Friday, September 14, 2012

Construction and Application of Genomic DNA Libraries in Plant

Crop failures, pathogenic outbreaks and famine are serious problems facing society. Understanding an organism's genome will help provide the genetic tools needed to solve these complex problems in a shorter time and with lesser effort. To efficiently study an organism's genome, it can be partitioned into a permanent and stable collection of DNA fragments, called a library. Such libraries provide convenient access to a genome for both laboratory and breeding applications. Genomic libraries can be used as substrates to physically map and sequence entire genomes, clone agriculturally important genes and to investigate gene expression patterns. Further, genomic libraries also provide powerful tools and resources for evaluating germplasm conservation stocks and biological diversity. Since the ongoing explosion of genetic data and molecular clone resources has opened new scientific possibilities with unfamiliar terms and acronyms to researchers venturing across all avenues of applied science

Serious consideration must be given to the various kinds of libraries that can be constructed with respect to different insert sizes, fragmentation methods, vectors, coverage, downstream operations, etc. Though downstream applications (see the Applications section of this chapter) differ for types of libraries, fragmentation size is the first consideration in genomic library construction. Genomic DNA libraries are classified as shotgun libraries, medium-size insert libraries and large insert libraries depending on insert sizes. A shotgun library, with a smaller insert size (generally 1.5–10 kbp), is made using high-copy plasmid vectors. It is the most common resource for complete sequencing of large genomic DNA clones (e.g. cosmid, PAC, BAC and YAC clones) and large DNA fragments (PCR products and restriction fragments). With the introduction of capillary sequencers for large-scale, high-throughput DNA sequencing, the shotgun library approach is considered a standard method for generating sequence-ready sublibraries for genome projects and positional cloning studies.

Cosmid and fosmid libraries contain medium-sized inserts (35–45 kbp). Cosmids are hybrids of λλ-phage and plasmids. They can replicate in the cell like a plasmid due to the plasmid replication origin and be packaged like a phage because of existing cos sites. Since most of the λλ-phage structure has been deleted, they can carry DNA inserts up to 45 kbp. This type of library is useful for cloning studies because of its simplicity and economic efficiency for construction in comparison with large insert library construction. The fosmid cloning system is similar in size to cosmids and the vector is derived from the endogenous E. coli F1 factor that maintains inserts in a single-copy state adding to the fidelity and stability of the DNA insert. The fosmid system is useful for easy generation of medium-sized insert (45 kbp) genomic libraries produced from small amounts of source DNA such as flow-sorted chromosomal DNA (Gingrich et al., 1996).

Large-insert genomic DNA libraries are essential for physical mapping, positional cloning and genome sequencing of complex genomes. There are two principal large-insert cloning systems: yeast and bacterial artificial chromosome systems (YACs and BACs). The YAC cloning system was first developed in 1987 (Burke et al., 1987), and uses Saccharomyces cerevisiae as the host and maintains large inserts (up to 1 Mb) as linear molecules with a pair of yeast telomeres and a centromere. Although used extensively in the late 1980s and early 1990s, this system has several disadvantages (Anderson, 1993; Zhang and Wing, 1997). The recombinant DNA in yeast can be unstable. DNA manipulation is difficult and inefficient. Most importantly, a high level of chimerism—cloning of two or more unlinked DNA fragments in a single molecule—is inherent within the YAC cloning system. These disadvantages impede the utility of YAC libraries and subsequently this system has been gradually replaced by the BAC cloning system introduced in 1992 (Shizuya et al., 1992).

The BAC system uses a derivative of the E. coli F-factor as a vector and E. coli as the host, making BAC cloning and subsequent down-stream procedures efficient and easy to perform. Recombinant DNA inserts of up to 200 kb can be efficiently cloned and stably maintained as single-copy plasmids in E. coli. BAC libraries have been developed for essentially all major crop plants, as well as model plant species, such as Arabidopsis thaliana and rice, and can be accessed from the laboratories that made the libraries or through various stock centres (e.g. AGI, CUGI and ABRC).

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