12 research outputs found
Unassigned Codons, Nonsense Suppression, and Anticodon Modifications in the Evolution of the Genetic Code
The origin of the genetic code is a central open
problem regarding the early evolution of life. Here, we
consider two undeveloped but important aspects of possible
scenarios for the evolutionary pathway of the translation
machinery: the role of unassigned codons in early stages of
the code and the incorporation of tRNA anticodon modifications.
As the first codons started to encode amino acids,
the translation machinery likely was faced with a large
number of unassigned codons. Current molecular scenarios
for the evolution of the code usually assume the very rapid
assignment of all codons before all 20 amino acids became
encoded. We show that the phenomenon of nonsense
suppression as observed in current organisms allows for a
scenario in which many unassigned codons persisted
throughout most of the evolutionary development of the
code. In addition, we demonstrate that incorporation of
anticodon modifications at a late stage is feasible. The
wobble rules allow a set of 20 tRNAs fully lacking anticodon
modifications to encode all 20 canonical amino
acids. These observations have implications for the biochemical
plausibility of early stages in the evolution of the
genetic code predating tRNA anticodon modifications and
allow for effective translation by a relatively small and
simple early tRNA set
A mathematical model of kinetoplastid mitochondrial gene scrambling advantage
We model and discuss advantages of pan-editing, the complex way of expressing
mitochondrial genes in kinetoplastids. The rapid spread and preservation of pan-editing
seems to be due to its concomitant fragment dispersal. Such dispersal prevents losing
temporarily non expressed mitochondrial genes upon intense intraspecific competition,
by linking non expressed fragments to parts which are still needed. We mathematically
modelled protection against gene loss, due to the absence of selection, by this kind of
fragment association. This gives ranges of values for parameters like scrambling extent,
population size, and number of generations still retaining full genomes despite limited
selection. Values obtained seem consistent with those observed. We find a quasi-linear
correlation between dispersal and number of generations after which populations lose
genes, showing that pan-editing can be selected to effectively limit gene loss under
relaxed selective pressure
On distinguishing between canonical tRNA genes and tRNA gene fragments in prokaryotes
Automated genome annotation is essential for extracting biological information from sequence data. The identification and annotation of tRNA genes is frequently performed by the software package tRNAscan-SE, the output of which is listed for selected genomes in the Genomic tRNA database (GtRNAdb). Here, we highlight a pervasive error in prokaryotic tRNA gene sets on GtRNAdb: the miscategorization of partial, non-canonical tRNA genes as standard, canonical tRNA genes. Firstly, we demonstrate the issue using the tRNA gene sets of 20 organisms from the archaeal taxon Thermococcaceae. According to GtRNAdb, these organisms collectively deviate from the expected set of tRNA genes in 15 instances, including the listing of eleven putative canonical tRNA genes. However, after detailed manual annotation, only one of these eleven remains; the others are either partial, noncanonical tRNA genes resulting from the integration of genetic elements or CRISPR-Cas activity (seven instances), or attributable to ambiguities in input sequences (three instances). Secondly, we show that similar examples of the mis-categorization of predicted tRNA sequences occur throughout the prokaryotic sections of GtRNAdb. While both canonical and non-canonical prokaryotic tRNA gene sequences identified by tRNAscan-SE are biologically interesting, the challenge of reliably distinguishing between them remains. We recommend employing a combination of (i) screening input sequences for the genetic elements typically associated with non-canonical tRNA genes, and ambiguities, (ii) activating the tRNAscan-SE automated pseudogene detection function, and (iii) scrutinizing predicted tRNA genes with low isotype scores. These measures greatly reduce manual annotation efforts, and lead to improved prokaryotic tRNA gene set predictions
On distinguishing between canonical tRNA genes and tRNA gene fragments in prokaryotes
Automated genome annotation is essential for extracting biological information from sequence data. The identification and annotation of tRNA genes is frequently performed by the software package tRNAscan-SE, the output of which is listed for selected genomes in the Genomic tRNA database (GtRNAdb). Here, we highlight a pervasive error in prokaryotic tRNA gene sets on GtRNAdb: the miscategorization of partial, non-canonical tRNA genes as standard, canonical tRNA genes. Firstly, we demonstrate the issue using the tRNA gene sets of 20 organisms from the archaeal taxon Thermococcaceae. According to GtRNAdb, these organisms collectively deviate from the expected set of tRNA genes in 15 instances, including the listing of eleven putative canonical tRNA genes. However, after detailed manual annotation, only one of these eleven remains; the others are either partial, noncanonical tRNA genes resulting from the integration of genetic elements or CRISPR-Cas activity (seven instances), or attributable to ambiguities in input sequences (three instances). Secondly, we show that similar examples of the mis-categorization of predicted tRNA sequences occur throughout the prokaryotic sections of GtRNAdb. While both canonical and non-canonical prokaryotic tRNA gene sequences identified by tRNAscan-SE are biologically interesting, the challenge of reliably distinguishing between them remains. We recommend employing a combination of (i) screening input sequences for the genetic elements typically associated with non-canonical tRNA genes, and ambiguities, (ii) activating the tRNAscan-SE automated pseudogene detection function, and (iii) scrutinizing predicted tRNA genes with low isotype scores. These measures greatly reduce manual annotation efforts, and lead to improved prokaryotic tRNA gene set predictions
On distinguishing between canonical tRNA genes and tRNA gene fragments in prokaryotes
Automated genome annotation is essential for extracting biological information from sequence data. The identification and annotation of tRNA genes is frequently performed by the software package tRNAscan-SE, the output of which is listed for selected genomes in the Genomic tRNA database (GtRNAdb). Here, we highlight a pervasive error in prokaryotic tRNA gene sets on GtRNAdb: the mis-categorization of partial, non-canonical tRNA genes as standard, canonical tRNA genes. Firstly, we demonstrate the issue using the tRNA gene sets of 20 organisms from the archaeal taxon Thermococcaceae. According to GtRNAdb, these organisms collectively deviate from the expected set of tRNA genes in 15 instances, including the listing of eleven putative canonical tRNA genes. However, after detailed manual annotation, only one of these eleven remains; the others are either partial, non-canonical tRNA genes resulting from the integration of genetic elements or CRISPR-Cas activity (seven instances), or attributable to ambiguities in input sequences (three instances). Secondly, we show that similar examples of the mis-categorization of predicted tRNA sequences occur throughout the prokaryotic sections of GtRNAdb. While both canonical and non-canonical prokaryotic tRNA gene sequences identified by tRNAscan-SE are biologically interesting, the challenge of reliably distinguishing between them remains. We recommend employing a combination of (i) screening input sequences for the genetic elements typically associated with non-canonical tRNA genes, and ambiguities, (ii) activating the tRNAscan-SE automated pseudogene detection function, and (iii) scrutinizing predicted tRNA genes with low isotype scores. These measures greatly reduce manual annotation efforts, and lead to improved prokaryotic tRNA gene set predictions
The contours of evolution: In defence of Darwin's tree of life paradigm
Both the concept of a Darwinian tree of life (TOL) and the possibility of its accurate reconstruction have been much criticized. Criticisms mostly revolve around the extensive occurrence of lateral gene transfer (LGT), instances of uptake of complete organisms to become organelles (with the associated subsequent gene transfer to the nucleus), as well as the implications of more subtle aspects of the biological species concept. Here we argue that none of these criticisms are sufficient to abandon the valuable TOL concept and the biological realities it captures. Especially important is the need to conceptually distinguish between organismal trees and gene trees, which necessitates incorporating insights of widely occurring LGT into modern evolutionary theory. We demonstrate that all criticisms, while based on important new findings, do not invalidate the TOL. After considering the implications of these new insights, we find that the contours of evolution are best represented by a TOL