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Principles of Plant Genetics and Breeding

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238 CHAPTER 14<br />

rDNA is that DNA is universal <strong>and</strong> hence the transfer <strong>of</strong><br />

DNA is possible, in theory, across biological barriers.<br />

Eukaryotes are genetically complex organisms, <strong>and</strong> not<br />

directly amenable to certain biotechnological procedures.<br />

Consequently, scientists pick plants <strong>and</strong> animals<br />

that are easy to manipulate, <strong>and</strong> invest in a thorough<br />

genetic analysis <strong>of</strong> their genomes. The information<br />

derived from such studies is extrapolated to more complex<br />

organisms. Arabidopsis thaliana was the first plant<br />

to be completely sequenced. This flowering plant has a<br />

short life cycle <strong>of</strong> about 6 weeks from germination to<br />

seed maturity. It produces seed pr<strong>of</strong>usely. Because <strong>of</strong> its<br />

small size, Arabidopsis can be cultured in trays or pots in<br />

a limited space. Two <strong>of</strong> the most important higher<br />

plants used for food that are being developed as model<br />

plants are rice (Oriza sativa) <strong>and</strong> maize (Zea mays).<br />

There are several subspecies <strong>of</strong> rice, the most widely<br />

eaten being O. indica, followed by O. japonica.<br />

Through both public <strong>and</strong> private sector efforts, the<br />

genome sequences <strong>of</strong> the two subspecies have been<br />

developed. Genomes <strong>of</strong> all cereals (wheat, barley, rice,<br />

corn, etc.) are structurally similar (see next Section).<br />

Genomic colinearity <strong>and</strong> its application in<br />

plant breeding<br />

Genomic colinearity is a term used to refer to the<br />

conservation (through evolutionary history) <strong>of</strong> gene<br />

content, order, <strong>and</strong> orientation between chromosomes<br />

<strong>of</strong> different species, or between non-homologous chromosomes<br />

within a single species. The term is sometimes<br />

used as synonymous with synteny, the phenomeonon <strong>of</strong><br />

conservation <strong>of</strong> gene order within related genomes.<br />

Synteny has been observed in the Brassicaceae using<br />

molecular techniques. An important discovery is the<br />

synteny found between dicots <strong>and</strong> monocots. The<br />

sequence information for Arabidopsis thaliana (a<br />

Brassica species) <strong>and</strong> rice have revealed a conserved<br />

gene order across this vast evolutionary distance. It is<br />

estimated that about 75% <strong>of</strong> genes in dicot species<br />

<strong>and</strong> about 40% in monocots occur in regions that have<br />

colinearity with Arabidopsis that could be exploited by<br />

breeders.<br />

Synteny has also been discovered in Fabaceae involving<br />

soybean, peanut, mungbean, lentil, common bean,<br />

common pea, <strong>and</strong> alfalfa. Some conservation <strong>of</strong> the<br />

genomic region in this family with Arabidopsis has been<br />

established. Synteny in the Poaceae is one <strong>of</strong> the clearest<br />

to be documented. Using the rice genome as the inner<br />

circle, researchers have graphically aligned the genomes<br />

<strong>of</strong> several cereals (sorghum, maize, wheat) by colinearity<br />

into concentric circles such that colinear regions in<br />

different species lie along any radius. Rice has become a<br />

model species for comparative genomic studies <strong>of</strong> species<br />

with large genomes (e.g., wheat, sugarcane) that are<br />

difficult to study by traditional genetics. Another advantage<br />

is that the so-called “orphan crops” (e.g., millet),<br />

which would otherwise not receive significant attention<br />

as the major cereals do, can now benefit from the<br />

information obtained from rice.<br />

The applications <strong>of</strong> colinearity to plant breeding<br />

include the following:<br />

1 The prediction <strong>of</strong> the location <strong>of</strong> genes controlling a<br />

particular function in crop species.<br />

2 The facilitation <strong>of</strong> genome mapping by transferring<br />

markers from a well-mapped genome to a less studied<br />

one.<br />

3 Mapping will facilitate marker-assisted selection or<br />

map-based cloning.<br />

4 After cloning a locus <strong>of</strong> agronomic importance, colinearity<br />

may provide opportunities for accumulating<br />

alleles <strong>of</strong> that locus from other distantly related species.<br />

This collection <strong>of</strong> genes may be a source for genetic<br />

engineering applications for crop improvement.<br />

Functional genomics<br />

Once DNA sequences have been obtained, the next<br />

important task is to underst<strong>and</strong> their function.<br />

Structural genomics focuses on sequencing <strong>of</strong> the<br />

genome; functional genomics focuses on gene function.<br />

The genome is essentially a set <strong>of</strong> instructions for<br />

making proteins <strong>of</strong> various kinds. Because most genes<br />

are expressed as proteins, one <strong>of</strong> the common ways <strong>of</strong><br />

underst<strong>and</strong>ing gene function is by tracking protein<br />

expression by cells (called proteomics). Genes may provide<br />

instructions for making specific proteins. However,<br />

in the process <strong>of</strong> carrying out these instructions, additional<br />

proteins can be produced, as has already been<br />

noted. Linking gene to function is a complex undertaking.<br />

Underst<strong>and</strong>ing the genome structure alone is<br />

insufficient; it is critical to identify the proteins they<br />

encode. Many techniques have been developed for<br />

deciphering gene function.<br />

Bioinformatics in plant breeding<br />

Bioinformatics may be defined as a knowledge-based<br />

theoretical discipline that attempts to make predictions<br />

about biological function using data from DNA

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