Barley for Food and Health: Science, Technology, and Products
Barley for Food and Health: Science, Technology, and Products
Barley for Food and Health: Science, Technology, and Products
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GENETICS AND NUTRIENT COMPOSITION 57<br />
genetics <strong>and</strong> by environmental growing conditions, including farming practices<br />
such as irrigation <strong>and</strong> genetic–environment interaction. By using superior<br />
germplasm <strong>and</strong> controlling certain environmental factors such as moisture available<br />
through irrigation, a reasonably uni<strong>for</strong>m barley crop can be produced from<br />
one growing season to the next. In this chapter we provide a brief introduction<br />
to the genetics of barley, the application of genetics in breeding barley <strong>for</strong> food,<br />
<strong>and</strong> the chemical components or nutrients of the barley kernel relative to their<br />
location in the kernel.<br />
GENETICS AND NUTRIENT COMPOSITION<br />
To date, a great amount of scientific in<strong>for</strong>mation on the barley genome has been<br />
generated. This is partially due to the international importance of barley as a<br />
major crop but also to the dedication of scientists who have a genuine interest<br />
in studying this fascinating grain. The genetics of barley has probably been<br />
explored to a greater extent than that of any other cereal grain. According to<br />
Hockett <strong>and</strong> Nilan (1985), cultivated <strong>and</strong> wild barleys are major experimental<br />
organisms among flowering plants <strong>for</strong> genetic studies. The wide use of barley in<br />
genetic studies was attributed to (1) its diploid nature, (2) its low chromosome<br />
number (2n = 14), (3) the relatively large chromosomes (6 to 8 μm), (4) the<br />
high degree of self-fertility, <strong>and</strong> (5) the ease of hybridization. The reader is<br />
referred to the following <strong>for</strong> discussion of barley genetics in greater depth than is<br />
presented in this book: Smith 1951; Nilan 1964, 1974; Hockett <strong>and</strong> Nilan 1985;<br />
Wettstein-Knowels 1992; Nilan <strong>and</strong> Ullrich 1993; <strong>Barley</strong> Genetics Newsletter<br />
1996; Kleinhofs <strong>and</strong> Han 2002; <strong>and</strong> Ullrich 2002.<br />
The barley genome contains a vast number of genes that determine food quality<br />
traits, many of which have not yet been characterized. Some traits are unique<br />
to barley, but most are shared with other cereal grain species. There are several<br />
anatomical traits that are controlled by one or two genes (simply inherited),<br />
but most traits involving complex physiological processes are inherited quantitatively,<br />
being controlled by several to many genes. Genetic knowledge of many<br />
simply inherited traits dates back many years, whereas reliable data about the<br />
inheritance of the quantitative traits have been limited until relatively recently<br />
with the development of comprehensive molecular marker–based genetic maps<br />
<strong>and</strong> the technology of quantitative trait locus (QTL) analysis (Ullrich 2002). The<br />
importance of this technology was summed up by Nilan (1990): “The comprehensive<br />
molecular marker linkage maps have proved a powerful tool in barley <strong>for</strong><br />
identifying QTL loci <strong>for</strong> agronomic <strong>and</strong> quality traits, determining QTL effects<br />
<strong>and</strong> action, <strong>and</strong> facilitating genetic engineering.”<br />
In discussing genetically controlled traits in a biological system such as barley,<br />
it is customary to identify the genes(s) <strong>and</strong> the location on the chromosome<br />
involved when this in<strong>for</strong>mation is known. <strong>Barley</strong> has seven pairs of chromosomes<br />
that have been defined based on their sizes <strong>and</strong> characteristics (Nilan<br />
1964; Ramage 1985). Chromosomes 1 through 5 differ in their sizes measured