Original research
The Mitochondrial Genome of Raphanus sativus and Gene Evolution of Cruciferous Mitochondrial Types

https://doi.org/10.1016/j.jgg.2013.01.003Get rights and content

Abstract

To explore the mitochondrial genes of the Cruciferae family, the mitochondrial genome of Raphanus sativus (sat) was sequenced and annotated. The circular mitochondrial genome of sat is 239,723 bp and includes 33 protein-coding genes, three rRNA genes and 17 tRNA genes. The mitochondrial genome also contains a pair of large repeat sequences 5.9 kb in length, which may mediate genome reorganization into two sub-genomic circles, with predicted sizes of 124.8 kb and 115.0 kb, respectively. Furthermore, gene evolution of mitochondrial genomes within the Cruciferae family was analyzed using sat mitochondrial type (mitotype), together with six other reported mitotypes. The cruciferous mitochondrial genomes have maintained almost the same set of functional genes. Compared with Cycas taitungensis (a representative gymnosperm), the mitochondrial genomes of the Cruciferae have lost nine protein-coding genes and seven mitochondrial-like tRNA genes, but acquired six chloroplast-like tRNAs. Among the Cruciferae, to maintain the same set of genes that are necessary for mitochondrial function, the exons of the genes have changed at the lowest rates, as indicated by the numbers of single nucleotide polymorphisms. The open reading frames (ORFs) of unknown function in the cruciferous genomes are not conserved. Evolutionary events, such as mutations, genome reorganizations and sequence insertions or deletions (indels), have resulted in the non-conserved ORFs in the cruciferous mitochondrial genomes, which is becoming significantly different among mitotypes. This work represents the first phylogenic explanation of the evolution of genes of known function in the Cruciferae family. It revealed significant variation in ORFs and the causes of such variation.

Introduction

Mitochondrial genomes are important for supplying ATP in the development and reproduction of plants. Plant mitochondrial genomes vary greatly in size, ranging from 187 kb (Oda et al., 1992) to over 11.3 Mb (Sloan et al., 2012). The organization of plant mitochondrial genomes is known to be reversibly dynamic compared with their counterparts in animals or fungi, which affects the molecular pool of the peculiar mitochondrial genomes of seed plants due to mediation via various large repeated sequences (> 1 kb). Irreversible restructuring of plant mitochondrial genomes has certainly occurred to form genomes corresponding to individual energetic systems, mediated by short repeats (Andre et al., 1992; Newton et al., 2004). The mitochondrial genomes of plant species have lost numerous genes and acquired many genes during their evolution. Based on these properties, conducting sequence analysis of these genomes is important for identifying vital activities in seed plants. Analysis of the evolution of mitochondrial genes or ORFs over a long time-span is necessary to reveal evolutionary sequelae indicative of plant speciation. In this study, the mitochondrial genome of Raphanus sativus (sat), of the Cruciferae family, was sequenced and annotated. How evolutionary factors such as mutation, insertion/deletion and rearrangement resulted in the mitochondrial types within this plant family was also explored.

R. sativus (radish), a member of the Cruciferae family, is widely cultivated in Asia, especially in China, Japan and South Korea (Curtis, 2011). The enlarged root of the radish is commonly consumed as a fresh food or can be utilized as fodder as well as green manure (Noreen and Ashraf, 2009). Classifications of radish mitotypes and certain genes in radish mitochondria have been reported (Sakai and Imamura, 1992; Albaum et al., 1995; Kim et al., 2007). The involvement of short repeat sequences in dynamic mtDNA rearrangements has also been studied in the radish (Lee et al., 2009).

The Cruciferae family was chosen to analyze the evolution of mitochondrial genes because it holds an important position among angiosperm families. The family contains key vegetable and oilseed crops as well as several model organisms, including Arabidopsis thaliana and certain Brassica species (Bailey et al., 2006). As a model for plant biology, the Cruciferae have been extensively studied. Among this family, the first reported mitochondrial genome was that of A. thaliana (termed as the tha mitotype) (Unseld et al., 1997). The mitotype sequence of Brassica napus (nap) was subsequently published (Handa, 2003). In a previous study, we sequenced the mitochondrial genomes of five Brassica mitotypes: cam (B. rapa), ole (B. oleracea), pol (B. napus cultivar Polima), jun (B. juncea) and car (B. carinata) (Chang et al., 2011; Chen et al., 2011).

In this study, we present the complete mitochondrial genome sequence for the radish (referred to as sat). We provide the annotated mitotype of the common radish that is most commonly cultivated in Northern China and analyzed the properties of sat. The sat mitotype and six previously reported cruciferous mitotypes were used as a representative sample of Cruciferae family mitochondrial genomes to analyze the evolution of genes with known functions and the variation among ORFs with unknown functions within this family. The reasons for this variation were also explored.

Section snippets

The mitochondrial genome of R. sativus

For mitochondrial genome sequencing of R. sativus, 40,774 sequencing reads was applied, and the total sequence of 18,393,340 bp was obtained. The initial assembly of the sequences was unsuccessful because of the limited coverage ratio for the mitochondrial genome. Further sequencing to increase the coverage resulted in a coverage ratio of 76×. Assembly of these sequences using Newbler resulted in four large contigs (mean length of 58,414 bp). These contigs were joined through Sanger sequencing

Discussion

In the present study, the genes of the sat mitochondrial genome were identified (Table S1), and a tripartite genome structure and large structural differences from other cruciferous mitochondrial genomes were demonstrated at the sequence level. These data will undoubtedly benefit research on the mechanisms of energetic metabolism, development and reproduction of R. sativus. The examination of mitochondrial genome and gene evolution in the Cruciferae revealed that cruciferous mitotypes have lost

Mitochondrial DNA isolation and sequencing

Mitochondrial DNA was isolated from the seeds of the local Nanjing variety of radish (chuanxinhong) according to the methods described by Chen et al. (2011) and stored in our laboratory. Genome sequencing was performed using the GS-FLX platform (Roche, CT, USA). Newbler v.2.6 (454 Life Science Corp, CT, USA) was used to assemble the reads into contigs. Sanger sequencing of PCR products was conducted to join the contigs to form the complete genome.

Sequence data analysis

The mitochondrial sequences were annotated with

Acknowledgements

This work was supported by the National Basic Research Program of China (973 Program) (No. 2011CB109300), the National Natural Science Foundation of China (No. 30970289), the National Key Technology R&D Program (Nos. 2010BAD01B02 and 2011BAD13B09) in China and the Special Fund for Independent innovation of Agricultural Science and Technology in Jiangsu province (Nos. CX (10) 1030 and CX (11) 1026). The authors thank Shanghai Majorbio Bio-pharm Biotechnology Company (China) for their help on the

References (31)

  • I.S. Curtis

    Genetic engineering of radish: current achievements and future goals

    Plant Cell Rep.

    (2011)
  • A.E. Darling et al.

    progressiveMauve: multiple genome alignment with gene gain, loss and rearrangement

    PLoS ONE

    (2010)
  • H. Handa

    The complete nucleotide sequence and RNA editing content of the mitochondrial genome of rapeseed (Brassica napus L.): comparative analysis of the mitochondrial genomes of rapeseed and Arabidopsis thaliana

    Nucleic Acids Res.

    (2003)
  • S. Kim et al.

    Identification of a novel mitochondrial genome type and development of molecular markers for cytoplasm classification in radish (Raphanus sativus L.)

    Theor. Appl. Genet.

    (2007)
  • T. Kubo et al.

    The complete nucleotide sequence of the mitochondrial genome of sugar beet (Beta vulgaris L.) reveals a novel gene for tRNACys(GCA)

    Nucleic Acids Res.

    (2000)
  • View full text