Sequence data from cDNA and genomic clones, coupled with analyses of expressed sequence tag databases, indicate that the CesA (cellulose synthase) gene family from barley (Hordeum vulgare) has at least eight members, which are distributed across the genome. Quantitative polymerase chain reaction has been used to determine the relative abundance of mRNA transcripts for individual HvCesA genes in vegetative and floral tissues, at different stages of development. To ensure accurate expression profiling, geometric averaging of multiple internal control gene transcripts has been applied for the normalization of transcript abundance. Total HvCesA mRNA levels are highest in coleoptiles, roots, and stems and much lower in floral tissues, early developing grain, and in the elongation zone of leaves. In most tissues, HvCesA1, HvCesA2, and HvCesA6 predominate, and their relative abundance is very similar; these genes appear to be coordinately transcribed. A second group, comprising HvCesA4, HvCesA7, and HvCesA8, also appears to be coordinately transcribed, most obviously in maturing stem and root tissues. The HvCesA3 expression pattern does not fall into either of these two groups, and HvCesA5 transcript levels are extremely low in all tissues. Thus, the HvCesA genes fall into two general groups of three genes with respect to mRNA abundance, and the co-expression of the groups identifies their products as candidates for the rosettes that are involved in cellulose biosynthesis at the plasma membrane. Phylogenetic analysis allows the two groups of genes to be linked with orthologous Arabidopsis CesA genes that have been implicated in primary and secondary wall synthesis.

Cellulose synthase-like CslF genes have been implicated in the biosynthesis of (1,3;1,4)-beta-d-glucans, which are major cell wall constituents in grasses and cereals. Seven CslF genes from barley (Hordeum vulgare) can be divided into two classes on the basis of intron-exon arrangements. Four of the HvCslF genes have been mapped to a single locus on barley chromosome 2H, in a region corresponding to a major quantitative trait locus for grain (1,3;1,4)-beta-d-glucan content. The other HvCslF genes map to chromosomes 1H, 5H, and 7H, and in two cases the genes are close to other quantitative trait loci for grain (1,3;1,4)-beta-d-glucan content. Spatial and temporal patterns of transcription of the seven genes have been defined through quantitative polymerase chain reaction. In developing barley coleoptiles HvCslF6 mRNA is most abundant. Transcript levels are maximal in 4- to 5-d coleoptiles, at a time when (1,3;1,4)-beta-d-glucan content of coleoptile cell walls also reaches maximal levels. In the starchy endosperm of developing grain, HvCslF6 and HvCslF9 transcripts predominate. Two peaks of transcription are apparent. One occurs just after endosperm cellularization, 4 to 8 d after pollination, while the second occurs much later in grain development, more than 20 d after pollination. Marked varietal differences in transcription of the HvCslF genes are observed during endosperm development. Given the commercial importance of cereal (1,3;1,4)-beta-d-glucans in human nutrition, in stock feed, and in malting and brewing, the observation that only two genes, HvCslF6 and HvCslF9, are transcribed at high levels in developing grain is of potential relevance for the future manipulation of grain (1,3;1,4)-beta-d-glucan levels.

A major part of the daily caloric intake of human societies around the world is derived from a diverse range of foods prepared from members of the grass family, including wheat (Triticum aestivum), rice (Oryza sativa), sorghum (Sorghum bicolor), the millets (Panicum miliaceum and Pennisetum americanum), barley (Hor- deum vulgare), and sugar cane (Saccharum officinarum). Grasses cover perhaps 20% or more of the earth's land surface (Gaut, 2002), and many of these are used as forage and fodder for the production of sheep, cattle, and other domesticated livestock. Maize (Zea mays )i s also used widely in animal feed diets, while sorghum, switchgrass (Panicum virgatum), and several other perennial grasses are attracting considerable attention as future biomass energy crops (McLaren, 2005). The grasses are noteworthy for the unusual compo- sition of their cell walls, because walls of grasses have less pectin and xyloglucan, but more heteroxylan, than walls from other higher plants. Most significantly, walls of the grasses contain as major constituents the (1,3;1,4)-b-D-glucans, which are not widely distributed outside the Poaceae. The compositions of walls from selected barley organs are shown in Table I. In many cases, constituents of cell walls in the grasses are closely linked with their widespread adoption, utility, and future potential in agricultural practice and en- ergy production. The noncellulosic polysaccharides of walls in grasses are important components of dietary fiber, which is highly beneficial for lowering the risk of serious human health conditions, including colorectal cancer, high serum cholesterol and cardiovascular disease, obesity, and non-insulin-dependent diabetes (Braaten et al., 1994; Brennan and Cleary, 2005). Con- versely, the noncellulosic wall polysaccharides from walls of cereals and grasses have antinutritive effects in monogastric animals such as pigs and poultry (Brennan and Cleary, 2005) and are often considered to be undesirable components of raw materials in the