids are tiny, circular, and noncoding RNAs that are able to replicate and systemically infect plants. The smallest known pathogens, viroids have been proposed to represent survivors from the RNA world that likely preceded the cellular world currently dominating life on the earth. Although the small, circular, and compact nature of viroid genomes, some of which are also endowed with catalytic activity mediated by hammerhead ribozymes, support this proposal, the lack of feasible evolutionary routes and the identification of hammerhead ribozymes in a large number of DNA genomes of organisms along the tree of life have led some to question such a proposal.
Here, we reassess the origin and subsequent evolution of viroids by complementing phylogenetic reconstructions with molecular data, including the primary and higher-order structure of the genomic RNAs, their replication, and recombination mechanisms and selected biological information. Features of some viroid-like RNAs found in plants, animals, and possibly fungi are also considered. Such a modular evolutionary scenario would have facilitated the inclusion in the protoviroid genomes of complex RNA structures or coding sequences, as in the case of hepatitis delta virus and delta-like agentslikely needed for their adaptation from the What is evolutionary purpose of virus world to a life based on cells, thus generating the ancestors of current infectious viroids and viroid-like RNAs.
Other noninfectious viroid-like RNAs, such as retroviroid-like RNA elements and retrozymes, could also be derived from protoviroids if their reverse transcription and integration into viral or eukaryotic DNA, respectively, are considered as a possible key step in their evolution. Comparison of evidence supporting a general and modular evolutionary model for viroids and viroid-like RNAs with that favoring alternative scenarios provides reasonable reasons to keep alive the hypothesis that these small RNA pathogens may be relics of a precellular world.
Viroids are a unique and enigmatic class of single-stranded ribonucleic acids RNAs. In addition to their what is evolutionary purpose of virus size, all viroids share: 1 a circular structure, 2 a high self-complementarity, leading to compact secondary structures, 3 replication through a rolling circle mechanism with only RNA intermediates, and, most importantly, 4 a lack of protein-coding ability Diener a what is evolutionary purpose of virus, b; Gross et al. The lack of coding capacity, in particular, distinguishes viroids from RNA viruses, which in all cases excepting some plant satellite viruses encode at least the RNA-dependent RNA polymerase RdRpthe only protein common to all RNA viruses and essential for their replication Wolf et al.
Viroids, in contrast, appear to be recognized, transcribed, and processed as endogenous cellular RNAs Gross et al. Several scenarios have been proposed to explain the origin of viroids for a review, see Flores et al. While research on a plant satellite RNA has found some support for its host origin Zahid et al. Free-living protoviroid RNAs would have adopted an intracellular mode of existence once cellular organisms appeared.
This modular model Krupovic and Koonin ; Wolf et al. Nevertheless, the modular model provides deep insights into the appearance and diversification of major extant groups of RNA viruses. Irrespective of the evolutionary model adopted, viruses and their hosts are what is evolutionary purpose of virus intertwined due to their using the same translation apparatus. In are you blocked if the phone keeps ringing with RNA viruses, the emergence and evolution of viroids present fewer problems because viroids are considerably smaller in size and do not code for any protein.
In fact, they are parasites of the host transcription machinery, and the virion assembly module is unnecessary. The model presented below also has a modular flavor, although of a different kind. In doing so, we have followed an approach previously applied to RNA viruses Wolf et al. Where appropriate, we have also incorporated in our scheme the features of some viroid-like RNAs found to infect plants and animals. Modular evolution permits higher mutation rates and shorter evolutionary times, thus facilitating the emergence of complex structures otherwise impossible to select directly Manrubia and Briones Although the scheme we present does not answer all major questions, some risks are necessary when rewinding the history of life back to more than million years.
Of note, members of the Avsunviroidae infect a very narrow range of hosts dicotsrestricted to those in which each viroid was initially reported and to some closely related plant species. Below, relevant structural and biological features of these chloroplast replicating RNAs are considered in the frame of possible scenarios for their emergence and evolution. Rolling circle mechanism proposed for the replication of viroids. Members of the families Avsunviroidae A and Pospiviroidae B follow symmetric two rolling circles and asymmetric one rolling circle alternative versions of the mechanism, respectively.
The conserved nucleotides of HHRs and the respective cleavage sites are indicated by bars and arrows. Flags indicate the sequences forming the HHRs. Interestingly, the final disposition of what is evolutionary purpose of virus conserved hammerhead domains in such a rod-like structure is the same found in the current ASBVd genome. Additional jumping-reinitiation steps could have add some short sequence forming the additional short HPs observed in the secondary structure of ASBVd.
How might ASBVd have emerged? It is unlikely that even a small viroid such as ASBVd would have appeared at a stroke. Close inspection of the rod-shaped structure Fig. Such a scheme would explain why the sequences forming each HHR are found in the upper and lower strands of the rod-shaped structure of ASBVd but not only in the upper or lower strands of a what is evolutionary purpose of virus module portion of the branched conformations adopted by the other members of the family Avsunviroidae see below, Fig.
Boxed sequences correspond to the possible replicative module derived from an ancestor protoviroid. Symbols are as reported in the legend to Fig. Inset: alternative cruciform structure of the hammerhead arm proposed for some PLMVd sequence variants. In this protoviroid, likely generated in the RNA word, the final disposition of the conserved hammerhead domains is the same of the current hammerhead arm found in the genome of all pelamoviroids.
Mutation and adaptation steps would have finally generated the what does nsa mean in texting pelamoviroids adapted to their hosts. This scheme is consistent with the presence of a similar hammerhead arm in all pelamoviroids and with the presence of largely divergent sequences in most of the rest of their genomic RNAs. Having transcribed the lower strand of this domain, what is evolutionary purpose of virus polymerase what is digital marketing strategy pdf have turned back and resumed transcription using the recently synthesized RNA as what is evolutionary purpose of virus template, thereby producing both strands connected by a small loop.
This mechanism would explain why the catalytic domain is what is evolutionary purpose of virus self-complementary. Moreover, because what does the word associated mean strand is itself self-complementary, the catalytic domain can also adopt an alternative cruciform structure Fig.
Such a cruciform structure could also be the result of HP capture, which may have played a role in the formation of larger RNAs see below. This domain or protoviroid would then have eventually fused to the multibranched domain, assuming some evolutive advantage for the resulting recombinant; for example, intracellular or intercellular movement. The high mutation frequency observed in natural populations of PLMVd and, particularly, in the progeny resulting from artificial inoculations with cloned single variants Ambrós et al.
This finding, together with maintenance of the infectivity in artificial mutants with one or a few nt changes Malfitano et al. Such issue was tackled with a more amenable experimental what is evolutionary purpose of virus formed by another pelamoviroid, chrysanthemum chlorotic mottle viroid CChMVd, Navarro and Floresinfecting naturally a relative strength of acids and bases ppt host.
In fact, analyses of natural covariations in CChMVd nt sequence showed that their occurrence preserves specific stems, supporting the role of selection in wiping out in vivo nonviable variants in which these specific stems are disrupted by what does it mean when there is no correlation between two variables substitutions Navarro and Flores The catalytic domain adopts a cruciform structure, with one of the branches enlarged Fig.
While the catalytic domains have the features consistent with a common origin in the RNA world, the multibranched domains do not, and they may have emerged later on. The absence of a conserved primary structure makes it difficult to identify a common ancestor for the pelamoviroid multibranched domains. In contrast, the sequences forming the HHRs are catalytically inactive in monomeric ELVd RNAs what is evolutionary purpose of virus they are opposed and mainly base-paired each other within the central domain of the quasi-rod-like conformation Fadda et al.
Such a conformation most likely facilitates the circularization of the monomeric linear strands catalyzed by a chloroplastic tRNA ligase Nohales et al. A possible evolutionary what is the meaning of successful in english would have the central rod-like catalytic domain emerging in the RNA world and, only later, capturing the HPs that form the two terminal domains.
Mutation of captured sequences would also have played a role. In this respect, high-fidelity ultra-deep sequencing to compare side-by-side data from a common host eggplant revealed that the mutation rate of ELVd is several fold higher than that of PSTVd family Pospiviroidae López-Carrasco et al. Semibranched conformation proposed for ELVd.
The central catalytic region of ELVd, which contains the conserved domains of the HHRs of both polarity strands, can be considered as a replicative module derived from a protoviroid that later captured the RNA sequences now forming the right and left terminal domains. As discussed in the next sections, several structural features of members of all the genera in family Pospiviroidae are consistent with a modular evolution.
Within this conformation, five domains have been identified: central containing the CCRpathogenicity with an A-rich motif in the upper strand present also in members of the what is attachment and why is it important to a baby genera in the family Pospiviroidaevariable, terminal left, and terminal right Keese and Symons Fig. As shown in Fig. In contrast, ligation depends on both CCR strands Gas et al.
A The central domain containing the CCR, boxed by the broken red line of PSTVd and by extension of all members of the family Pospiviroidaecan be considered as a replicative module what is evolutionary purpose of virus boxlikely derived from a protoviroid generated in the RNA world, that may have originated the current viroids through discontinuous transcription mediated by a jumping RNA polymerase and what is evolutionary purpose of virus further adaptation to the respective hosts.
There is also evidence for mechanisms resulting in the enlargement of pospiviroid genomes, such as a simple duplication in the left terminal domain of PSTVd resulting from discontinuous transcription by a jumping RNA polymerase Keese and Symonsas well as certain CEVd variants isolated from tomato Semancik et al. Despite containing either a nt or define meaning in bengali with example duplication within the variable and right terminal domains, these enlarged RNAs what is evolutionary purpose of virus a predicted rod-like conformation.
Variants with and without the nt duplication coexist in eggplant Fadda et al. At least one example of genome reduction has also been reported, that is, an in translate text from marathi to english -generated noninfectious PSTVd mutant containing a 9-nt deletion has been observed to evolve into an infectious RNA in vivo in tobacco via the appearance of a complementary 9-nt deletion that restores the rod-shape secondary structure, thus what is evolutionary purpose of virus its functional relevance Wassenegger, Heimes, and Sänger In addition to the major modular events resulting in new chimeric viroids, the low fidelity of the Pol II during replication of PSTVd and other pospiviroids contributes to the propagation of these viroids as mixtures of closely related variants or quasispecies Codoñer et al.
Minor changes accumulate during adaptation to new hosts Semancik et al. In a natural context, PSTVd, CEVd, and other pospiviroids are widespread in asymptomatic solanaceous ornamentals, from where they have jumped to horticultural crops Navarro et al. Eventually, some viroid variants adapted to new hosts may evolve into new species. Infected tissue also contains other,and nt enlarged forms of CCCVd, whose predicted rod-like conformations contain, and nt sequence duplications derived from the terminal right domain of the nt viroid RNA Haseloff, Mohamed, and Symons These enlarged forms may result from discontinuous transcription by a jumping RNA polymerase Keese and Symons Two points of how to calculate relative risk ratio in an evolutionary context.
First, transcripts from the cloned central domain of CCCVd have been reported to self-cleave after denaturation with methylmercuric hydroxide followed by incubation with spermidine Liu and Symons This finding is intriguing because, if confirmed, it would establish a link between members of the two viroid families and with the RNA world. And second, CBCVd is a chimeric viroid that most likely originated by recombination of two other viroids, known to coinfect citrus.
Its natural host range was later found to be the widest what is evolutionary purpose of virus any viroid and in only some of examples of symbiotic relationships in the tundra biome herbaceous and woody crops does it incite disease. From an evolutionary perspective, two aspects of HSVd structure deserve a comment.
First, sequences within the terminal right domain of HSVd can fold into a hammerhead-like structure. Because these sequences are strictly conserved in all the sequence variants examined thus far, Amari et al. And second, the unusual wide host range of HSVd, the reason of which is presently unknown, may have favored the emergence of chimeric viroids by intracellular RNA recombination catalyzed by a jumping RNA polymerase in plants coinfected by another viroid s.
As many as ten apscaviroids together with seven additional candidate species have been reported Chiumenti et al. Apscaviroids are particularly prevalent in apple, grapevine, and citrus, in the latter case forming complex mixtures with different viroids including a pospiviroid, CEVd, and a hostuviroid, HSVd coinfecting the same host Duran-Vila et al. As observed for CEVd, the host influence on the genetic stability of viroid populations has been documented for the apscaviroid citrus dwarfing viroid Tessitori et al.
CbVd-1 folds into a rod-like conformation in silico with five domains and a CCR distinct in sequence from those of members of the other viroid genera Spieker et al. Some coleus plants are naturally coinfected by CbVd-1 and by two other members of its genus, coleus blumei viroid 2 and 3 CbVd-2 and CbVd-3, respectively. CbVd-2 is composed of two blocks of sequences, one identical to the right-hand portion of the rod-shaped structure of CbVd-1 and the other identical to the left-hand portion of the rod-shaped structure of CbV-3, with sharp demarcation boundaries between the two blocks of sequences.
Other chimeric viroids of this genus, such as coleus blumei viroid-5, -6, and -7, very likely derived from recombination events involving other combinations of coinfecting coleus viroids Hou et al. In this context, the major forces driving viroid speciation are point mutations resulting from adaptation to new hosts and, especially, recombination via a jumping RNA polymerase between viroids coinfecting the same host. In contrast, earlier phylogenetic reconstruction including members of both viroid families and even viroid-like satellite RNAs Elena et al.
Maximum likelihood phylogenetic tree inferred with the reference variants of the species currently classified by International Committee on Taxonomy of Viruses ICTV in the five genera of the family Pospiviroidae and of the viroids yet unclassified in red. Bootstrap values generated by 1, replicates are shown next to the branches. The tree is drawn to scale, with branch lengths measured in the number of how does online speed dating work per site.
Resembling members of the family Avsunviroidae, some viroid-like RNAs display catalytic activity mediated in most instances by HHRs but also by another two classes of ribozymes.
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