Core Ideas Percentage coverage of TEs is significantly negatively correlated with that of genes. Different TE superfamilies exhibit distinct distribution patterns in mulberry genome. Copia elements may have a dominant influence on the regulation of mulberry genes. TE‐containing genes assigned to pathways were mainly in metabolism‐related pathways. TEs are a driving force in the formation of alternatively spliced genes. Mulberry (Morus notabilis C. K. Schneid) leaves have been used as the food for the domesticated silkworm, Bombyx mori, for more than 5000 yr, and the mulberry–silkworm relationship is one of the best‐known and oldest models of plant defense–insect adaptation. The availability of a genome assembly of mulberry provides us with an opportunity to mine the characteristics and distribution of transposable elements (TEs) in this species and to examine their relationship to genes and gene expression. In this study, a significantly correlated inverse relationship between the percentage coverage of genes and TEs was observed. The TE‐rich regions appeared to have a lower percentage of putatively expressed genes. Distribution patterns between different TE superfamilies were detected in the mulberry genome. The Copia elements (the TE making up the greatest proportion of the mulberry genome) were significantly overrepresented within genes in the mulberry genome, and they may have a dominant influence on evolution of the mulberry genome. Approximately 96.93% (330/344) of the TE‐containing genes assigned to pathways were assigned to metabolism‐related pathways. The TE‐related alternative splicing events accounted for 7.58% (402/5,302) of all alternative splicing types in the mulberry genome, suggesting that TEs are one of the driving forces in the formation of the alternatively spliced genes. The results will be valuable in improving our understanding of the important roles of TEs in mulberry genome evolution.
The evolutionary dynamics of long terminal repeat (LTR) retrotransposons in tree genomes has remained largely unknown. The availability of the complete genome sequences of the mulberry tree (Morus notabilis) has offered an unprecedented opportunity for us to characterize these retrotransposon elements. We investigated 202 and 114 families of Copia and Gypsy superfamilies, respectively, comprising 2916 intact elements in the mulberry genome. The tRNAMet was the most frequently used type of tRNA in both superfamilies. Phylogenetic analysis suggested that Copia and Gypsy from mulberry can be grouped into eight and six lineages, respectively. All previously characterized families of such elements could also be found in the mulberry genome. About 95% of the identified Copia and Gypsy full elements were estimated to have been inserted into the mulberry genome within the past 2–3 million years. Meanwhile, the estimated insertion times of members of the three most abundant families of the Copia superfamily (908 members from the three most abundant families) and Gypsy superfamily (783 members from the three most abundant families) revealed divergent life histories. Compared with the situation in Gypsy elements, three families of Copia elements are under positive selection pressure, which suggested that Copia elements may have a dominant influence in the evolution of mulberry genes. Analysis of insertion and deletion dynamics suggested that Copia and Gypsy elements exhibited a very long half-life in the mulberry genome. The present work provides new insights into the insertion and deletion dynamics of LTR retrotransposons, and it will greatly improve our understanding of the important roles transposable elements play in the architecture of the mulberry genome.
Horizontal transposable element transfer (HTT) events have occurred among a large number of species and play important roles in the composition and evolution of eukaryotic genomes. HTTs are also regarded as effective forces in promoting genomic variation and biological innovation. In the present study, HTT events were identified and analyzed in seven sequenced species of Rosales using bioinformatics methods by comparing sequence conservation and K/K value of reverse transcriptase (RT) with 20 conserved genes, estimating the dating of HTTs, and analyzing the phylogenetic relationships. Seven HTT events involving long terminal repeat (LTR) retrotransposons, two HTTs between Morus notabilis and Ziziphus jujuba, and five between Malus domestica and Pyrus bretschneideri were identified. Further analysis revealed that these LTR retrotransposons had functional structures, and the copy insertion times were lower than the dating of HTTs, particularly in Mn.Zj.1 and Md.Pb.3. Altogether, the results demonstrate that LTR retrotransposons still have potential transposition activity in host genomes. These results indicate that HTT events are another strategy for exchanging genetic material among species and are important for the evolution of genomes.
The effect of topographic correction (TOC) has a profound influence on the quantitative application of remote sensing image. With regard to the invalid evaluation of the TOC model with such a single topographic correction assessment (TCA) method, we have selected five TCA indexes from five different perspectives: the difference in mean radiance radiometry between sunlit and shaded slopes, the cosine empirical relationship, stability, heterogeneity, and outlier ratio. The entropy weight method was used to assign weight to each TCA indexes, and the comprehensive evaluation value (CEV) of TOC for each band of remote sensing image was obtained by weighted superposition. After that, the weight of each band of the remote sensing image is determined by the entropy weight method, and the CEV of the TOC of the remote sensing image is obtained by weighting and superposition, so as to realize the effect evaluation of the six TOC models of C, SCS + C, VECA, Teillet, Minnaert, and Minnaert + SCS. The results indicate that the proposed method can effectively evaluate the correction effect of the TOC model. Results indicate that the SCS + C model has the best correction effect, while the Minnaert model performs the worst. The results generated from the Minnaert + SCS, Teillet, and Minnaert models typically show inferior quality. The SCS + C, VECA, and C models are better suited for generating images with high spectral fidelity, and these three correction models are recommended for TOCs over mountainous areas.
Reverse transcriptase (rt) fragments from LINE retrotransposons in the mulberry genome were analyzed in terms of heterogeneity, phylogeny, and chromosomal distribution. We amplified and characterized conserved domains of the rt using degenerate primer pairs. Sequence analyses indicated that the rt fragments were highly heterogeneous and rich in A/T bases. The sequence identity ranged from 31.8% to 99.4%. Based on sequence similarities, the rt fragments were categorized into eight groups. Furthermore, similar stop codon distribution patterns among a series of clones in the same group indicated that they underwent a similar evolutionary process. Interestingly, phylogenetic analyses of the rt fragments isolated from mulberry and 13 other plant species revealed that two distantly related taxa (mulberry and Paeonia suffruticosa) grouped together. It does not appear that this phenomenon resulted from horizontal transposable element transfer. Fluorescence in situ hybridization analysis revealed that most of the rt fragments were concentrated in the subtelomeric and pericentromeric regions of the mulberry chromosomes, but that these elements were not abundant in the mulberry genome. Future studies will focus on the potential roles of these elements in the subtelomeric and pericentromeric regions of the mulberry genome.
Copia and Gypsy play important roles in structural, functional and evolutionary dynamics of plant genomes. In this study, a total of 106 and 101, Copia and Gypsy reverse transcriptase (rt) were amplified respectively in the Morus notabilis genome using degenerate primers. All sequences exhibited high levels of heterogeneity, were rich in AT and possessed higher sequence divergence of Copia rt in comparison to Gypsy rt. Two reasons are likely to account for this phenomenon: a) these elements often experience deletions or fragmentation by illegitimate or unequal homologous recombination in the transposition process; b) strong purifying selective pressure drives the evolution of these elements through “selective silencing” with random mutation and eventual deletion from the host genome. Interestingly, mulberry rt clustered with other rt from distantly related taxa according to the phylogenetic analysis. This phenomenon did not result from horizontal transposable element transfer. Results obtained from fluorescence in situ hybridization revealed that most of the hybridization signals were preferentially concentrated in pericentromeric and distal regions of chromosomes, and these elements may play important roles in the regions in which they are found. Results of this study support the continued pursuit of further functional studies of Copia and Gypsy in the mulberry genome.
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