Chapter 8 Barley Breeding History, Progress, Objectives, and Technology EUROPE Wolfgang Friedt 1 POsiTiOn Of BARLEY PRODUCTiOn AnD BREEDinG Barley (Hordeum vulgare) is one of the leading cereal crops of the world and it is clearly number 2 in Europe, next to bread wheat (Triticum aestivum). For example, in Germany, approximately 1.9 million hectares of spring and winter barley contrast to 3 million hectares of winter wheat. The remaining small grain cereals, that is, oats, rye, and triticale, together cover around 1 million hectares. On a global scale, the barley area harvested worldwide has declined from a maximum of more than 80 million hectares in the 1970s to nowadays less than 60 million hectares (Table 8.1.1). During the same time, the cultivation area in Europe went down from approximately 53 to 29 million hectares in 2008 (Table 8.1.2). This represents 51% of the world barley hectarage (56.8 million hectares). The major European barley-growing countries, France, Germany, Russia, Spain, Ukraine, and the United Kingdom, account for about three-quarters of the total annual barley grain production of about 105 million tons, representing 67% of the world production (157,644,721 t in 2008). Almost two-thirds (62%) of the total Barley: Production, Improvement, and Uses. Edited by Steven E. Ullrich © 2011 Blackwell Publishing Ltd. F European barley production comes from the European Union (EU) member states (EU 27) covering 50% of the cultivation area in Europe. The average grain yield per unit area in the EU (27) is 25% higher than the European grand mean (45.4 vs. 36.2 dt/ha; cf. FAO 2009). These igures demonstrate the outstanding importance of barley for Europe and of the European crop for global barley grain production. The high yield potential of modern cultivars and improved cultivation practices basically allow continuously increasing barley production in all areas. For example, in Germany, the average grain yield of spring barley on a farm scale has grown from less than 2.0 t/ha in the early nineteenth century to a current level of 6.1 t/ha. Over the last four decades, barley yield in Europe has been increased by 60%, in Western Europe by 80% even. The corresponding relative yield increases worldwide amount to 53% (Table 8.1.1; FAO 2009). In recent decades, the productivity of barley cultivars has risen by an annual rate of approximately 1%–2%, which is due to (i) genetic breeding progress in terms of more productive cultivars, (ii) more eficient disease and insect control, (iii) improved fertilization schemes, and (iv) improved agricultural production technology (harvest, storage, etc.) in general. Yield increases go along with a substantial improvement of malting quality and signiicant enhancement of agronomic traits leading to better yield stability, for example, due to resistances against lodging, diseases, and insects. Barley cultivars released and grown in Europe today are characterized by a considerable spectrum of resistance against 158 Ullrich—Production, Improvement, and Uses c08.indd 158 8/30/2010 8:28:50 PM Barley Breeding History, Progress, Objectives, and Technology 159 Table 8.1.1 Barley production: comparison of global production igures with those of Europe (total and western), 1966–2008 (FAO2009) World Europe (Total) Western Europe Perioda Area Harvested (M ha)b Grain Yield (t/ha)c Area Harvested (M ha)b Grain Yield (t/ha)c Area Harvested (M ha)b Grain Yield (t/ha)c 1966–1968 1976–1978 1986–1988 1996–1998 2006–2008 Change (%)d 61.6 81.1 77.6 62.0 56.2 91 1.7 2.1 2.2 2.4 2.6 153 34.3 53.4 48.7 30.9 28.7 84 2.0 2.3 2.4 2.9 3.2 160 5.1 6.3 5.3 4.3 4.0 78 3.3 3.8 4.9 5.9 5.9 179 a Averages of three consecutive years each. 106 ha. c Metric tons per hectare. d Relative values 2006–2008 versus 1966–1968. b virus diseases—particularly soil-borne yellow mosaic viruses—and fungal pathogens such as powdery mildew (Blumeria graminis), leaf rust (Puccinia hordei), or scald (Rhynchosporium secalis). The number of resistant cultivars has been signiicantly increased throughout the past few decades (Table 8.1.3). For example, a comparison of rankings for powdery mildew attack and grain yield of current winter barley cultivars with those on the market in Germany two or three decades ago clearly shows the drastic change in the varietal performance: some modern winter barley cultivars even combine a very high yield potential with a pronounced mildew resistance, which was not achievable before (cf. Friedt et al. 2000; Anonymous 2009). The main reason for strongly improved mildew resistance particularly in the spring cultivars is seen in the wide use of effective resistance genes. Various genes have been exploited throughout the decades with some emphasis on the Mla locus complex. Recently, however, the interest of breeders has focused on the mlo gene. Today, probably more than one third of the spring barley cultivars in Europe carry this gene conferring practically complete ield resistance against powdery mildew. A similar situation also exists with regard to resistances against other diseases, such as scald (R. secalis), leaf or brown rust (P. hordei), and yellow mosaic-inducing viruses (barley mild mosaic virus [BaMMV] and BaYMV), some of the most important diseases of winter barley in Europe. Whereas in the 1980s only a few cultivars showed satisfactory resistance, many cultivars expressing a high level of resistance, that is, low susceptibility (1–3 on a 1–9 scale according to the German Plant Variety Ofice) have been released more recently (Table 8.1.3). For improving rust and scald resistance, various genes have been used. For example, Rph7 provides fairly stable leaf rust resistance and has been transferred to various cultivars. New genetic sources for scald resistance have been identiied showing low disease incidence in different regions of Europe (cf. Friedt and Rasmussen 2004). Also, Pyrenophora net blotch and Ramularia leaf blotch have to be mentioned as spreading fungal diseases, which occasionally and regionally cause high infections and damage to European barley crop. The increased level of disease resistance in present-day cultivars also relects a general tendency in agricultural practice, as farmers are under a constantly increasing pressure to reduce the use of agrochemicals and pesticides for environmental reasons, thus enforcing focus on the resistance properties of varieties. Therefore, breeding for resistance in barley is a very important task on a global scale, since average yield losses worldwide of up to 30% have to be faced due to fungal and viral diseases along with insect 2 Ullrich—Production, Improvement, and Uses c08.indd 159 8/30/2010 8:28:51 PM F Barley: Production, Improvement, and Uses 160 Table 8.1.2 Major igures of barley production in European countries in 2008 (FAO2009) Country 131 Albania Austria Belarus Belgium Bosnia and Herzegovina Bulgaria Croatia Czech Republic Denmark Estonia Finland France Germany Greece Hungary Ireland Italy Latvia Lithuania Luxembourg Malta Moldova Montenegro The Netherlands Norway Poland Portugal Romania Russian Federation Serbia Slovakia Slovenia Spain Sweden Switzerland Macedonia a Ukraine United Kingdom Total or average European Union (EU) (EU 27) 132 Area Harvested (ha) 1400 185,857 612,639 55,016 22,723 222,659 61,200 482,395 717,300 141,200 585,500 1,799,300 1,961,700 150,000 332,000 181,200 330,067 131,200 332,500 9739 400F 130,179 790 50,200 128,240 1,206,560 43,100 386,706 9,420,800 92,417 213,050 19,229 3,462,400 407,700 33,112 47,351 4,167,200 1,032,000 29,157,029 14,473,752 Production (t) 25.0 52.1 36.1 77.0 33.4 39.4 44.5 46.5 47.3 24.7 36.4 67.6 61.0 25.3 44.5 69.0 37.5 23.4 29.2 54.2 40.0 27.1 15.2 61.8 41.3 30.0 23.2 31.3 24.6 37.3 41.8 39.9 32.5 44.2 61.2 34.4 30.3 59.5 3500 967,921 2,212,480 423,800 75,841 878,000 272,100 2,243,865 3,396,000 349,100 2,128,600 12,171,300 11,967,100 380,000 1,478,200 1,249,700 1,236,697 307,100 970,400 52,816 1600 353,124 1200 310,200 530,000 3,619,460 99,800 1,209,410 23,148,450 344,141 891,317 76,788 11,261,100 1,801,000 202,700 162,779 12,611,500 6,144,000 36.2 45.4 105,533,089 65,662,080 Source: http://faostat.fao.org/. a Former Yugoslav Republic of Macedonia. Major seed producers (1000 t): Belarus (250), Czech Republic (104), Denmark (132), Finland (136), France (235), Germany (322), Poland (235), Russia (3000), Spain (850), Ukraine (700), and the United Kingdom (143). The global annual seed production varied from 9.5 to 9.8 million tons in 2006–2008. Of the total global annual seed production, 6.8 (70%) is generated in Europe and 2.8 MT (29%) in the EU (27). pests. Therefore, in order to avoid necessary applications of chemicals for the prevention of respective yield losses and to ensure economic grain production, future barley breeding for disease and pest resistances should even be further intensiied. F Grain Yield (dt/ha) CURREnT sPRinG BARLEY CULTiVARs COMBininG HiGH QUALiTY AnD DisEAsE REsisTAnCE Modern European spring barley varieties trace back to European landraces from Bavaria (south- Ullrich—Production, Improvement, and Uses c08.indd 160 8/30/2010 8:28:51 PM Barley Breeding History, Progress, Objectives, and Technology Table 8.1.3 diseasesa 161 Number of winter barley cultivars listed in Germany expressing pronounced resistances to major pathogens and Year No. of Listed Cultivars Mildew Resistanta Scald Resistanta Brown Rust Resistanta Yellow Mosaic Resistanta 1980 1986 1994 2002 2009 35 46 69 84 67b 3 3 14 32 33 3 1 5 28 18 0 0 3 18 19 n.d. n.d. 19 49c 52c Source: Beschreibende Sortenliste, Bundessortenamt, Hannover, Germany (German Plant Variety Ofice). a Low susceptibility: scores 1–3 (scale 1–9: 1 = minimum susceptibility, 9 = maximum susceptibility) for powdery mildew (B. graminis), scald (R. secalis), brown rust (P. hordei), and yellow mosaic viruses (BaYMV and BaMMV). b Comprising 34 six-row and 33 two-row cultivars. c Four cultivars each are also resistant against BaYMV-2. n.d., not detected. ern Germany), Moravia (today Czech Republic), Sweden, and the United Kingdom. Cycles of cross-breeding irst focused on hybridizations among the landraces, leading, for example, to early varieties like “Isaria” (1924) and “Kenia” (1931). Later, more distant and exotic barley materials were included, for example, Hordeum laevigatum and “arabische,” two major sources of disease resistance inally leading to outstanding spring varieties like “Aramir” (released in Germany 1974) and the mlo11-carrying variety “Apex” (1983). A third cycle of cross-breeding comprised the Czechoslovakian short-straw, heavy-tillering mutant “Diamant” (Fig. 8.1.1), which also contributed to better malt quality due to high proteolytic enzyme activity (Fischbeck 3 1992, cit. Friedt et al. 2000). This led to the important cv. “Trumpf” (1973), also named “Triumph,” which acquired a wide distribution in Europe. Numerous progenies of crosses between Trumpf and representatives of the “Aramir group” became popular cultivars. Major members of this cycle were “Blenheim” (UK, 1985), “Carmen” (A, 1985), “Prisma” (NL, 1985), “Natasha” (F, 1986), and “Cheri” (D, 1987). An outstanding exponent of this variety group is “Alexis” (D, 1986), a widely grown cultivar with additional genes for mildew resistance, for example, the mlo gene of the Diamant mutant “Helena” (Fig. 8.1.1). On the other hand, the improved quality of Alexis as compared to Trumpf was probably derived from parents like 133 “Proctor” or Isaria (Fischbeck 1992, cit. Friedt et al. 2000). More recent outstanding spring barley cultivars were “Scarlett” (1995) and “Barke” (1996), both classiied at maximum malt quality (score 9) and thereby representing an improvement in comparison to Alexis (score 8). “Marnie,” a later spring malt barley cultivar, combines very high grain yield (ca. 10% superior to Barke) with improved malting quality (superior lytic parameters) and resistances against powdery mildew, leaf rust, scald, and net blotch. Marnie’s outstanding disease resistance derives from Israeli H. spontaneum used as a cross parent (J. Breun, pers. comm.). Malting quality is an economically very important but also genetically complex trait, comprising a number of different quality parameters. For a reliable description of this complex and its stability, multilocation trials are necessary, which need to be conirmed over several years of cultivation. As many as 29 out of 47 (62%) spring barley cultivars in Germany are classiied by very high malting quality (extract content) represented by a score of 8–9 (Anonymous 2009). Malt quality is deined by the following speciic criteria (desired expression): Kohlbach index, that is, enzyme activity (high), extract difference (low), extract content (high), and protein content (low). Results of EBC trials with European varieties released during the past few decades clearly show that modern, high-yielding cultivars have a drastically increased Kohlbach index and extract content Ullrich—Production, Improvement, and Uses c08.indd 161 8/30/2010 8:28:51 PM F Barley: Production, Improvement, and Uses 162 Isaria (24) Rika (51) Baladi ~~~~~ Ragusa R ~~~~~~ (29) Rika (51) CP Sulu S l ~~~~ Diamant (65) **** RiBaRi Helena (83) Proctor (52)) (5 St **** F1 1400 W409 /250 U1 1455/12714 Drossel/RiBaRi Him T253/Volla Him.T253/Volla Br. 1453e16 Aufis (74) Trumpf (73) Br. 1622d54 Br . 2629 (Piruette/ Br. 1622) Br. 2476 Br. 1400 W1455 (Helena/RiBaRi) (W409/Proctor) Georgie (77) Br. 1747 Rupee Br. 1622 W5604 (Medusa/ Diamant) Hanna Alexis (86) Br. 2730 Kym (81) Libelle (90) Amazone (86) Br. 3791 Scarlett (95) Barke (96) _____ ~~~~ ***** ( ) Year of release Selection from landraces Source of resistance Mutation fig. 8.1.1. Pedigree of major German spring barley cultivars as important parents of modern high malting quality varieties. along with a strongly reduced extract difference and a lower protein content (R.S. Schildbach, pers. comm.). These are the reasons for the observed progress regarding extract content as the major overall quality trait. However, the barley grain is not only used for feeding and beer production but also for producing malt whisky. Breeding lines have been developed, which combine all the relevant trait expressions desired for malt whisky production, for example, sprit yield, enzymatic potential (diastatic power), and low content of glycosidic nitrile. Cultivar “Optic,” derived from the cross Chad × (Corniche × Force), has been the leading spring barley in England and in Scotland. Its successors, “Publican” and “Forensic,” are considered to be particularly suitable for British (Scottish) conditions, helping growers with extra yield and also providing quality advantages to end F users, for example, distillers (Barley Malt News 5 2009; http://www.newfarmcrops.co.uk). Modern variety breeding aims at the development of cultivars with high adaptability and elasticity so that they may be successfully grown in widely different environments. Therefore, cultivars developed under continental climatic conditions as prevalent in large parts of Germany should also be competitive in other European countries and elsewhere. Such successful modern cultivars represent optimal genetic complexes determining the necessary trait combinations providing high crop stability and performance. As shown in Table 8.1.4, actual German spring barley cultivars are characterized by such optimal trait compositions, for example, “Streif” or “Tocada,” combining broad resistances against abiotic and biotic stresses with high yield poten6 tial and superior malting quality. 4 Ullrich—Production, Improvement, and Uses c08.indd 162 8/30/2010 8:28:51 PM Barley Breeding History, Progress, Objectives, and Technology 163 Table 8.1.4 Major positive traits of spring barley cultivars listed in Germany in 2009 Cultivar (Year of Release) Disease Resistance Crop Stability Grain Yield Malt Extract Annabell (1999) Braemar (2002) Average Average High High–very high High-very high High Average-high Very high High High–very high Average-high Very high Average-high High–very high Grace (2008) Marthe (2005) Quench (2006) Streif (2007) Tocada (2008) Moderate BR and NB resistance Resistant: PM (high), BR (moderate) Broad partial resistances Broad resistance, particularly to PM and NB Resistant: PM (high), scald (moderate) Broad resistance, particularly against PM and NB Moderate NB and BR resistance Low culm and ear cracking Low lodging, ear and culm cracking Low lodging, ear and culm cracking Low lodging, ear and culm cracking Low lodging, ear and culm cracking Low lodging, ear and culm cracking 134 BR, brown rust (P. hordei); NB, net blotch (Pyrenophora teres); PM, powdery mildew (B. graminis). CURREnT WinTER BARLEY VARiETiEs COMBininG HiGH YiELD AnD MALTinG QUALiTY Malta Carsten 1565 Strengs Aurea(SG) W1106 Br. 127a14 Br. 59c21 Alpha Dea Herfordia SvP 674 Malta Modern European winter barley varieties are mainly traced back to two sources of six-row barley landraces, one from The Netherlands and one from the Canadian winter barley sort “Mammut.” The variety “Friedrichswerther Berg” (1904), a progeny of hybridization between these two pools, occupies a central position in Central European winter barley breeding. Subsequent breeding cycles based on combinations of six-row winter types and both twoand six-row spring barleys, including, for example, the mildew-resistant Dalmatian landrace “Ragusa,” led to a series of important varieties such as “Dea” (1953) and “Dura” (1961). Much later, it became obvious that Ragusa also imparted resistance against BaMMV to the German winter barley gene pool. “Vogelsanger Gold,” another major variety derived from a cross of a mildew-resistant spring barley and repeated backcrosses, together with cv. Dura, were major cross parents for later widely grown six-row winter barleys like “Corona” (1980) and more recent cultivars (cf. Fischbeck 1992, cit. Friedt et al. 2000). Varieties like Dea were also parents of crosses with two-row spring barley cultivars like “Ingrid,” which inally in 1968 led to the development of cv. “Malta” (see Fig. 8.1.2), the Br. 301a W5907 Br. 269c22 Marinka (85) Labea (92) Angora (90) Tiffany (96) Jura (95) Regina (95) fig. 8.1.2. Pedigree section of important two-rowed German winter barley cultivars as major parents of modern, quality-improved two-row winter barley varieties (J. Breun, pers.comm.). major parent of subsequent widely distributed two-row winter-type malting barley cultivars such as “Sonja” (1974), “Igri” (1976), “Marinka” (1985), “Trixi” (1987), and progenies thereof. A more recent cross between the two-row winter barleys “Labea” (1992) and Marinka led to cultivars like “Jura” (1995) and “Tiffany” (1996) (Fig. 8.1.2; cf. Friedt et al. 2000). Particularly the latter two closely related cultivars rapidly gained importance in Germany in the late 1990s, since they combined a fairly good malt quality (extract content 7, equivalent to spring barley Apex) with a suficient yield performance (cf. Friedt and Rasmussen 2004). Winter malting Ullrich—Production, Improvement, and Uses c08.indd 163 8/30/2010 8:28:51 PM F Barley: Production, Improvement, and Uses 164 barley cultivation in Western Europe is still expanding at the expense of spring barley. Among the currently leading commercial winter two-row varieties are “Campanile,” “Finesse,” “Finita,” “Malwinta,” “Passion,” and “Reni.” Examples of currently leading six-row winter types are “Fridericus,” “Highlight,” “Lomerit,” and “Naomie.” It is worth mentioning that hybrid breeding is gaining importance in European winter barley breeding. For example, the highyielding six-row hybrid cv. “Zzoom” was added to the German variety list in 2008 (Anonymous 2009). BROAD-BAsED ViRUs-REsisTAnT WinTER BARLEY CULTiVARs Considerable breeding progress has been made with regard to virus resistance of European barley varieties. Besides barley yellow dwarf virus (BYDV), particularly yellow mosaic-inducing soil-borne viruses transmitted by the plasmodiophorid Polymyxa graminis have recently expanded due to the extension of cereal cultivation and infested acreage. They cause damage to barley crops in many European growing regions. Among the insect-transmitted viruses vectored by aphids, BYDV has an outstanding relevance for barley cultivation and is expected to gain importance with advancing global warming. There is no doubt that breeding of virus-resistant cultivars is the best, cost-effective, and environmentally compatible approach to prevent virus infection. Numerous sources of resistance or tolerance to the major barley viruses were identiied in the past few decades and have been used in classical breeding as well as by molecular breeding approaches (cf. Ordon et al. 2009). Since these viruses were identiied in the late 1970s, the breeding goal of high-yielding virusresistant cultivars has been achieved quite rapidly (cf. Werner et al. 2000), starting from the identiication of resistant sources within exotic cultivars or unadapted germplasm and followed by usually biparental crosses to combine resistance with other agronomic and quality traits. At the beginning of the 1990s, some important cultivars like F “Jana” were already resistant to BaMMV and BaYMV. Furthermore, extensive screening and genetic analysis enabled the release of cv. “Tokyo” ([(Fallon × 13060) × 87 – 5381 B] × Swift]; R. Hemker, pers. comm.), only a few years after the irst detection of the resistance-breaking strain BaYMV-2 in Europe. The resistance of “Tokyo” was derived from line number 13060, which has “Mokusekko 3,” the donor of rym5 in its pedigree. Due to its agronomic shortcomings, susceptibility to scald, and the relatively limited distribution of BaYMV-2, this cultivar has not achieved great acceptance by growers. However, resistant lines derived from Tokyo with better agronomic performance are being released. Winter barley is particularly attractive to growers due to its high yield potential as compared to spring types. On the other hand, due to the more recent history of this crop, the variation for disease resistance or malting quality is rather limited in comparison to spring barley. Nevertheless, great improvements have been achieved as may be recognized from Table 8.1.5. Recently listed German winter barley cultivars such as “Jade” (two-row) and “Christelle” (six-row) exhibit superior trait compositions, combining quantitative resistances against major diseases with outstanding yield and promising malting quality. From that, it can be expected that future varieties will combine their maximal yield potential with suficient stability (e.g., lodging and stress resistances) as well as suficient malting quality. Ultimately, such a trend can be expected to entirely replace spring barley with winter barley in many European barley-growing areas. BiOTECHnOLOGY-BAsED AnD MARKER-AssisTED APPROACHEs: EVOLUTiOn Of BREEDinG METHODs Acceleration of barley breeding via haploidy Combinations of different resistance genes or the introgression of novel resistances from nonadapted germplasm into adapted cultivars’ background are classically achieved by sexual Ullrich—Production, Improvement, and Uses c08.indd 164 8/30/2010 8:28:51 PM Barley Breeding History, Progress, Objectives, and Technology Table 8.1.5 165 Major positive traits of new winter barley cultivars, released in Germany in 2009 Cultivar Disease Resistance Crop Stability Grain Yield Grain Quality Christelle (six-row) Mildew, net blotch, brown rust, BYMV Mildew, brown rust, BYMV Mildew, BYMV Mildew, net blotch, scald, brown rust, BYMV Mildew Low culm cracking Very high Moderate test weight Low spike cracking Very high Very high Very high Low–moderate test weight Moderate–high test weight Low–moderate test weight Low culm and spike cracking Lodging resistant Lodging resistant, low culm cracking Low spike cracking High-very high Moderate test weight High High High test weight Moderate–high test weight Moderate-high Moderate–high test weight Low culm cracking High Moderate test weight Kathleen (six-row) Semper (six-row) Souleyka (six-row) Anisette (two-row) Canberra (two-row) Jade (two-row) Lucie (two-row) Zephyr (two-row) Mildew, BYMV Mildew, net blotch, scald, BYMV Mildew, scald, brown rust, BYMV Scald, BYMV BYMV, resistant against barley yellow mosaic viruses. recombination, that is, crosses between selected parental lines followed by phenotypic selection in the segregating offspring. In this case, the success of breeding entirely depends on extensive ield and/or greenhouse tests for resistance to the respective pathogen(s). However, since barley is damaged by many pathogens, which often show a rapid adaptation to their hosts’ resistance, breeding for resistance is a very complex task, and the identiication of desired recombinants by phenotypic selection, for example, in pedigree selection schemes, has almost reached the limits of manageability. Thus, methods of plant biotechnology like anther and microspore culture allowing the rapid production of homozygous doubled-haploid (DH) lines and cultivars were highly welcome and have been implemented into barley breeding schemes. For example, via anther culture, the spring barley cv. “Henni” (D, 1995), the two-row winter barley “Anthere” (D, 1995), and the sixrow cultivars “Uschi” (D, 1997), “Sarah” (D, 1997), “Carola” (F, 1997) and “Nelly” (D, 1998) had been released earlier than expected and became widely grown in Europe (E. Laubach, pers. comm.). It is obvious that a substantial time gain can be achieved by the application of this biotechnology step in barley breeding (Fig. 8.1.3). Consequently, the “haploid breeding method” is widely applied now and has gained great importance for barley breeding as demonstrated by the fig. 8.1.3. Generalized breeding scheme for spring and winter barley assisted by the application of the “haploid method” (E. Laubach, pers. comm.). 127 growing number of DH varieties released (E. Laubach, pers. comm.). Molecular markers and marker-assisted selection In addition, the development of molecular markers, which to some extent allow the transfer of selection steps from the phenotypic (ield) to the genotypic (laboratory) level, offers new Ullrich—Production, Improvement, and Uses c08.indd 165 8/30/2010 8:28:52 PM F Barley: Production, Improvement, and Uses 166 F opportunities for a more eficient barley breeding aiming at desired combinations of resistance, yield, and quality (Friedt et al. 2002). For example, to enhance the resistance against BYDV, Habekuss et al. (2009) used DH lines and molecular markers to combine resistance genes Ryd2 and Ryd3 and a quantitative trait locus 7 (QTL) from cv. “Post” on chromosome 2H. DH lines combining Ryd2 and Ryd2 are reported to show lower virus titer and reduced symptom expression as compared with parental materials. Another example is yellow mosaic-inducing viruses: barley cv. Taihoku A has been described as resistant to yellow mosaic viruses reported in Germany (i.e., BaMMV, BaYMV, BaYMV-2, and BaMMV-Teik). This cultivar carries the BaMMV resistance gene rym13 on chromosome 4H, which may be responsible for the resistance against the whole virus complex. By bulked segregant analysis using simple sequence repeats (SSRs) and ampliied fragment length polymorphisms (AFLPs), Humbroich et al. (2009) constructed a map with the closest marker linked to the BaMMV/BaMMV-Teik resistance locus at a distance of 1 cM. These markers are useful tools to introduce resistance to BaMMV-Teik into adapted breeding lines carrying other resistance genes, which are not effective against all BaMMV/ BaYMV strains known in Europe. For a long time, powdery mildew has been known as an important pathogen of barley in almost every growing region so that resistance against mildew is of utmost importance. To identify molecular markers, Korell et al. (2008) used a cDNA-AFLP approach to study near-isogenic barley lines differentiated by alleles of the resistance gene Mlg located on chromosome 4H. Based on the identiication of a short differential fragment (37 bp), which turned out to be part of 8 a nucleoside diphosphate kinase, a CAPS marker cosegregating with Mlg was developed. Due to its codominance, clear banding pattern, and close linkage, this marker is well suited for markerassisted selection procedures. While DNA marker analyses gain increasing importance in plant breeding and become more widely adopted in cultivar development, the capacity for high-throughput analyses at low cost is crucial for its practical application. Automation of the analysis processes is a way to meet these requirements. For this purpose, the company Svalöf Weibull AB, Sweden, has developed a fully automated PCR system. It was evaluated on 9 barley lines and was shown to be capable of analyzing up to 2200 samples per day at costs of EUR 0.24 per analysis for marker-assisted selection and quality control of genetically modiied plants (Dayteg et al. 2007). 10 Genome analysis and GM barley Today, sequencing of the barley genome has become a realistic option. Necessary steps to establish and improve genomics tools have been initiated and assembled, coordinated by an international consortium (cf. Schulte et al. 2009). A suitable reference genome sequence will be an excellent foundation for “selection with markers and advanced recombination technology” (SMART) or marker-assisted breeding, leading to future genomics-based barley improvement. In the foreseeable future, access to highly sophisticated breeding tools with a broad genetic diversity (cf. Fig. 8.1.4) as an absolute basis for gain of selection will probably become the major limiting factor for further breeding progress. Therefore, the conservation, maintenance, and evaluation of plant genetic resources are urgently needed social tasks to build the foundation for future reasonable plant breeding, crop improvement, and entirely successful agriculture. This will be even more the case if expected climate changes will lead to additional requirements of new varieties such as drought or heat resistances. Within the international network of gene banks, for example, the Institute of Plant Genetics and Crop Plant Research (IPK), Gatersleben, Germany, plays a major role not only for barley conservation but also for barley genomics (see Stein et al. 2007; Schulte et al. 2009; cf. Sato et al. 2009). Genetic transformation is not only necessary for developing new cultivars with speciically modiied traits such as improved disease and stress resistance or grain quality. Stable genetic transformation represents also an optimal Ullrich—Production, Improvement, and Uses c08.indd 166 8/30/2010 8:28:52 PM 1 2 3 4 5 6 10 cm 167 10 cm 10 cm 10 cm 10 cm 10 cm 10 cm 10 cm Barley Breeding History, Progress, Objectives, and Technology 7 8 fig. 8.1.4. Barley spike phenotypes: 1, two-rowed cv. “Barke” (wild type); 2–8, mutant phenotypes; 2, awnless; 3, intermedium erectoides; 5, brachytic; 6–8, “irregular” spikes. Source: Dr. S. Gottwald. approach to the detailed elucidation of plant gene function(s). This is considered particularly relevant in barley, a widely used experimental model plant. So, Hensel et al. (2008) presented details of the establishment of a protocol for Agrobacteriummediated gene transfer to immature embryos, enabling the highly eficient generation of transgenic barley. Advancements were made by comparing the inluence of various treatments and cocultivation conditions, leading to transgenic lines expressing the respective transgene product at low T-DNA copy numbers. In particular, the newly established protocol turned out to be useful for transformation of various spring and winter barley genotypes besides the highly amenable cv. “Golden Promise.” A very useful tool is now available for functional gene analyses as well as genetic engineering approaches in barley cultivar development. Nevertheless, it cannot be expected that transgenic barley will play a major role in European agriculture in the near future. Strong reservations against GM crops and GM food among some consumers and farmers have caused political decisions by European authorities to the disadvantage of the application of GM technology up to the present. 128 COnCLUsiOns AnD PERsPECTiVEs Barley breeding in Europe has been extremely successful throughout the last century. Cycles of cross-breeding, which irst made use of hybridization between European landraces, and later exploiting more distant germplasms providing valuable disease resistances, and inally combining high-yield and high-quality varieties, have led to highly productive modern cultivars, both in spring and winter barley. In most cases, the practical breeding techniques comprised little more than manual hybridization, careful observation, precise testing, and conscious selection. More recently, breeding programs have been enhanced by the implementation of modern biotechnology tools, like the “haploid method.” Now, it is becoming obvious that marker-assisted selection procedures—particularly PCR-based techniques combined with fast and high-throughput analysis, also called SMART breeding—will further enhance the process of selecting resistant varieties with superior agronomic performance in the future. As a consequence, existing breeding schemes including “fast track” procedures will even be accelerated and carried out more eficiently than up to now. Ullrich—Production, Improvement, and Uses c08.indd 167 8/30/2010 8:28:52 PM F Barley: Production, Improvement, and Uses 168 In addition to methodological progress, “new” traits including special qualities (”functional barley”) for alternative, for example, chemical purposes, such as high amylose and waxy barley, would potentially allow the diversiication and extension of barley cultivation and use. Furthermore, hulless barley may help to improve feed quality regarding digestability and nutrient contents, probably combined with phytase activity or low phytate content for better phosphate use. In this regard, a better understanding of gene and genome functions will enable more straightforward breeding approaches like the direct transfer of donor genes into suitable receptor cultivars. In this context, it is worth mentioning that a 11 new EMS-induced mutant population comprising 10,279 M2 individuals of two-rowed spring barley cv. Barke was described recently (Gottwald 12 et al. 2009). This new TILLING resource has potential for use in fundamental research as well as in applied breeding. ACKnOWLEDGEMEnTs Expertise on varieties and pedigrees provided by Josef Breun, Herzogenaurach, Germany, and Dr. Eberhard Laubach, Nordsaat, Germany, is gratefully acknowledged. I wish to thank Prof. Dr. Frank Ordon for detailed information regarding disease resistance and marker-assisted selection. REfEREnCEs F Anonymous. 2009. ••. In •• Bundessortenamt (ed.). Beschreibende Sortenliste 2009. Landbuch-Verlag, 13 Hannover, Germany. Behn, A., L. Hartl, G. Schweizer, G. Wenzel, and M. Baumer. 2004. QTL mapping for resistance against nonparasitic leaf spots in a spring barley doubled haploid 14 population. Theor. Appl. Genet. 108:1229–1235. Dayteg, C., S. Tuvesson, A. Merker, A. Jahoor, and A. Kolodinska-Brantestam. 2007. Automation of DNA marker analysis for molecular breeding in crops: practical experience of a plant breeding company. Plant Breed. 126:410–415. Food and Agriculture Organization (FAO) of the United Nations. 2009. Agricultural Statistics Web site, available at http://faostat.fao.org/. Friedt, W. and M. Rasmussen. 2004. ••. Symp. EUCARPIA Cereal Section, Salsomaggiore, Italy, •• ••, ••. Friedt, W., K. Werner, and F. Ordon. 2000. ••. pp. 271–279. In •• •• (ed.). Proc. 8th Int. Barley Genet. Symp., Adelaide, Vol. 1. ••, ••. Friedt, W., K. Werner, B. Pellio, C. Weiskorn, M. Krämer, and F. Ordon. 2002. Strategies of breeding for durable disease resistance in cereals. Prog. Bot. 64:138–167. Gottwald, S., P. Bauer, T. Komatsuda, U. Lundqvist, and N. Stein. 2009. TILLING in the two-rowed barley cultivar “Barke” reveals preferred sites of functional diversity in the gene HvHox1. BMC Res. Notes 2:258. doi: 10.1186/1756-0500-2-258. Habekuss, A., C. Riedel, E. Schliephake, and F. Ordon. 2009. Breeding for resistance to insect-transmitted viruses in barley—an emerging challenge due to global warming. J. f. Kulturplanzen 61:53–61. Hensel, G., V. Valkov, J. Middlefell-Williams, and J. Kumlehn. 2008. Eficient generation of transgenic barley: the way forward to modulate plant–microbe interactions. J. Plant Physiol. 165:71–82. Humbroich, K., H. Jaiser, A. Schiemann, P. Devaux, A. Jacobi, L. Cselenyi, A. Habekuss, W. Friedt, and F. Ordon. 2009. Mapping of resistance against barley mild mosaic virus-Teik (BaMMV)—an rym5 resistance breaking strain of BaMMV—in the Taiwanese barley (Hordeum vulgare) cultivar Taihoku A. Plant Breed. doi: 10.1111/j.1439-0523.2009.01721.x. Korell, M., T.W. Eschholz, C. Eckey, D. Biedenkopf, K.H. Kogel, W. Friedt, and F. Ordon. 2008. Development of a cDNA-AFLP derived CAPS marker co-segregating with the powdery mildew resistance gene Mlg in barley. Plant Breed. 127:102–104. Ordon, F., U. Kastirr, F. Rabenstein, and T. Kühne. 2009. Virus resistance in cereals: sources of resistance, genetics and breeding. J. Phytopathol. 157:535–545. Sato, K., N. Nankaku, and K. Takeda. 2009. A high-density transcript linkage map of barley derived from a single population. Heredity 103:110–117. Schmalenbach, I., J. Léon, and K. Pillen. 2009. Identiication and veriication of QTLs for agronomic traits using wild barley introgression lines. Theor. Appl. Genet. 118: 483–497. Schulte, D., T.J. Close, A. Graner, P. Langridge, T. Matsumoto, G. Muehlbauer, K. Sato, A.H. Schulman, R. Waugh, R.P. Wise, and N. Stein. 2009. The International Barley Sequencing Consortium—at the threshold of eficient access to the barley genome. Plant Physiol. 149: 142–147. Stein, N., M. Prasad, U. Scholz, T. Thiel, H. Zhang, M. Wolf, R. Kota, K.R. Varshney, D. Perovic, I. Grosse, and A. Graner. 2007. A 1000-loci transcript map of the barley genome: new anchoring points for integrative grass genomics. Theor. Appl. Genet. 114:823–839. Werner, K., B. Pellio, F. Ordon, and W. Friedt. 2000. ••. Plant Breed. 119:517–519. Werner, K., W. Friedt, and F. Ordon. 2005. Strategies for pyramiding resistance genes against the barley yellow mosaic virus complex (BaMMV, BaYMV, BaYMV-2). Mol. Breed. 16:45–55. 15 16 17 18 19 20 21 Ullrich—Production, Improvement, and Uses c08.indd 168 8/30/2010 8:28:53 PM Barley Breeding History, Progress, Objectives, and Technology Werner, K., W. Friedt, and F. Ordon. 2007. Localisation and combination of resistance genes against soil-borne viruses of barley (BaMMV, BaYMV) using doubled haploids and 22 molecular markers. Euphytica 158:323–329. NORTH AMERICA Rich D. Horsley and Bryan L. Harvey Large breeding programs for barley in North America exist in Canada, Mexico, and the United States. This portion of the chapter will focus on the programs in Canada and the United States because of the working knowledge and associations the authors have with these programs and the individuals working on them. The two largest programs in Mexico are the national program, which is the Instituto Nacional de Investigaciones Forestales, Agricolas, y Pecurias (INIFAP), and the program overseen by the International Center for Agricultural Research in the Dry Areas (ICARDA). For many years, the ICARDA program was located at the headquarters of the International Maize and Wheat Improvement Center (CIMMYT) in El Batan, Mexico. The national program has been successful in developing malting barley cultivars for Mexico, such as Esmerelda, and the ICARDA program has been successful in developing germplasm with multiple disease resistance for use around the world. In 2007, the ICARDA barley breeder responsible for Mexico and South America was moved to the ICARDA headquarters in Aleppo, Syria; however, the responsibilities of this breeder for the region did not change. 169 BARLEY PRODUCTiOn in CAnADA Barley in Canada was introduced to New France by the irst governor, Samuel de Champlain, in 1606. Canada’s irst brewery was built in Quebec city in 1668 (Metcalfe 1995). Later, the British brought two-row brewing barleys to Upper Canada. Trade between Canada and the United States thrived and brought good returns to Ontario growers for their two-row malting barley. This trade was brought to an end by U.S. protectionist measures in the form of the McKinley Tariff Act in 1890. Therefore, Ontario growers were forced to turn to feed markets for their barley. Although the two-row barley cultivars were highly suited to malting, they were low yielding, and thus improvements were required to facilitate viable barley production in that area. The Hudson Bay Company introduced cereal grains, including barley, to their trading posts throughout Northwest Canada in the 1600s in an attempt to attain food self-suficiency at these posts. However, signiicant commercial production did not begin in Western Canada until the arrival of Selkirk settlers in southern Manitoba in 1812. Barley has been a major cereal grain in Western Canada since that time. Over 90% of Canada’s barley acreage is in Western Canada (Table 8.2.1). However, it is still an important crop in Eastern Canada where growing conditions are highly variable with a multiplicity of agro-ecoregimes. Canada today can be divided into speciic barley-growing regions (Fig. 8.2.1). Below are Table 8.2.1 Canadian barley production in thousands of tons, 2004–2008 (calculated from data provided by Statistics Canada; http://www.statcan.gc.ca; veriied July 22, 2009) Year Area 2004 2005 2006 2007 2008 Mean % Canada Maritimes Quebec Ontario Manitoba Saskatchewan Alberta British Columbia 12,567 171 368 339 1278 4681 5628 90 11,678 170 340 292 603 4969 5232 72 9573 113 302 291 1035 3397 4405 31 10,984 146 308 218 1195 3945 5114 58 11,781 123 258 192 1121 4594 5448 47 11,314 146 315 266 1122 4317 5165 60 100 1 3 2 10 38 46 <1 Ullrich—Production, Improvement, and Uses c08.indd 169 8/30/2010 8:28:53 PM F Barley: Production, Improvement, and Uses 170 fig. 8.2.1. Barley production regions in Canada: 1, Maritimes; 2, Quebec; 3, Ontario; 4, Southeastern Prairies; 5, South Central Prairies; and 6, Northern Montane. speciic characteristics and breeding priorities for each region. program in Ottawa has responsibility for this area. Québec region Maritime region This region is characterized by abundant moisture, high humidity, and moderate temperatures. Acid soils are typical for this area. Foliar diseases are common and lodging is often a problem. There are no longer any barley breeding programs based in this area; thus, growers in the region rely on germplasm developed elsewhere. The Agriculture and Agri-Food Canada (AAFC) F The eastern portion of this region is similar to the Maritimes, while the western portion has commonalities with Ontario. In both areas, resistance to foliar diseases and strong straw are important selection criteria. Breeding for this region is carried out by the private sector since all of the public breeding programs have been terminated. The Ottawa program also has responsibility for this area. Ullrich—Production, Improvement, and Uses c08.indd 170 8/30/2010 8:28:53 PM Barley Breeding History, Progress, Objectives, and Technology Ontario region This region also is favored by adequate moisture; however, temperatures are considerably higher and midsummer drought is often encountered. Thus, in addition to disease resistance and straw strength, early maturity is desirable to avoid the hot dry periods of July and August. A small area of winter barley is produced in the southwestern portion of this province. This region is serviced by breeding programs at AAFC’s Ottawa station, the private sector, and a small program at the University of Guelph. Western Canada Western Canada is a vast area comparable in size to continental Europe. Clearly, such a large area has many ecozones contained within its borders. It can, however, be divided into three broad areas: the warm, moist Southeastern Prairies region; the cool Northern Montane region; and the drier South Central Prairies region. Southeastern Prairies The Southeastern Prairies region is characterized by adequate moisture levels, in most years, high temperatures and humidity in midsummer. Diseases such as stem rust (incited by Puccinia graminis f. sp. tritici [Eriks. and E. Henn.] D.M. Henderson), fusarium head blight (FHB, incited by Fusarium graminearum Schwabe), and spot blotch (incited by Cochliobolus sativus [Ito and Kuribayashi] Drechs. ex Dastur) are favored by such conditions. The AAFC’s Brandon programs play a key role in servicing this area. The University of Saskatchewan’s Crop Development Centre also provides cultivars for this area. Cultivars developed for Minnesota and North Dakota also do well here. 171 Drechslera teres [Sacc.] Shoemaker f. teres and D. teres f. maculata Smedeg.) is the most frequently found. This area is less suited to six-row cultivars and short-statured cultivars for those producers who windrow their crop prior to combining. This is prime two-row malting barley production territory. This area is the prime focus of the Crop Development Centre in Saskatoon but is also serviced by the Brandon and Lacombe programs. Northern Montane The Northern Montane region typically also has adequate moisture but has cooler temperatures and shorter growing seasons. Scald (incited by Rhynchosporium secalis [Oudem.] J.J. Davis) is a serious leaf disease in this region. Lower temperatures generally do not favor spot blotch and FHB. The Alberta Field Crop Development Centre, in Lacombe, is strategically located to service this area. The area is also serviced by the programs in Brandon and Saskatoon. Of the 11–12 million tons of barley produced in Canada each year, over 90% is grown in Western Canada. Approximately 80% is used for feed, most of that domestically. Canada’s exports of barley are typically in the range of 2–3 million tons. The largest portion of this is malting barley. Major customers for malting barley are the United States, China, Colombia, and South Africa. United States and Japan are major importers of Canadian malt, amounting to several hundred thousand tons annually. Markets for feed barley are more variable with Saudi Arabia, Japan, and several Middle Eastern countries as the most consistent buyers. Food barley consumes a very small but growing amount of barley and it constitutes an important niche for some growers. Detailed statistics can be obtained from the Canadian Grain Commission reports (http:// www.grainscanada.gc.ca/; veriied July 22, 2009). 23 BARLEY PRODUCTiOn in THE UniTED sTATEs South Central Prairies The South Central Prairies region is characterized as having reduced rainfall, high evapotranspiration rates, and frequent droughts. Foliar diseases are less common and net blotch (incited by Barley was introduced into the United States along the Eastern Coast shortly after the irst settlers arrived in the seventeenth century and Ullrich—Production, Improvement, and Uses c08.indd 171 8/30/2010 8:28:53 PM F Barley: Production, Improvement, and Uses 172 its production moved west with the growth of 24 the population (Poehlman 1985). One factor hastening the movement of barley to the west was the disease FHB. This pathogen attacks not only barley but also wheat (Triticum spp.) and maize (Zea mays L.). As maize production would move into an area, the level of FHB in the barley would increase; thus, its production would move west to areas where maize was not grown. This movement of barley west to avoid FHB continues today. The Red River Valley region of North Dakota and Minnesota was the center of six-row barley production in the United States from the 1940s to the 1990s. In the 1990s, aboveaverage precipitation and an increase in maize production in the Red River Valley provided the conditions conducive to FHB in barley. Thus, many growers in the region quit growing barley because of the high susceptibility of the crop to FHB and other market factors. The center of six-row barley production in the United States is now north central North Dakota, and new cultivars are being developed that are adapted to the more arid regions of western North Dakota and eastern Montana. The area of barley production in the United States can be subdivided into four regions: the East, Upper Midwest, West, and Southwest. The Midwest and West regions account for over 80% of the area sown to barley (United States Department of Agriculture National Agricultural Statistics Service [USDA-NASS]; available at http://www.nass.usda.gov/index.asp; veriied May 23, 2009). Speciics mentioned below on barley production and cultivars grown in the United States and in many states can be obtained from the Web site of the USDA-NASS (available at http://www.nass.usda.gov/index.asp; veriied May 23, 2009). North Dakota is the largest barley-producing state in the United States followed by Idaho and Montana (USDA-NASS). Over 90% of the barley sown in the Upper Midwest region of Minnesota, North Dakota, and South Dakota are six-row malting barley cultivars. Production in this region is typically done under dryland conditions, and the reason for the preponderance of six-row barley goes back to the origin of the barley F improvement efforts in the early 1900s. The barleys found to be best adapted to the region and to have acceptable malt quality at the time were six-row barley introductions that could be traced back to the Manchuria region of China (Rasmusson 1985). In the West region, two-row barley cultivars are typically grown, and barley for malting is often grown under irrigation. Of the early barley introduced into much of this region, two-row barley accessions from central Europe were best adapted and formed the germplasm pool from which many malting barley cultivars were developed. The area sown to malting versus nonmalting types varies by state. Idaho and Montana, the two largest barley-producing states in the region, and Wyoming sow between 65% and 80% of their barley area to malting cultivars. Other states in the region may sow less than 10% of their barley area to malting cultivars. In the East region, nonmalting winter barley cultivars are typically grown. Maryland, Pennsylvania, and Virginia sow the most barley in the region; yet, their combined production area accounts on average for only 3.6% (5-year average, 2004–2008) of the U.S. total. In the Southwest region, nonmalting spring barley is typically grown and most of the production is in the irrigated desert regions of Arizona and Southern California. The estimated total area sown to barley in this region is around 1% of the U.S. total. Unlike Canada, most of the barley produced in the United States is used domestically. From 2004 to 2007, less than 15% of the annual U.S. production of 4.8 million tons is exported, and most of the barley exported is feed barley (http:// usda.mannlib.cornell.edu/usda/fas/grainmarket//2000s/2008/grain-market-1211-2008.pdf and http://www.nass.usda.gov/ QuickStats/index2.jsp; veriied July 22, 2009). BREEDinG PROGRAMs in CAnADA Early barley improvement was carried out through introductions, initially from Europe and later from Manchuria. Selection carried out on Ullrich—Production, Improvement, and Uses c08.indd 172 8/30/2010 8:28:53 PM Barley Breeding History, Progress, Objectives, and Technology these Manchurian six-row landraces, irst at Ontario Agricultural College in Guelph and later at Macdonald College in Québec, gave rise to important cultivars in Canadian barley history. Most notable of these was OAC 21, which was widely grown throughout both Eastern and Western Canada. It was the standard for malting quality for six-row barley for many decades. Successful hybridization did not start until the 1920s; among the irst products of this program from Macdonald College was Montcalm. This cultivar was the irst Canadian malting barley developed by hybridization. It was widely adapted and high yielding and had excellent malting quality for its time. Not surprisingly, it soon dominated the malting barley acreage and held that position until the mid-1960s. In Western Canada, agricultural colleges were established in all three Prairie Provinces by 1918. As in the East, initial focus was on the introduction of cultivars from other sources. OAC 21, for example, was widely distributed by these colleges. Hybridization programs were quickly established, however, aimed at producing cultivars suitable for prairie conditions. The AAFC research stations also became active at this time. Notable among these was the station at Brandon, Manitoba, a program that has remained extremely important to this day. Barley breeding in Canada is primarily carried out by public sector agencies due to the lack of proitability of private sector breeding in self-pollinated crops like barley. Typically, less than 10% of the barley acreage is sown with pedigreed seed; thus, royalty collection is insuficient to generate proitability. These public programs have a long history of collaboration with each other and with programs in the United States. A number of annual and periodic meetings within Canada provide networking opportunities for Canadian barley scientists. The triennial North American Barley Researchers Workshop is a venue in which barley scientists from both countries exchange views and information, setting the stage for ongoing shared targets and collaboration. Speciics on the public breeding programs in Canada are presented below. 173 AAfC, Ottawa This program originally served eastern Ontario and western Quebec but has expanded its original mandate area to include the Maritime Provinces, Quebec, and Ontario where programs have been eliminated or reduced in scope. This is an enormous geographic area even though it only constitutes about 6% of the Canadian barley production. Production here is used exclusively for feed except for very small amounts of malting barley in Quebec. Thus, the program has the daunting task of servicing a diversity of production conditions. In this feed market, no premiums are paid for quality, and thus the emphasis is on yield and yield stability through disease resistance, straw strength, and nutrient use eficiency. Important diseases here include powdery mildew (incited by Erysiphe graminis DC. f. sp. hordei Em. Marchal), scald, FHB, stem rust, net blotch, Septoria speckled leaf blotch (incited by Septoria passerinii Sacc. 25 and Stagonospora avenae f. sp. triticea T. Johnson), and barley yellow dwarf virus. This program and its predecessors have provided excellent varieties, primarily of six-row cultivars, to service the needs of producers in this region. University of Guelph As indicated earlier, this program was one of the original barley improvement programs in Canada and provided cultivars like OAC 21, which gave Canada wide importance for many years. This 26 program, in addition to cultivar improvement, developed the Hordeum bulbosum method of doubled-haploid barley production that had widespread use in barley genetics research (Kasha and Kao 1970). The program now is the responsibility of one breeder for a portion of his time. The focus has narrowed to primarily six-row, spring, feed types. Objectives are high yield, disease resistance, and early maturity. Despite the reduction in resources, this program continues to play an important role for southwestern Ontario producers. AAfC Brandon The AAFC program in Brandon, Manitoba, is another program that has been historically Ullrich—Production, Improvement, and Uses c08.indd 173 8/30/2010 8:28:53 PM F Barley: Production, Improvement, and Uses 174 important to barley production in Western Canada. This program has an excellent record of producing widely adapted six-row cultivars for both feed and malting uses. Conquest and Bonanza are the best known of these cultivars and dominated the six-row malting area for many years. This station also provided valuable service by incorporating sources of resistance for a number of diseases into adapted materials. More recently, this role has expanded to tworow barley lines, and the broadly grown AC Metcalfe is an example of the important cultivars emanating from Brandon. Alberta Agriculture field Crop Development Centre, Lacombe, AB This program is the result of the evolution of AAFC’s programs at Lethbridge, Beaverlodge, and Lacombe and at the University of Alberta, all of which have been terminated (except Lacombe). Soon after its inception, a federal provincial agreement was signed and resources from both governments were contributed. The vast majority of its efforts have been devoted to developing high yield potential cultivars for Alberta’s hog and cattle industries. A number of cultivars have been released with good disease resistance and adaptation to high manure rates and adequate rainfall. University of saskatchewan Since its inception during the early part of the twentieth century, this has been the only Saskatchewan-based breeding program. Its primary focus has been the dryer portions of the prairies. The program was given a signiicant boost with the establishment of the Crop Development Centre in 1971. This recognized the importance of barley in the province where the largest acreage and the second largest production in Canada exists. The program has been preeminent in the production of highquality two-row malting barley as typiied by Harrington and a number of its successors. Harrington set the standard for quality in high enzyme activity, rapid modiication barley cultivars. It was also highly suited to the harsh environment of the dry prairies. The feed and food portion of the program pioneered a number of barley utilization areas in Canada including hulless feed barley, both two- and six-row; hulless waxy food barley; two-row forage barley; semidwarf six-row feed barley; pure amylopectin hulless, waxy food barley; hulless barley with resistance to both surfaceborne and loose smut (incited by Ustilago tritici [Pers.] Rostr.) for organic production; hulless malting barley; and low phytate barley. The program is one of the few programs to date to utilize molecular markers routinely for breeding, especially disease resistance, and for puriication of breeder seed. F supporting facilities In addition to the above breeding programs, there are several units that are important contributors to the breeding programs. Pathology support is provided by AAFC’s scientists with the lead role taken by the station in Winnipeg and support from Lacombe, Charlottetown, and Ottawa. These scientists have developed testing protocols for all of the important barley diseases in Canada and supervise and collaborate in screening for resistance. This is of course a vital resource for breeders. Plant Genetic Resources Canada, with headquarters in Saskatoon, contains Canada’s gene bank. Canada has global responsibility for barley and oat, and thus has a large collection of cultivated and wild species of barley. There is excellent collaboration between breeders and gene bank scientists in the collection characterization and utilization of barley germplasm. Malting barley is an important cash crop for Canada, and thus malting cultivars are sown on far more area than end usage would justify. Given Canada’s signiicant role in the export market for malting barley and malt, the development of high-quality potential cultivars is essential. Malting barley breeders have their own screening laboratories but rely on others for screening advanced materials. This role is carried by several organizations. The Grain Research Laboratory of the Canadian Grain Commission develops testing Ullrich—Production, Improvement, and Uses c08.indd 174 8/30/2010 8:28:54 PM Barley Breeding History, Progress, Objectives, and Technology protocols, screens advanced lines from all programs, and collates the data for cultivar registration testing evaluation. Members of the industry also contribute to this testing though their own laboratories. The Brewing and Malting Barley Research Institute (BMBRI) is an industry-based organization that coordinates collaborative testing of candidate malting lines for registration in Canada. They also assist in arranging plantscale testing post registration. The Canadian Malting Barley Technical Centre is an independent membership-based facility that provides pilot-scale malting and brewing tests on a fee for service basis. The Canadian Wheat Board is the marketing agency for Canadian malting barley and facilitates plant-scale testing of newly registered cultivars before inal acceptance in the domestic and export markets. All of these agencies located at Winnipeg collaborate to provide quality targets for breeders that can be accessed at the BMBRI Web site (http://www.bmbri.ca/; veriied May 23, 2009). These are similar to those published by the American Malting Barley Association (AMBA) in the United States and are listed elsewhere in this chapter. There are unfortunately no similar networks for feed and food barley; thus, breeders are forced to set up any necessary testing on a case-by-case basis. BREEDinG PROGRAMs in THE UniTED sTATEs Barley breeding in the United States is largely done by public institutions located in the northern tier of states. Institutions with breeders working solely on barley include Montana State University, North Dakota State University (NDSU), Oregon State University, the University of California, Davis, the University of Minnesota, Washington State University, and the United States Department of Agriculture Agricultural Research Service (USDA-ARS) at Aberdeen, Idaho. Public programs with breeders working on barley as well as other small grains are located at the University of Georgia, University of 175 Maryland, University of Nebraska, Utah State University, and Virginia Polytechnic Institute and State University (VPI). All of these programs have development of wheat (Triticum aestivum L.) cultivars as their top priority and have much smaller efforts in barley. The programs in California, Minnesota, Montana, North Dakota, and Washington work primarily on developing spring barley cultivars; the programs in Georgia, Maryland, Nebraska, Oregon, and Virginia work almost exclusively in developing winter barley cultivars; and the USDA-ARS program in Idaho works on developing both spring and winter barley cultivars. In addition, the Minnesota, North Dakota, and Idaho USDA-ARS programs work primarily on developing malting barley cultivars. Unlike many areas of the world where only malt made from two-row barley is used for brewing, the brewing industry in the United States uses substantial amounts of malt made from six-row barley. The breeding program at the University of Minnesota focuses on the development of six-row barley cultivars, whereas the NDSU and USDA-ARS programs work on the development of two- and six-row cultivars. The public programs in California, Georgia, Maryland, Nebraska, Utah, and Virginia develop primarily nonmalting barley cultivars, and the Montana, Oregon, and Washington programs work on barley for malting and nonmalting uses. The VPI, Idaho USDA-ARS, Oregon, and Washington breeding programs also work on hulless barley for food, feed, and industrial uses. The program at VPI is particularly working on hulless barley for ethanol production in collaboration with the USDA-ARS Eastern Regional Research Center at Wyndmoor, Pennsylvania. Breeding in the private sector is dominated by three companies that develop cultivars for production across more than one barley-growing region in the United States. The companies and the headquarters of their breeding programs are Busch Agricultural Resources, LLC (BAR, a member of Anheuser-Busch InBev), located in Fort Collins, Colorado; MillerCoors, located in Burley, Idaho; and Westbred, LLC, headquartered in Bozeman, Montana. The BAR and MillerCoors programs breed malting barley, Ullrich—Production, Improvement, and Uses c08.indd 175 8/30/2010 8:28:54 PM F Barley: Production, Improvement, and Uses 176 whereas Westbred develops feed, hulless food, and forage-type barley cultivars. The MillerCoors program develops cultivars almost exclusively for use within their company, whereas BAR develops cultivars that are used by brewers in addition to Anheuser-Busch InBev. Breeding priorities in the United states Malting The choice of parents to use for crossing is critical for developing acceptable barley cultivars because barley cultivars often are adapted to speciic production regions and are designed for particular uses, such as malting and brewing. Designation of a cultivar as a “malting-type” is assigned by a particular company that would use the cultivar or by an organization such as AMBA, located in Milwaukee, Wisconsin. AMBA is a nonproit trade association composed of dues paying malting and brewing companies. A primary goal of AMBA is to ensure an adequate supply of high-quality two- and six-row malting barley for the malting and brewing industries in the United States (http://www.ambainc.org/about/bh.htm; veriied May 23, 2009). This is accomplished, in part, by providing grant funds to public breeding programs to support their cultivar development efforts. AMBA also coordinates pilot-scale and plant-scale evaluation programs of advanced breeding lines or cultivars to determine their suitability for their member companies. These two programs will be discussed in greater detail later in the chapter. Evaluation of malt quality of breeding lines prior to entering into AMBA’s testing programs is done by the USDA-ARS Cereal Crops Research Unit in Madison, Wisconsin, and in limited cases in industry laboratories. Evaluation of breeding lines developed by the public breeding programs at a single site by the USDA-ARS allows AMBA’s members and other stakeholders to make comparisons between lines that are all malted and evaluated under similar conditions. Malting barley in the United States is unique from most other crops in the country in that it is marketed and stored on an identity preserved basis. Malting barley cultivars are kept segregated F by cultivar, and often times a cultivar is additionally segregated based on production area or by a speciic quality trait such as grain protein level. Segregation is done because cultivars used for malting and brewing must meet a long list of speciic criteria before they are accepted. In addition, malting is done by cultivar and often by cultivar grown at a speciic location, and many beer brands have speciic compositions of different barley cultivars in their brewing blend. A unique feature of malting barley in North America is that the life of cultivars can easily be 10 years or more. Examples of this are Larker, Morex, and Robust six-row barleys, and Klages and Harrington two-row barleys. Larker, Morex, Robust, and Klages were developed in the United States, and Harrington was developed at the University of Saskatchewan in Saskatoon, Canada. Guidelines for speciic quality traits, which include measurements on barley and malt, are provided to the public breeding programs by AMBA (Table 8.2.2). Quality traits that receive the most attention in malting barley breeding programs are barley grain protein and kernel plumpness, and malt extract, enzymatic activity (α-amylase and diastatic power), wort protein, measures of carbohydrate modiication (β-glucan content and wort viscosity), and protein modiication (Kohlbach index) in the malt. A detailed description of these malt parameters and others can be found in Kunze (2004). Because of the long list of parameters that must be met, development of malting barley cultivars is typically done using crosses between parents that already have acceptable quality. Shortfalls in malt quality of either parent usually appear in the progeny. This need to make crosses between parents with acceptable quality has led to very narrow germplasm bases because variability already has been severely restricted by adaptation to speciic production regions. For example, Horsley et al. (1995) stated that in the early 1990s, all cultivars developed by six-row barley breeding programs in the Upper Midwest United States and the eastern Prairie Provinces of Canada could be traced back to 15 accessions obtained in the early 1890s. However, even within the narrow germplasm bases of malt barley programs, gains Ullrich—Production, Improvement, and Uses c08.indd 176 8/30/2010 8:28:54 PM Barley Breeding History, Progress, Objectives, and Technology Table 8.2.2 Barley and malt quality speciications provided to barley breeders in the United States by the American Malting Barley Association, Inc., Milwaukee, Wisconsin, in May 2008 (http://www.ambainc.org/ni/Guidelines.pdf; veriied 22 July 2009) 135 Barley factors Plump kernelsa Thin kernelsb Germinationc Protein Skinned and broken kernels Malt factors Total protein On 7/64” sieve Two-Row Barley Six-Row Barley >90% <3% >98% ≤13% <5% >80% <3% >98% ≤13.5% <5% ≤12.8% >70% ≤13.3% >60% Measures of malt modiication β-Glucan (ppm) <100 Fine–coarse extract <1.2 difference Kohlbach index 40%–47% Turbity (NTU) <10 Viscosity (absolute <1.50 cP) 136 Congress wort Soluble protein Extract (ine grind db) Color (°ASBC) Free amino nitrogen Malt enzymes Diastatic power (°ASBC) α-amylase (20° DU) <120 <1.2 42%–47% <10 1.50 4.4%–5.6% >81.0% 5.2%–5.7% >79.0% 1.6–2.5 >190 1.8–2.5 >200 >120 >140 >50 >50 177 malting barley. However, feed quality attributes are being developed for several classes of livestock. Using marker-assisted selection (MAS), cv. Valier was developed to incorporate excellent agronomic performance and improved feed characters for beef cattle (Blake et al. 2002). Incorporating disease resistance genes becomes especially important because application of expensive fungicides to feed barley can be cost prohibitive. Breeders developing hulless barley for human consumption look to develop cultivars that are high in soluble iber to reduce the risk of cardiovascular disease (Behall et al. 2004). The trait that receives the most attention is breeding for increased grain β-glucan content. Breeders developing hulless barley for the poultry feed market work to develop cultivars with reduced grain βglucan content (Newman et al. 1987). Unlike the central USDA-ARS-CCRU laboratory available 27 for testing malt quality, there is no central laboratory available for determining the feed or food quality of breeding lines. Each of the breeding programs is responsible for determining the quality of their lines themselves. Likewise, there is no single organization such as AMBA that is providing desired quality characteristics to breeders for feed and food barley. Resistance to diseases still are being made. Rasmusson and Phillips (1997) theorized that gains are made in the narrow germplasm bases due to de novo variation and elevated epistasis. In addition, for an excellent review on gains and diversiication in barley obtained through breeding, the reader is encouraged to read Fischbeck (2003). Feed and food In developing cultivars designed for livestock feed, the choice of parents is of less importance because growers are usually not paid premiums for speciic quality parameters as is done for A high priority in many breeding programs is the development of cultivars with resistance to one or more pathogens. Because barley is considered a low-input crop, the easiest and most cost-effective method of disease control is growing resistant cultivars. The list of barley diseases caused by bacteria, fungi, and viruses is plentiful (Mathre 1997). Good sources of genes for resistance to diseases and other pests are often unadapted accessions or the wild ancestors of barley, such as Hordeum vulgare subsp. spontaneum. The USDAARS National Small Grains Collection (NSGC) in Aberdeen, Idaho, is the part of the USDAARS’s National Plant Germplasm System that has a large collection of cultivated and wild ancestors and relatives of barley and other small grain cereals from around the world. In June 2009, the Ullrich—Production, Improvement, and Uses c08.indd 177 8/30/2010 8:28:54 PM F Barley: Production, Improvement, and Uses 178 USDA-ARS-NSGC had 27,925 accessions of H. vulgare subsp. vulgare, 1507 accessions of H. vulgare subsp. spontaneum, and over 440 additional Hordeum accessions (http://www.arsgrin.gov/cgi-bin/npgs/html/site_holding.pl?NSGC; veriied July 22, 2009). Another important utility that the USDAARS provides to barley breeders, pathologists, and other stakeholders is tracking and identiication of different rust pathogens in the United States, and screening and identiication of new sources of resistance to new rust pathotypes. This role is overseen by the USDA-ARS Cereal Disease Laboratory (CDL) located in St. Paul, Minnesota. Each year, the USDA-ARS-CDL publishes a series of bulletins that report on the current situation of several cereal rust diseases in the United States (http://www.ars.usda.gov/ Main/docs.htm?docid=9757; veriied July 22, 2009). Each barley-growing region of the United States has different disease proiles and priorities when breeding for disease resistance. In the Upper Midwest United States, the high-priority prevalent diseases are wheat stem rust, FHB, spot blotch, and the net form of net blotch (incited by Pyrenophora teres Drechs.). All cultivars developed for this region must have resistance to the prevalent pathotypes of wheat stem rust due to the devastation this disease can cause. Since 1993, major breeding efforts have been ongoing in the region to develop cultivars with improved resistance to FHB and accumulation of the mycotoxin deoxynivalenol (DON) that is produced by the causal organism. The best sources of resistance to FHB and DON accumulation being used by Midwestern U.S. breeding programs have originated from China, Japan, and Switzerland (Prom et al. 1996; Urrea et al. 2005). The list of foliar diseases causing signiicant losses in the western and southwestern United States is much smaller than that in the other regions due to the much drier environment. The disease receiving the most attention since the late 1980s is stripe rust, incited by Puccinia striiformis f. sp. tritici. In 1975, a pathotype capable of attacking most cultivars in the United States spread F from Europe to Colombia, and the irst disease caused by this pathotype was irst seen in the United States in winter 1991 in Texas (Roelfs et al. 1992). From Texas, the disease spread quickly to California, Idaho, Montana, Oregon, and Washington (Chen 2004). Because breeders and pathologists were aware that this disease would eventually reach the United States, they were proactive and were able to identify and incorporate resistance into their germplasm (Roelfs and Huerto-Espino 1994; Chen 2004). Complexes of 28 diseases that are more problematic in the western United States are a series of different root and crown rots (e.g., rhizoctonia, pythium, and fusarium). The effects of this group of diseases are especially noticed in reduced tillage and direct seed systems as well as during dry years or in dry areas of the ield such as hilltops because roots are unable to uptake suficient water (Cook and 29 Veseth 1991). In the eastern United States, the predominant diseases are barley leaf rust (incited by Puccinia hordei Otth.), powdery mildew, and barley yellow dwarf virus. A dificulty in breeding for barley leaf rust is that multiple leaf rust resistance genes have been overcome by changes in pathogen races of P. hordei. Resistance to insects and nematodes In the United States, the aphids bird cherry-oat aphid (Rhopalosiphum padi [L.]), green bug (Schizaphis graminum [Rondani]), and Russian wheat aphid (RWA) (Diuraphis noxia [Kurdjumov]) are the major insect pests of barley (Porter et al. 1999). Breeding for resistance to RWA receives the most attention, and damage from this pest occurs predominantly in the intermountain west states of Colorado, Idaho, Montana, and Wyoming. A large screening effort for identiication of accessions in the USDAARS-NSGC with resistance to RWA was conducted in the glasshouse by USDA-ARS scientists at Stillwater, Oklahoma (Webster et al. 1991). Over 100 lines with RWA resistance were identiied and germplasm lines were rapidly developed and made available to breeders (Mornhinweg et al. 1995, 1999). In a collaborative effort Ullrich—Production, Improvement, and Uses c08.indd 178 8/30/2010 8:28:54 PM Barley Breeding History, Progress, Objectives, and Technology between USDA-ARS researchers at Stillwater, Oklahoma, and Aberdeen, Idaho, additional RWA germplasm lines and cultivars have been developed. The cultivar Burton (Bregitzer et al. 2005) is an example of this successful collaboration of ARS scientists with university experiment station researchers in the region. Three nematode genera (Heterodera, Meloidogyne, and Pratylenchus) are known to cause economic damage to barley (Mathre 1997). However, breeding for resistance to nematodes in the United States is not a priority. Resistance to abiotic stresses As stated earlier, barley is considered by growers to be a low-input crop; thus, breeding for resistance to abiotic stresses is a high priority for many barley breeders. A partial list of these stresses includes drought and looding, high and low temperatures, mineral deiciencies and toxicities, poor soil tilth (structure), and preharvest sprouting (PHS). A dificulty when breeding for abiotic stresses is that plants often are exposed to multiple stresses at the same time, and it is not unusual that common responses to the stresses are provoked in plants (Langridge et al. 2006). Thus, breeding for abiotic stresses generally is considered more dificult than breeding for diseases or other pests. Internationally, functional genomic technologies are being used to gain a better understanding of how plants respond to abiotic stresses (Langridge et al. 2006). However, establishment of effective breeding strategies based on this information will be challenging. sELECTinG sOURCEs Of REsisTAnCE TO ABiOTiC AnD BiOTiC sTREssEs in CAnADA AnD THE UniTED sTATEs After sometimes exhaustive searches, genotypes, including cultivars, unadapted genotypes, landraces, and wild barley accessions that are resistant or tolerant to most economically important biotic and abiotic stresses usually can be found (Ullrich et al. 1995). However, sources of the resistance genes are often poorly adapted to their 179 intended region, and desirable genes may show linkage drag to undesirable traits. When a new disease or pest arises or resistance breaks down in currently grown cultivars, breeders can develop improved cultivars in a short time if they have been taking a proactive approach to the problem. Examples of this approach include work done by USDA-ARS researchers on RWA and barley stripe rust beginning in the late 1980s. The length of time needed to develop improved cultivars also is reduced if the sources of resistance can be found in adapted germplasm. When a loss of resistance is unanticipated, it can easily take more than 10 years to incorporate new resistance into acceptable cultivars. In the case of malting barley, this time frame is grossly underestimated because stringent quality parameters must be maintained. This is the current case for developing improved cultivars with resistance to FHB and DON accumulation. Breeding for these problems in the Midwest United States began in 1993, and the irst breeding lines that combined acceptable agronomic performance, disease resistance, and malt quality did not enter advanced testing by the malting and brewing industries until 2008. Not only is barley considered a low-input crop by producers, but many barley breeders are expected to develop improved cultivars using limited inancial and physical resources. The tworow barley breeding program at NDSU in the United States is an example where the existing barley genetic resources were poorly adapted to the production area when the program was started in the early 1970s (Horsley et al. 2008). Improved two-row lines were developed by making crosses between unadapted two-row germplasm and adapted Midwest U.S. six-row malting cultivars. The most important selection criteria in early generations were low grain protein (Foster et al. 1967) and high kernel plumpness. The irst cultivar released from this program was Bowman (Franckowiak et al. 1985). Additional two-row cultivars released by this program are Stark, Logan, Conlon, Rawson, and Pinnacle. Conlon was added to the list of cultivars recommended for production as a malting barley by AMBA in 2000. Ullrich—Production, Improvement, and Uses c08.indd 179 8/30/2010 8:28:54 PM F Barley: Production, Improvement, and Uses 180 BREEDinG METHODs AnD TECHniQUEs in CAnADA AnD THE UniTED sTATEs F No two breeding programs use the same breeding scheme. A detailed example of a breeding scheme used to develop malting barley cultivars at NDSU in the United States can be found in Horsley et al. (2008). The scheme uses a modiied-pedigree breeding method and off-season nurseries to reduce the length of time needed to develop new cultivars by up to 3 years. Variations to the scheme above can be made at any stage to accommodate speciic resources and breeding goals. For example, doubled-haploids or MAS based on DNA markers can be used to shorten the length of time needed to develop new cultivars by several years. These methods, as well as barley transformation, are described in greater detail in other chapters. Off-season or glasshouse nurseries also can reduce the length of time needed to develop new cultivars by providing for two or more ield generations per year. Locations used for offseason nurseries by breeders in Canada and the United States include Arizona and California in the southwestern United States, Argentina, and New Zealand. Finally, nonmalting or feed barley cultivars typically are developed as much as 3–4 years faster than malting barley cultivars because 30 extensive quality testing is not required. Currently, MAS is gaining wider use and research continues to identify opportunities to incorporate this promising method. As stated earlier, MAS is being used at the University of Saskatchewan disease resistance breeding and puriication of breeder seed. In the United States, MAS is being used for identifying lines for resistance to such diseases as barley stripe rust, FHB, and Septoria speckled leaf blotch. A 4-year (2006– 2010) multistate collaborative project in the United States titled the Barley Cooperative Agricultural Project (CAP), funded by the USDA Cooperative State Research, Extension, and Education Service (CSREES), has as its primary goal to identify single nucleotide polymorphism (SNP) markers associated with economically important traits using association genetics methods. Information from this project will be used to generate PCR primers pairs for MAS at 31 the USDA-ARS high-throughput molecular marker laboratories located in Fargo, North Dakota, or Pullman, Washington. These laboratories are currently screening breeding lines using molecular markers for traits such as FHB and barley stripe rust resistance. COLLABORATiOn BETWEEn BREEDinG PROGRAMs in THE UniTED sTATEs AnD AMBA fOR DEVELOPinG MALTinG BARLEY CULTiVARs A long history of collaboration has existed between malting barley breeders in the Canadian and U.S. breeding programs and members of the malting and brewing industries. Development of improved malting barley cultivars is dependent on barley breeders knowing the desired levels of the numerous malt quality parameters required by maltsters and brewers. Communication of the maltsters’ and brewers’ needs is conveyed to the breeders through the Web and periodic meetings. As described earlier, AMBA provides breeders in the United States with baseline criteria that must be met by new cultivars before they will be recommended for use by their members. This collaboration is accomplished through industry testing of the malting and brewing qualities of advanced breeding lines and new cultivars that is coordinated by AMBA. This evaluation, referred to as pilot-scale and plant-scale evaluations, requires a minimum of 4 years of testing. Before a cultivar can be added to the “AMBA List of Recommended Malting Barley Varieties,” it must be rated as satisfactory in 2 years of pilot-scale evaluation and in 2 years of plant-scale evaluation. Breeding programs participating in the AMBA evaluation programs include the public programs in Idaho, Minnesota, Montana, North Dakota, Oregon, and Washington, and BAR. AMBA pilot-scale evaluation Based on favorable agronomic and malt quality characteristics, breeders are allowed to submit up to eight lines (four six- and four two-row lines) to Ullrich—Production, Improvement, and Uses c08.indd 180 8/30/2010 8:28:54 PM Barley Breeding History, Progress, Objectives, and Technology AMBA for pilot-scale evaluation. Seed for this evaluation is obtained from two of the best six locations of large increase plots that are grown speciically for AMBA’s tests by collaborating breeders. For this evaluation, 17 kg of each entry is divided among the malting and brewing members of AMBA, and malt quality is compared 32 to the checked cultivars. No brewing evaluation is done as part of the AMBA pilot-scale evaluation. Lines must be rated as satisfactory in 2 of 3 years of pilot-scale evaluation before they can be considered for advancement to the AMBA Plant Scale Evaluation Program. 181 the list in the 1980s (Azure, B1601, and Robust) and 1990s (Excel, Foster, and Stander) for production in the Midwest United States. All six of these cultivars are six-row. In the 2000s, seven new cultivars (Conlon, Drummond, Lacey, Legacy, Rasmusson, Stellar-ND, and Tradition) have been added to the list for production in the Midwest United States. Conlon is the only tworow cultivar. In addition, German cv. Scarlett was added to the list in 2007 based on input from AMBA’s malting members. CULTiVAR RELEAsE in CAnADA AMBA plant-scale evaluation Typically, no more than one six-row and one tworow lines from a breeding program are allowed in the plant-scale evaluation program at any one time. The plant-scale evaluation program includes evaluation of malting and brewing quality. Grain of the candidate for plant-scale evaluation is grown on approximately 250 ha in the region where it is intended to be grown. The production is done by local producers with the goal of producing a total 435–655 t of grain. Malt for plantscale brewing evaluation is typically produced by one of AMBA’s malting members. In subsequent years of the AMBA Plant Scale Evaluation Program, a different maltster will be responsible for producing the malt. This process ensures that multiple maltsters have input during the AMBA plant-scale evaluation process. The methods of brewing evaluation of plantscale candidates are different for each brewing company. Some companies may use small inclusions of the candidate line in their brewing blend, while others may use inclusion rates of 50% or more. Ultimately, each brewing member wants to identify any deiciencies the candidate line may have during the malting and brewing processes and in the inished malt and beer. Even though the addition of a cultivar to the AMBA List of Recommended Malting Barley Varieties requires support from only one of AMBA’s member companies, very few cultivars get added to the list because of the stringent quality requirements. For example, only three cultivars were added to Cultivar registration is compulsory in Canada for a number of ield crops including barley. Registration is administered by the Variety Registration Ofice of the Canadian Food Inspection Agency (CFIA; http://www. inspection.gc.ca/english/plaveg/variet/ vartoce.shtm; veriied July 22, 2009). This ofice recognizes and relies on registration recommending committees to develop testing procedures, to evaluate data, and to make recommendations on the suitability of candidate cultivars for registration. In Western Canada, for example, the recognized committee is the Prairie Registration Recommending Committee for Oat and Barley. Details of testing protocols are posted on the Web site http://www.pgdc.ca/ (veriied July 22, 2009). Membership on this committee includes scientists from government, universities, and the private sector; industry specialists; provincial government specialists; and representatives of farmer organizations. This committee sets the standards for testing and tests are conducted under its auspices. It also determines its operating procedures subject to ratiication by the CFIA. The committee is divided into three evaluation teams for data evaluation purposes: the agronomy team evaluates ield performance; the disease team evaluates disease resistance; and the quality team evaluates suitability for end-use processing and manufacturing. The composition of these yield trials, including checks, is approved by the committee annually and administered by a test coordinator. There are three yield trials: the 33 Ullrich—Production, Improvement, and Uses c08.indd 181 8/30/2010 8:28:54 PM F Barley: Production, Improvement, and Uses 182 Western Six-Row Barley Cooperative, the Western Two-Row Barley Cooperative, and the Hulless Barley Cooperative. Entry into these tests requires six station years of data from tests in Canada with appropriate checks. Cooperative preregistration trials are conducted by breeders and other cooperators at a number of locations, and candidates are evaluated over a 2-year period before consideration for registration. Data are collected on agronomic performance including yield, lodging, maturity, seed weight, and so on. In the case of malting barley, an additional year of collaborative malting tests is required. Quality evaluation is conducted on seeds from selected sites of the yield trials. Micro malts are produced and wet chemistry analyses are conducted by the Grain Research Laboratory. In parallel to these tests, a collaborative malting trial is coordinated by the BMBRI. Promising entries from irst-year cooperative testing plus successful entries from previous collaborative testing are grown in separate plots, with commercial checks, at several sites across Western Canada. Seed from selected sites is malted and analyzed by the Grain Research Laboratory and by industry laboratories. Disease resistance data are collected on the yield trials when infections justify and separate nurseries are grown at various sites. For example, a stem rust nursery is grown at the Winnipeg AAFC station; a scald nursery is grown at Lacombe; and an FHB nursery is grown at the Brandon AAFC station. Coordinators are assigned for each disease and are responsible for test coordination and data collation. When considering candidate cultivars, each evaluation team passes expert judgment on their area of expertise. A inal judgment is then made by the committee in plenary on the basis of all the data as a total package. Thus, these experts now consider all data as opposed to their respective discipline data so they are able to balance weaknesses in one area with strengths in another to reach a inal recommendation to either support or not support the application for registration. This information is transmitted to the registration ofice and a inal decision is made there, usually taking the advice of the committee. F Postrelease adaptation and commercial acceptance testing in Canada Adaptation testing generates more information on cultivar adaptation than is typically necessary for a breeder to feel comfortable in releasing a new cultivar and for cultivar registration purposes. However, this information is found to be very useful by many producers who use it to assist in their decision to adopt new cultivars. The type of testing in these yield trials is similar to advanced generation yield trials but with a greater number of stations involved and less data are recorded. These trials are usually conducted by agencies other than breeding organizations. Commercial acceptance testing is necessary for malting cultivars and for special use food or feed cultivars. The malting and brewing industry, for example, will not take on new cultivars without such testing, in which they support and actively participate. Typically, they have seen several years of micromalting data on new cultivars but no brewing information. Thus, commercial scale ields are contracted and selected barley is malted and brewed under commercial conditions. Most maltsters and brewers want to see at least 2 years of successful malting and brewing before they will begin to incorporate a new cultivar into their blends. Again, these tests, while critical to the success of a cultivar, are beyond the purview of the breeder other than to provide seed sources and to take a keen interest in the process. seed production in Canada Pedigreed seed production in North America is among the best in the world. The role of the professional seed grower is vital to the success of any new cultivar. They increase the seed that the commercial farmer uses and maintain the genetic integrity of the cultivar in doing so. The Canadian Seed Growers Association (CSGA) and the CFIA are coregulators of the pedigreed seed certiication system. The CSGA sets the standards and the CFIA oversees production to ensure that standards are met. For barley, there are the following classes of seed: Ullrich—Production, Improvement, and Uses c08.indd 182 8/30/2010 8:28:54 PM Barley Breeding History, Progress, Objectives, and Technology 1. 2. 3. 4. 5. Breeder seed, produced under the supervision of a recognized breeder; Select seed, produced from breeder seed by qualiied select growers; Foundation seed, produced by registered seed growers; Registered seed, and Certiied seed. The reader is referred to the CSGA Web site, http://www.seedgrowers.ca (veriied July 22, 2009), for more detailed information. CULTiVAR RELEAsE AnD inTELLECTUAL PROPERTY issUEs in THE UniTED sTATEs Unlike many areas of the world, there is no formal process in the United States administered by the federal government for determining if a line can be released as a named cultivar for commercial production. Each university or private breeding program has its own criteria and process for releasing cultivars. While an experimental line is usually evaluated in one or more Cooperative Regional Yield Trials coordinated by the USDAARS, this is not required before it is released as a named cultivar. The determination of whether a cultivar will be added to the AMBA List of Recommended Malting Barley Varieties is described previously. In general, most cultivars from the public programs are released to seed grower organizations (e.g., Crop Improvement Associations) in their states. The breeder and foundation seeds of these releases are handled by the State Crop Improvement Association or by the university or state agricultural experiment station that released the cultivar. Production of the registered and certiied seed classes is handled by growers often contracted by seed companies, but ields and seed are inspected and certiied by the State Crop Improvement Association or State Department of Agriculture. Production of the foundation seed class for cultivars released by private companies is not consistent. Some companies may contract the production of the foundation class themselves 183 with local growers or they may contract a university to handle this production. Protection of intellectual rights for public and private breeders can be done by obtaining a plant variety protection (PVP) certiicate administered by the Plant Variety Protection Ofice of the USDA Agricultural Marketing Service (http:// www.ams.usda.gov/AMSv1.0/; veriied July 28, 2009). Less commonly, utility patent protection may be obtained through the United States Patent and Trademark Ofice of the United States Department of Commerce (http://www.uspto. gov/main/patents.htm; veriied July 28, 2009). The decision to obtain a PVP certiicate or a utility patent for a new barley cultivar, especially in the public sector, is not consistent across universities, agencies, programs, or even cultivars nor is the institution of royalties for growing protected cultivars. REfEREnCEs Behall, K.M., D.J. Scholield, and J.G. Hallfrisch. 2004. Lipids signiicantly reduced by diet containing barley compared to whole wheat and brown rice in moderately hypercholesterolemic men. JAMA 23:55–62. Blake, T., J.G.P. Bowman, P. Hensleigh, G. Kushna, G. Carlson, L. Welty, J. Eckhoff, K. Kephart, D. Wichman, and P.M. Hayes. 2002. Registration of ‘Valier’ barley. Crop Sci. 42:1748–1749. Bregitzer, P., D.W. Mornhinweg, R. Hammon, M. Stack, D.D. Baltensperger, G.L. Hein, M.K. O’Neill, J.C. Whitmore, and D.J. Fielder. 2005. Registration of Burton barley. Crop Sci. 45:1166–1167. Chen, X. 2004. Epidemiology of barley stripe rust and races of Puccinia striiformis f. sp. hordei: the irst decade in the United States. Cereal Rusts Powdery Mildews Bull. ••:••– ••. Available at http://www.crpmb.org/2004/1029chen/ 34 (accessed August 17, 2008). Cook, R.J. and R.J. Veseth. 1991. Wheat Health Management. APS Press, St. Paul, MN. Fischbeck, G. 2003. Diversiication through breeding. pp. 29–52. In R. von Bothmer, T. van Hintum, H. Knüpffer, and K. Sato (eds.). Diversity in barley (Hordeum vulgare). Elsevier Science B.V., Amsterdam, The Netherlands. Foster, A.E., G.A. Peterson, and U.J. Banasick. 1967. Heritability of factors affecting malting quality of barley, Hordeum vulgare L. Emend. Lam. Crop Sci. 7:611–613. Franckowiak, J.D., A.E. Foster, V.D. Pederson, and R.E. Pyler. 1985. Registration of Bowman barley. Crop Sci. 25:883. Horsley, R.D., P.B. Schwarz, and J.J. Hammond. 1995. Genetic diversity in malt quality of North American six-row spring barley. Crop Sci. 35:113–118. Ullrich—Production, Improvement, and Uses c08.indd 183 8/30/2010 8:28:54 PM F Barley: Production, Improvement, and Uses 184 35 36 37 38 F Horsley, R.D., J.D. Franckowiak, and P.B. Schwarz. 2008. Barley breeding. In M.J. Carena (ed.). Handbook of Plant Breeding: Cereals, Vol. 3. Springer, Berlin (in press). Jorgensen, J.H. 1992. Discovery, characterization and exploitation of Mlo powdery mildew resistance in barley. Euphytica 63:141–152. Kasha, K.J. and K.N. Kao. 1970. High frequency haploid production in barley (Hordeum vulgare L.). Nature 225: 874–876. Kunze, W. 2004. Technology Brewing and Malting, 3rd international ed.—in English. VLB, Berlin. Langridge, P., N. Paltridge, and G. Fincher. 2006. Functional genomics of abiotic stress tolerance in cereals. Brief Funct. Genom. Proteom. 4:343–354. Mathre, D.E. (ed.). 1997. Compendium of Barley Diseases, 2nd ed. American Phytopathological Society, St. Paul, MN. Metcalfe, D.R. 1995. Barley. pp. 82–87. In A.E. Slinkard and D.R. Knott (eds.). Harvest of Gold: the History of Field Crop Breeding in Canada. University of Saskatchewan Press, Saskatchewan, Canada. Mornhinweg, D.W., D.R. Porter, and J.A. Webster. 1995. Registration of STARS-9301B barley germplasm resistant to Russian wheat aphid. Crop Sci. 35:603. Mornhinweg, D.W., D.R. Porter, and J.A. Webster. 1999. Registration of STARS-9577B Russian wheat aphid resistant barley germplasm. Crop Sci. 39:882–892. Newman, R.K., C.W. Newman, and H. Graham. 1987. Nutritional implication of β-glucans in barley. pp. 773– 780. In S. Yasuda and T. Konishi (eds.). Barley Genetics V. Proc. 5th Int. Barley Genet. Symp., Okayama, Japan, October 6–11, 1986. Poehlman, J.M. 1985. Adaptation and distribution. pp. 1–17. In D.C. Rasmusson (ed.). Barley. Agronomy Monograph 26. American Society of Agronomy, Madison, WI. Porter, D.R., D.W. Mornhinweg, and J.A. Webster. 1999. Insect resistance in barley germplasm. pp. 51–61. In S.L. Clement and S.S. Quisenberry (eds.). Global Genetic Resources for Insect-Resistant Crops. CRC Press, Boca Raton, FL. Prom, L.K., B.J. Steffenson, B. Salas, J. Moss, T.G. Fetch, and H.H. Casper. 1996. Evaluation of selected barley accessions for resistance to fusarium head blight and deoxynivalenol concentration. pp. 764–766. In •• Scoles and •• Rossnagel (eds.). Proc. of the V International Oat Conference & VII International Barley Symposium, University of Saskatchewan, Saskatoon, Canada. July 30– August 6, 1996. University Extension Press, Saskatoon, Canada. Rasmusson, D.C. (ed.). 1985. Barley. Agronomy Monograph 26. ASA and CSSA, Madison, WI. Rasmusson, D.C. and R.L. Phillips. 1997. Plant breeding progress and genetic diversity from de novo variation and elevated epistasis. Crop Sci. 37:303–310. Roelfs, A.P. and J. Huerto-Espino. 1994. Seedling resistance in Hordeum t barley stripe rust from Texas. Plant Dis. 78:1046–1049. Roelfs, A.P., J. Huerta-Espino, and D. Marshall. 1992. Barley stripe rust in Texas. Plant Dis. 76:538. Ullrich, S.E., D.M. Wesenberg, H.E. Blockelman, and J.D. Franckowiak. 1995. International cooperation in barley germplasm activities. pp. 157–170. In R.R. Duncan (ed.). International Germplasm Transfer: Past and Present. CSSA and ASA, Madison, WI. Urrea, C.A., R.D. Horsley, B.J. Steffenson, and P.B. Schwarz. 2005. Agronomic characteristics, malt quality, and disease resistance of barley germplasm lines with partial fusarium head blight resistance. Crop Sci. 45:1235–1240. Weaver, J.C. 1950. American Barley Production. Burgess 39 Publications, Minneapolis, MN. Webster, J.A., C.A. Baker, and D.R. Porter. 1991. Detection and mechanisms of Russian wheat aphid (Homoptera: Aphidadae) resistance in barley. J. Econ. Entomol. 842(2):669–673. AUSTRALIA David M.E. Poulsen and Reg C.M. Lance HisTORY—BARLEY PRODUCTiOn in AUsTRALiA Barley is the second most important cereal crop, after wheat, in Australia. It has expanded from the irst crop of 3.24 ha, sown at Farm Cove after the arrival of the First Fleet in 1788 (Sparrow and Doolette 1975, 1987 loc. cit.), to an industry that 40 annually produces 4.5–5.5 million tons of grain (Lovett 1997; Powell 1997; Stuart 1997; Healy 2001). Lovett (1997) stated that the net value of Australian barley production was greater than A$1 billion per year. The barley-growing regions are conined to the southern temperate cereal-growing regions with rainfall varying from 250 to 600 mm (see Fig. 9.5.1, Chapter 9 Australia; Paynter and Fettell 2008) The igure indicates regional production as tons per square kilometer based on the Australian Bureau of Statistics 2006 Agricultural Census data and is for the 2005/2006 harvest. The areas of greatest production are southern Western Australia; the southern Eyre Peninsula, Yorke Peninsula, midnorth and upper southeast of South Australia; the “Mallee” and Wimmera regions of Victoria; southern New South Wales; and the Darling Downs of Queensland. Other 41 production regions include northern New South Wales and Tasmania. Most Australian barley cultivars are two-row spring types with white aleurone. In most areas, Ullrich—Production, Improvement, and Uses c08.indd 184 8/30/2010 8:28:54 PM Barley Breeding History, Progress, Objectives, and Technology the crops are sown in late autumn or early midwinter, lower in spring, and are harvested in early summer. Australian barley production over the 5-year period 2002–2006 inclusive averaged 7 million 42 tons of which 2.619 MT was used domestically and 4.590 MT was exported (Table 8.3.1). Only 0.169 MT was used for malt and other human uses, and most (2.257 MT) are used for feed. Of the exported grain, 2.461 MT was feed, 1.485 MT was malting barley, and 0.627 MT was malt (in grain equivalents). This malt is mostly exported to Asian countries, vis-à-vis, Vietnam, Japan, South Korea, the Philippines, and Thailand (Emalt.com 2008). So, in total, malt production was 0.796 MT. Although Australia is not a major malting barley producer, its exports represent approximately 25% of the world trade. The largest barley production is in South Australia (SA), closely followed by Western Australia (WA), then Victoria (Vic), New South Table 8.3.1 Australian barley production and disposal (5year average, 2002–2006) Australian Barley Production 137 and Use 5-Year Average (2002–2006) 000 t Production 7115 Domestic use As malt and other human use Feed Seed 2619 169 2257 193 Export Feed barley Malting barley Malt (grain equivalent) 4590 2461 1485 627 Source: Australian Bureau of Agriculture and Resource Economics (ABARE) and Australian Bureau of Statistics (ABS). Table 8.3.2 185 Wales (NSW), Queensland (Qld), and Tasmania (Tas). Yields are highest in the southern regions 43 and lowest in New South Wales and Queensland (Table 8.3.2). For practicality of research management, the Grains Research and Development Corporation (GRDC) has divided the Australian cropping environment into three regions, namely, the northern region, the southern region, and the western region (Davies 1993). The northern 44 region includes Queensland and northern New South Wales. Southern New South Wales, Victoria, South Australia, and Tasmania make up the southern region. The dividing line between the northern and southern regions is drawn at Dubbo (latitude 33° S). Only Western Australian cropping areas are included in the western region. During the 1990s, approximately 63% of the Australian barley crop was produced in the southern region (Healy 2001). The western region contributed 23% of the crop, while the northern region produced approximately 14%. These igures often vary, as seasonal conditions in Australian barley-growing areas can luctuate widely (Powell 1997). In most seasons, 40%–50% of the Australian barley crop is accepted as malting barley (Lovett 1997; Stuart 1997; MacLeod 2001). However, only 700,000 t or 35% of malt barley production is malted within Australia (MacLeod 2001). Approximately two-thirds of the Australian malt production is exported annually (Powell 1997). Thus, approximately 88% of the total Australian malting barley crop is exported, making the country the third largest exporter of malt, after Europe and North America (MacLeod 2001). Consequently, the viability of the Australian barley industry is heavily reliant on maintaining its share of the international market (Stuart 1997). Australian barley production by state (5-year average, 2001–2005) State Units Qld NSW Vic Tas SA WA Australia Area Production Yield Hectares (’000) Tons (’000) Tons per hectare 113 175 1.55 778 1356 1.74 793 1477 1.86 8 26 3.06 1176 2242 1.91 1160 2126 1.83 4030 7402 1.84 138 F Source: Australian Bureau of Agriculture and Resource Economics (ABARE) and Australian Bureau of Statistics (ABS). Ullrich—Production, Improvement, and Uses c08.indd 185 8/30/2010 8:28:55 PM Barley: Production, Improvement, and Uses 186 The main export market for Australian malting barley is the Asia Paciic region. China buys approximately 50% of the Australian malting barley crop each year (Stuart 1997). Furthermore, Sewell (1997) predicted that Chinese import requirements of malting barley from Australia are likely to reach 1.36 million tons by the year 2010. This has arisen through increased demand for beer within China and increased foreign investment in Chinese breweries (MacLeod 2001). Because brewhouse technology has been reined, quality speciications for the Chinese market have become more stringent (Spiel 1999). Japan is also a key market for Australian malting barley and commands premium prices. In the early 1980s, Australia’s share of the Japanese malting barley market was approximately 33% (Powell 1997). However, by 1997, signiicant improvements in the quality of Canadian and European cultivars led to Australian malting barley exports to Japan being reduced to 124,226 t, or 15.5% of Japan’s malt barley intake (Fukuda et al. 1999). Other important markets for Australian malting barley include Taiwan, the Philippines, South Korea, and several other Asian and South American countries (Stuart 1997). Australian malt is principally exported to Japan, Thailand, the Philippines, and South Korea (MacLeod 2001). While the Japanese market for Australian malt has declined in a similar pattern to that seen for malting barley, there have been signiicant increases in sales to Thailand, Vietnam, and South Korea (E-malt. com 2008). These three countries account for 47% of Australian malt exports. Increases in brewing capacity in Thailand, Vietnam, and South Korea should increase demand for Australian malt and malting barley but will also necessitate stable grain supplies and improved variety performance. In response to the above market signals, the Australian industry has taken several steps to improve the quality and reliability of its export barley and malt supplies. The accreditation of Australian malting cultivars is coordinated by the Malting and Brewing Industry Barley Technical Committee (MBIBTC) (Hawthorne 1999). This committee has established technical guidelines, F which are available to all breeding programs. Those guidelines include detailed experimental schedules and speciications to be met by potential cultivars before domestic and/or export malting accreditation is awarded (MBIBTC 1998). Malting barley usually attracts a price premium of A$30–A$50 per ton over feed barley in Australia. To qualify for malting grade, the grain must irstly be obtained from an accepted malting cultivar. Point-of-delivery assessment must also prove that the grain meets appropriate speciications, typically, a low proportion of small grain or “screenings,” protein between 8.5% and 12.0%, bright grain color, and absence of contaminants such as smut, sclerotes, weed seeds, and soil (ABB Grain 2008; CBH Group 2008; Grainco 2008). 45 HisTORY—BARLEY iMPROVEMEnT in AUsTRALiA The early history of Australian barley improvement was comprehensively described in review papers written by Sparrow and Doolette (1975, 1987 loc. cit.). Most early Australian cultivars were obtained from Europe. These tended to be late maturing and befell the consequences of terminal droughts or the hot dry inishes to the seasons so typical in southern Australia. A South Australian farmer, Samuel Prior, developed the irst Australian barley cultivar in 1903 from a selection of imported seed line of Chevalier (or alternatively, Spratt Archer). Prior’s Chevalier selection was recognized for its superior malting quality and subsequently proved to be well adapted to the dry South Australian environment. It became known as Prior and led to a massive expansion in barley production. Prior became the backbone of the Australian barley industry until the end of the 1960s. From the 1920s to the 1950s, barley breeding was conducted on a part-time basis in Victoria, New South Wales, and South Australia. Early breeding programs led to Prior A bred by Albert Pugsley and Research from the program at Werribee in Victoria, bred by A.R. Raw in 1942. However, in 1956, a barley improvement scheme Ullrich—Production, Improvement, and Uses c08.indd 186 8/30/2010 8:28:55 PM Barley Breeding History, Progress, Objectives, and Technology was established with funds provided by maltsters, brewers, and growers and with matching funds from the commonwealth government. This scheme led to the development of full-time breeding programs in South Australia and Victoria, and consequently, the release of several barley cultivars, which inally replaced Prior. The most signiicant of these was the malting barley Clipper, released from the Waite Agricultural Research Institute in 1968. The genotype × environment interaction studies of Finlay and Wilkinson (1963) had a signiicant inluence on the South Australian program. Improved yield over a range of environments was found in introductions from North Africa. This material was early maturing, tolerant of alkaline and sodic soils, disease resistant (especially cereal cyst nematode [CCN], mildew, and other leaf diseases), and tolerant of hot dry inishes. A similar well-adapted material was sourced from the United States, but this originally came from North Africa. New breeding programs were established in Western Australia in 1962 and in Queensland, New South Wales, and Tasmania in the late 1960s and early 1970s. Barley improvement in Queensland effectively began in 1967 when Dr. R.P. Johnston of the Department of Primary Industries initiated a series of trials to evaluate breeding lines from the Waite Institute (Poulsen 2001). A breeding program was designed to breed high-yielding malting and feed barley cultivars, targeting Queensland barley-producing regions. Funding management for barley research and development went through a series of iterations from the late 1970s through to the present day (Joppich 1985; Purse and McNee 1985; Smith 1987; Davies 1993). Originally, funding for different crops was governed through a series of research councils. However, in 1990, 4 federal research councils and 10 state wheat and barley committees were amalgamated to form the GRDC, with the overarching responsibility to drive research and development across the entire Australian grains industry (Davies 1993). This has led to the development of a coordinated barley research and development program for Australia, including strong barley improvement programs in each of the southern, western, and northern 187 regions (Lovett 2001). Further agronomic research of French and Schultz (1984a,b) has drawn attention to the importance of water use eficiency in water-limiting environments and the importance of understanding abiotic and biotic constraints, which may limit productivity and quality improvements. Nationally focused projects have been established. For example, the National Barley Molecular Marker Program (NBMMP), later to be amalgamated with the equivalent wheat program to become the Australian Winter Cereal Molecular Marker Program (AWCMMP), was set up to develop and implement molecular technologies on a national scale (Langridge 1997; Barr et al. 2001). The effects of developing focused barley breeding programs became clear during the 1980s, as new malting cultivars with speciic regional adaptation were released, replacing Clipper (Sparrow 1984). Schooner was released from the Waite Institute in 1983 and quickly dominated the southern States. In Western Australia, the locally bred cv. Stirling similarly took over from Clipper. A third cultivar, Grimmett, replaced Clipper in Queensland and in northern New South Wales. The result of the signiicant investment from the GRDC together with inputs from the malting and brewing industries and from marketing and grain handling organizations and authorities have 46 led to a signiicant improvement in the number and quality of malting and feed varieties released in the 1990s and early 2000s. Other cultivars have since been released in each of the GRDC regions, including high-yielding feed types. A representative malting barley pedigree chart is shown in Fig. 8.3.1, where possible varieties are listed in chronological order. Caution should be taken in attempting to determine the order of each cross; special expediency sometimes prevented the convention of putting the female parent irst. Some observations can be made. Early programs were sparse and based on a limited number of European parents. There was a limited use of backcrossing to incorporate disease resistance traits. Cycle times were often 12 or more years. The 1970s and 1980s saw the use of North African landraces providing both an appropriate Ullrich—Production, Improvement, and Uses c08.indd 187 8/30/2010 8:28:55 PM F Barley: Production, Improvement, and Uses 188 fig. 8.3.1. ••. phenological adaptation and disease resistances (such as CCN, mildew, and alkaline soil). New introgressions came from Europe, North America, and Japan. More varieties have been released with shorter cycle times. The use of molecular genetic markers has enabled more complex crosses to be managed more easily and with greater eficiency (e.g., Flagship, Sloop-SA, and Sloop-Vic). Barley Australia accredited varieties are shown as shaded rounded rectangles. The newer malting varieties include Western Australia: Gairdner, Baudin, Hamelin, and Vlamingh; South Australia: Sloop and the backcross derivative Sloop-SA; Tasmania: Shannon, Franklin, and Vertess; Queensland: Tallon and Lindwall; Victoria: Arapilies, Sloop-Vic, Buloke, F and Hindmarsh; and New South Wales: Cowabbie. Feed varieties and some malting lines are shown in Table 8.3.3. The table has been sorted by date of release (or introduction) and by name. RECEnT PROGREss in AUsTRALiAn BARLEY iMPROVEMEnT PROGRAMs Barley Breeding Australia (BBA)— a new paradigm BBA began operating on July 1, 2006 as the Australian national barley breeding program implementing a national plan for breeding improved varieties to beneit the barley industry. Ullrich—Production, Improvement, and Uses c08.indd 188 8/30/2010 8:28:55 PM Table 8.3.3 Australian feed barleys and other signiicant varieties Name Year Released Pedigree Breeder Trabut Abyssinian Commander Maltworthy Deputy Ablyn Noyep Anabee Resibee A14 A16 Bussell Ketch Lara Cutter Corvette Parwan Bandulla Beecher Cantala Malebo Waranga O’Connor Moondyne AB6 Windich Yerong Gilbert Brindabella Kaputar Namoi Dash Mundah Molloy Picola Fitzgerald Tilga Unicorn Doolup Lindwall Tantangara Wyalong Yambla Keel Binalong MacKay Cowabbie Milby Tulla Capstan Maritime Grout Urambie Vertess Yarra Fleet Hannan Lockyer Roe 1916 1920 1930 1938 1939 1952 1959 1962 1962 1968 1968 1968 1969 1971 1975 1976 1979 1981 1981 1981 1981 1981 1984 1987 1988 1989 1990 1992 1993 1993 1993 1995 1995 1996 1996 1997 1997 1997 1998 1998 1998 1998 1998 1999 2001 2001 2002 2002 2002 2004 2004 2005 2005 2005 2005 2006 2007 2007 2007 Algerian introduction selection Abyssinian introduction Coast selection Prior × Beaven’s Special Hero selection Abyssinian × Flyn Prior’s Chevalier selection Research selection Research selection Prior × Ymer Prior × Ymer Prior × Ymer Lenta × Noyep Research × Lenta Prior A × Proctor Bonus × CI-3576 ((((Plumage Archer × Prior) × Lenta)) × (Research × Lenta)) ((Prior × Lenta) × (Noyep × Lenta)) Atlas × Vaughn Kenia × Erectoides 16 Pallidum selection (CPI-11083) ((Plumage Archer × (Prior × Lenta)) × (Research × Lenta)) × Clipper WI-2231 × (Atlas 57 × A14) ((Dampier × A14) × Kristina)) × (Clipper × Tenn 65-117) CPI-71283 × 4* Clipper (Atlas 57 × A16) × Parwan M22 × Malebo (Armelle × Lud) × Luke ((((Weeah × CI-17115) × HCB27) × Jadar II) × Cantalla) ((5604 × 1025)) × ((Emir × Shabet) × CM67) (Sultan × Nackta) × (RM1508 × Godiva) (Chad × Joline) × Cask O’Connor × Yagan (Golden Promise × WI-2395) × 72S:267 VB75031 × Elgina Onslow × Tas 85-466 Forrest × Cantala 54C25 × 51C38 75S:323 × 74S:314 Triumph × Grimmett AB6 × Skiff Schooner × Stirling Skiff × FM 437 (CPI-18197 × Clipper) × WI-2645 Blenheim × (Skiff × O’Connor) Cameo × Koru (((AB6 × Franklin) × Franklin)) × (Rubin × Skiff) (((AB6 × Franklin) × Franklin)) × (Rubin × Skiff) Skiff × FM 437 (Waveney × WI-2875) × (Chariot × Chebec) 74S:314 × 74S:309 Cameo × Arupo “S” (Yagan × Ulandra) × Ulandra Franklin × Cooper (VB9108 × Alexis) × VB9104 Mundah × Keel × Barque B28719 × (Windich × Morex) Tantangara × VB9104 Doolup × (Windich × Morex) DANSW NSWDA DANSW RAC DANSW NSWDA RAC VicDA VicDA DAWA DAWA DAWA WARI VicDA WARI WARI VicDA VicDA VicDA NSWDA Vic-DPI DAWA DAWA CSIRO-PI DAWA NSWDPI Qld-DPI NSWDPI CIMMYT NSWDPI NFC DAWA DAWA Vic-DPI DAWA NSWDPI Kirin Australia DAWA Qld-DPI NSWDPI NSWDPI NSWDPI WARI NSWDPI Qld-DPI NSWDA NSWDA NSWDPI Uni Adelaide Uni Adelaide Qld-DPI NSWDPI UTas and TASDPIWE MBQIP Uni Adelaide DAFWA DAFWA DAFWA F 189 Ullrich—Production, Improvement, and Uses c08.indd 189 8/30/2010 8:28:55 PM Barley: Production, Improvement, and Uses 190 F BBA is an unincorporated joint venture between the GRDC, the Queensland Department of Primary Industries and Fisheries, the New South Wales Department of Primary Industries, the Department of Primary Industries, Victoria, the University of Adelaide, the Department of Agriculture and Food, Western Australia, and the Tasmanian Institute of Agricultural Research. BBA is governed through an advisory board formed by the parties and coordinated through a management committee. The formation of BBA rationalized the previous six state-based barley breeding programs into one national program with three regional nodes: BBA-North (QDPIWarwick, Queensland), BBA-South (University of Adelaide, Urrbrae, South Australia), and BBAWest (DAFWA, Kensington, Western Australia). The restructure aims to use resources clearly and eficiently in barley breeding so as to provide the foundation of a future vibrant barley industry (Lance et al. 2007; Reading 2007). The previous programs in Victoria, New South Wales, and Tasmania will be focusing their efforts on germplasm development and barley prebreeding. The breeding nodes are to focus on the development, release, and adoption of varieties meeting speciic market and regional needs. As such, they must achieve operational eficiencies through the adoption of a “best practice in breeding” framework. BBA-North is actively changing its primary focus from breeding malt varieties to breeding feed varieties, thereby meeting the future demand for feed grain in northeastern Australia. BBA-South has extended its trial network to cover the neutral to alkaline soils in New South Wales and Victoria and the alkaline soil regions of southern Western Australia. The most adapted barleys are mid season to earlier maturing. BBA-West, in addition to conducting trials in Western Australia, conducts yield trials and nurseries in New South Wales, Victoria, and Tasmania, on neutral to acid soils. As such, the focus is on medium- to later-maturing varieties suited to the medium- to high-rainfall zones. Germplasm from the NSWDPI, Victorian and Tasmanian breeding programs is being pro47 gressed and integrated into the operations of BBA-West and BBA-South. Barley Australia—malting barley accreditation Barley Australia was established in 2005 as the peak body for the Australian barley industry. It was the recognition of a future deregulated marketing environment for Australian barley that was a catalyst for its creation as it was recognized by the maltsters and marketers (purchasing the barley crop from the Australian farmers) that there was a need for a body to deal with “industry good” functions that had previously been undertaken by the statutory marketing companies that preexisted the new market environment, such as cooperative industry decisions on quality standards, variety suitability to malting, and a coordinated approach to industry-wide research. Barley Australia is entirely self-funded by the maltsters and marketers of Australian barley and is a testament to the members in their ability to cooperate for industry beneit even though in the Australian marketplace they are staunch competitors. Now in its fourth year, Barley Australia serves a signiicant purpose in the Australian industry as a “shop window” for the industry and disseminates information to the Australian barley industry stakeholders as well as to the international marketplace. Barley Australia is responsible for creating and managing the national varietal accreditation system as well as the Barley Australia Assured Quality trademark for HACCP-certiied 48 barley and malt out of Australia. More information on Barley Australia can be sought from the Web site http://www.barleyaustralia.com.au/. 49 The MBiBTC The MBIBTC has technical malting and brewing experts representative of Australia’s major malting and brewing companies. Varieties submitted to Barley Australia are evaluated by an MBIBTC member for the irst year of commercial evaluation. If deemed suitable, then the malt from the evaluation passes to Pilot Brewing Australia (PBA). PBA PBA conducts the last step in the process of evaluation of malting barley for either domestic or Ullrich—Production, Improvement, and Uses c08.indd 190 8/30/2010 8:28:55 PM Barley Breeding History, Progress, Objectives, and Technology export market suitability. PBA is malting and brewing industry funded and also cofunded by the GRDC. Samples for pilot brewing are taken from commercial-scale malting evaluation trials. The facility is able to replicate the requirements of both domestic and export brewers utilizing sugar or starch adjuncts. For a new malting variety to be accredited, it must pass two successful years of commercial malting evaluation and pilot brewing or commercial-scale brewing with all stages being reviewed by the MBIBTC. The major proportion of the northern region barley crop is purchased for domestic stock feed. Most northern feed barley is used for cattle since the region contains the highest concentration of 50 feedlots in Australia (Barr and Kneipp 1995). Feed for the monogastric industries is also a signiicant market, with approximately 30% of the grain purchased by stock feed manufacturers for pig and poultry rations (M. Covaceveitch, pers. comm.). AUsTRALiAn BARLEY BREEDinG OBJECTiVEs Barley breeding objectives have changed substantially in Australia over the past century. While the basic agronomic objectives of grain yield, lodging resistance, and appropriate maturity have changed relatively little, the evolution of on-farm cultural practices has resulted in a signiicant increase in the number of diseases affecting the crop. Objectives relating to grain quality have also increased in number and detail, as knowledge of grain quality for both malting and feed use has become more reined. See Table 8.3.4 for a comprehensive summary of breeding objectives/ traits. Agronomic traits Yield, lodging resistance, straw strength, height (short straw, medium height straw); phenology (late maturity, short straw for longer seasons; early maturity, taller straw for drier shorter 51 seasons); head loss, stem breakage. 191 Improved grain yield has been the primary objective of Australian barley breeding activities since the nineteenth century (Sparrow and Doolette 1975). From the outset, modern Australian programs were clearly designed with yield as the primary selection criterion (Ellis 1983; Johnston 1983; Portman 1983; Read 1983; Sparrow 1983; Vertigan 1983). However, it is considered extremely important that yield advances are made together with improvements in grain quality to protect the Australian share of the world barley market (GRDC 2001). Johnston (1974) had concluded that a combined malting and feed breeding program could service both markets, improving both yield and quality. As well as grain yield, Australian barley breeders have selected material for a range of agronomic traits. All of the programs made resistance to lodging or a high level of straw strength a key agronomic breeding objective (Portman 1983; Read 1983; Vertigan 1983). Johnston (1983) placed a very high priority on lodging resistance as the inability of barley cultivars to stand up in the ield was seen as a major limitation to the further expansion of the crop in Queensland. Plant height was established as another objective of the Australian programs. For example, short cultivars were desirable in the high- and mediumrainfall zones of Western Australia, while taller cultivars suited low-rainfall areas (Portman 1983). Controlling head loss caused by stem breakage was another signiicant agronomic trait targeted, in particular, for Western Australia, New South Wales, and Tasmania. Crop phenology targets of the different Australian programs were diverse and were driven by the amount of available moisture and the duration of optimal growing conditions. Read (1983) explained how longer-season, short-straw material was being developed for parts of southern New South Wales where early-sown cultivars were suitable. These types were also suitable for Tasmania and eastern Victoria. In Western Australia, longer-season cultivars were more suitable for the high- and medium-rainfall zones, while quick maturing barleys were required for the parts of the cropping area with low rainfall (Portman 1983). Most areas of South Australia Ullrich—Production, Improvement, and Uses c08.indd 191 8/30/2010 8:28:55 PM F Barley: Production, Improvement, and Uses 192 Table 8.3.4 Differences in the grain production environments and key selection targets and traits for GRDC breeding programs in northern, southern, and western regions Southern Region Western Region Northern Region Latitude 33° S–43° S Predominantly winter rainfall Lighter, shallower soils, sands in WA Winter cropping only Latitude 28° S–35° S Predominantly winter rainfall Lighter, shallower soils, sands in WA Winter cropping only Latitude 22° S–33° S Predominantly summer rainfall High proportion of heavy clay soil types Winter/summer crop rotations Double cropping Agronomic traits Yield, grain weight, plumpness, hectoliter weight, seedling vigor, phenological adaptation (bvp, ppd, vrn) Malting quality Malt extract, diastatic power (DP), apparent attenuation limit (AAL), low viscosity, increased free alpha amino nitrogen (FAAN) Feed quality Digestible energy (DE), metabolizable energy (ME), animal intake, low bloat, low phytate Resistance/tolerance to biotic stress Net form net blotch Spot form net blotch Scald Powdery mildew Leaf rust (part of SA, S-NSW, NE, and SW-Vic, Tas) BYDV Barley grass stripe rust Rhizoctonia 139 Major pests Cereal cyst nematode (SA, Vic) Resistance/tolerance to abiotic stress Drought/terminal drought Alkaline soil/boron toxicity tolerance (SA, Vic-Mallee) Acid soil/aluminum toxicity (sNSW, NE, and SW-Vic) Frost Salinity/sodicity tolerance Manganese deiciency tolerance Waterlogging tolerance Preharvest spouting tolerance Blackpoint/kernel discoloration tolerance Screening tolerance Winter cropping only Net form net blotch Spot form net blotch Scald Powdery mildew Leaf rust BYDV Net form net blotch Spot form net blotch Scald Powdery mildew Leaf rust Stem rust Common root rot Crown rot Root lesion nematode—Pratylenchus neglectus and P. sp. Root lesion nematode—Pratylenchus thornei and P. sp. Drought/terminal drought Alkaline soil/boron toxicity tolerance (WA-Mallee) Acid soil/aluminum toxicity Frost Salinity/sodicity tolerance Waterlogging tolerance Drought/terminal drought Frost Preharvest spouting tolerance Blackpoint/kernel discoloration tolerance Screenings tolerance Winter cropping only Preharvest spouting tolerance Blackpoint/kernel discoloration tolerance Screening tolerance and parts of Victoria required medium-quick maturing genotypes. Data from Queensland and northern New South Wales revealed that medium to medium-slow maturing European-type germ- F Winter/summer crop rotations, double cropping plasm gave better grain yield and quality in eastern environments, but that quicker maturity was required in western areas (•• Poulsen, unpub52 lished data). Ullrich—Production, Improvement, and Uses c08.indd 192 8/30/2010 8:28:56 PM Barley Breeding History, Progress, Objectives, and Technology Malting and feed quality Quality attributes include both physical and biochemical grain characteristics. Sparrow and Doolette (1975) suggested that early assessment of malting quality was aimed at identifying lines with plump and uniform grain, high malt extract, and quick and even germination when malted. That deinition remains true today, and a host of individual parameters is recognized as contributing to each of the above attributes. In 1983, the MBIBTC was formed to provide industry guidance to the breeding programs (Armitt and Way 1990). It has provided Australian barley breeders with guidelines to develop new malting cultivars (MBIBTC 1998) and has developed a rating system for malting barleys, which takes into account weighted factors for malt extract, protein modiication, diastatic power (DP), wort viscosity, and dimethyl sulide content (Armitt and Way 1990). The relative activities of speciic enzymes such as α-amylase, β-amylase, and β-glucanase and cellular constituents such as β-glucan and starch are also recognized as target parameters for malting barley improvement programs. Grain dormancy is essential to protect against preharvest sprouting caused by summer 53 storms (Poulsen et al. 1995). Sprouting interferes with the malting and brewing processes by causing uneven starch and cell wall degradation in the grain. The southern and western regions are less concerned about dormancy as those areas have predominantly winter rainfall patterns; however, the southern coast of Western Australia is prone to preharvest sprouting problems and a degree of dormancy is required. While some quality speciications are required for all malting barleys, there is now a dichotomy in domestic and export market requirements due to the different sources of carbohydrate adjuncts used in the brewing process (MBIBTC 1998). The domestic brewing industry requires moderate levels of DP; however, many Asian breweries require high DP values. It has gradually become recognized that speciic quality attributes can enhance livestock performance. Johnston (1974) commented that barley was seen as a source of energy and protein for 193 both the ruminant and nonruminant feeding industries. He concluded that high protein requirements of the feed industry could be better dealt with through agronomic management than breeding. Recent research has begun to deine attributes of barley affecting animal feeding performance. Bowman and Blake (1996) identiied considerable genetic variation in barley ruminant feed quality. Dry matter digestibility, starch digestibility, starch content, starch particle size, ground grain particle size, and grain hardness were identiied as key traits in cattle nutrition. Resistance to biotic stresses Early Australian priorities for disease resistance breeding were scald (Rhynchosporium secalis), covered smut (Ustilago segetum var. hordei), and powdery mildew (Blumeria graminis f. sp. hordei) (Sparrow and Doolette 1975). New South Wales Agriculture produced scald-resistant lines from the cross Commander × Prior during the 1940s. However, named varieties were not released. In the same era, covered smut and powdery mildew resistant (Mlk; now susceptible) (from the Indian variety Kwan) backcross-derived lines based on Prior and Research were produced at the Waite Institute (Pugsley and Vines 1946; Pugsley 1951), leading to Prior A (=Kwan × 6* Prior; Kwan was 54 thought to have at least three genes conferring resistance to covered smut), which has proven to be a cornerstone of barley adaptation and improvement in southern Australia. Additionally, Prior was one of the earliest of the European introductions and contains the Eam1 gene, which is prevalent in Australian germplasm (•• Franckowiak, pers. comm.) and is the source of earliness in 55 Erbet (=Prior × 7* Betzes). This inherent “earliness” enabled grain ill to be completed in a hot dry spring. In the 1970s and early 1980s, disease resistance was considered less important than grain yield or grain and malting quality. Most programs used disease-resistant parents, but effective resistances were seen as desirable, rather than essential, characteristics in new varieties (Johnston 1985; Gilmour 1993). Disease resistance breeding took higher priority in the New South Wales breeding Ullrich—Production, Improvement, and Uses c08.indd 193 8/30/2010 8:28:56 PM F Barley: Production, Improvement, and Uses 194 F program. Resistance to leaf diseases was seen as necessary for early-sown cultivars grown in the southern parts of the State (Read 1983). Development of scald-resistant material was also occurring in Western Australia (Portman 1983, 1985), South Australia (Sparrow 1983), and Victoria (Ellis 1983). By 1985, diseases were recognized as restricting barley production throughout Australia. Hirsch (1985) estimated yield losses from diseases in southern Australia between 1973 and 1983 to be more than 16% of the overall crop value, with a net cost of A$20–A$25 million. Rees (1985) noted that considerable changes had occurred in Queensland, and several diseases had become signiicant, including stem rust (Puccinia graminis), leaf rust (Puccinia hordei), scald, net-type net blotch (Pyrenophora teres f. sp. teres), spot-type net blotch (P. teres f. sp. maculata), crown rot (Fusarium pseudograminearum), and common root rot (Cochliobolus sativus). This was in direct contrast to an earlier statement that the only regular problem in northern barley crops was powdery mildew (Rees et al. 1981). The increasing disease levels were believed to have been due to changes in seasonal conditions, pathogen genetic diversity, cultivars, and agricultural practices. Severe stem rust occurred in Queensland barley crops during 1982–1984 when the highly susceptible cv. Galleon entered production on the Darling Downs (Dill-Macky 1992). Localized epidemics occurred in 1982 and 1984; however, a more general epidemic occurred in the Darling Downs and Burnett regions during the 1983 season. Shortly afterwards, Rees (1985) suggested that the predominance of a single cultivar in Queensland, Grimmett, was making the State’s crop generally vulnerable to disease, in the event that new pathogens or pathotypes became established. This threat was conirmed in 1988, when new leaf rust pathotypes caused a signiicant epidemic in Grimmett crops. A combination of cultivar susceptibility and suitable weather conditions led to highly damaging epidemics of leaf blotches in 1998 (Poulsen et al. 1999; Rees et al. 1999). The changing disease pressures in the northern region 56 led to changes in the NBIP breeding objectives. A stem rust research program was initiated after the 1982–1984 epidemics (Johnston 1985), and other expansions to the program occurred after the 1988 leaf rust and 1998 net blotch epidemics (Poulsen 2001). In each case, projects were developed to assess the impact of the diseases and to identify useful sources of genetic resistance (Cotterill et al. 1992; Dill-Macky 1992; Platz 57 2001). The most signiicant nematode pest of barley in Australia has been CCN (Heterodera avenae), which occurs naturally in South Australia and Victoria. The southern programs have bred for CCN resistance for many years (Ellis 1983; Sparrow 1983). The irst Australian CCNresistant barley cultivar was Galleon, a South Australian feed variety with the CCN resistance gene Ha4 (Karakousis et al. 2003c) introgressed from a North African landrace, CIho-3576 (Gall eon = [Clipper × Hiproly] × 3* WI-2231, where WI-2231 = Proctor × CIho-3576). The second resistant cultivar was Chebec, which contained the CCN resistance gene Ha2 (Barr et al. 1998) from Orge Martin (Chebec = [Orge Martin × 2* Clipper] × Schooner). Barleys are generally tolerant toward CCN, but resistance varieties reduce the number of CCN in the soil, which has a signiicant and positive effect on subsequent intolerant and susceptible wheat crops. There are no major insect pests at present in Australia that can be addressed by genetic tolerance or resistances. Some pests such as Russian leaf aphid are regarded as “quarantine threats,” and there are prebreeding programs to overcome potential incursions. Resistance to abiotic stresses Breeding for abiotic stress has generally been associated with soil characteristics. In the 1980s, the Waite Institute breeding program was selecting for manganese-eficient cultivars to address deiciency problems in some South Australian soils (Sparrow 1983). Conversely, the New South Wales program, now followed by the BBA-West program, has been developing cultivars adapted to aluminum and manganese toxicity for production on acid soils (Read 1985). Parts of Victoria, South Australia, and Western Australia have Ullrich—Production, Improvement, and Uses c08.indd 194 8/30/2010 8:28:56 PM Barley Breeding History, Progress, Objectives, and Technology alkaline and sodic soils of marine origin, and these are associated with boron toxicity. The breeding programs in those States have developed conventional and molecular screening methodologies and have incorporated boron tolerance into elite germplasm (Jefferies et al. 1997; Barr et al. 2000). inTEGRATiOn Of TECHnOLOGiEs in AUsTRALiA Barley breeding methods in Australia Many barley breeders use pedigree breeding systems. Speciic parents are hybridized by controlled pollination, and individual lines are selected from segregating material in the F2 and subsequent generations (Anderson and Reinbergs 1985). Selection usually commences with simple traits and progresses to multienvironment yield trials. Pedigree breeding strategies form the core component of most, if not all, of the Australian barley improvement programs. Various modiications may be applied to pedigree breeding strate- 195 gies. Riggs et al. (1982) designed a pedigree program in which F4 bulks were tested in ield trials to bring yield evaluation forward by 1 year and to improve resource use eficiency in their program. The NBIP uses a variation of this technique in its cross-evaluation strategy (Johnston 1983; Poulsen et al. 1995b). The Western Australian program uses several breeding strategies that are utilized where appropriate (Fig. 8.3.2). The basic “conventional” method is an F2 progeny method with a reselection phase at the F5. Almost 50% of the stage 2 program is derived from doubled haploids (DHs). Marker-assisted selection (MAS) coupled with DH, single-seed descent (modiied, SSDm) and male sterile facilitated recurrent selection (MSFRS) are seen as the methods of choice for increasing the rate of genetic gain and for reducing the time to release of new varieties. Backcross breeding aims to recover as much of a desirable recurrent genotype as possible while introgressing a new gene from a less-adapted parent (Harlan et al. 1922, cited by Anderson and 58 fig. 8.3.2. ••. A, agronomic; Y, yield; Q, quality; D, disease. 129 130 Ullrich—Production, Improvement, and Uses c08.indd 195 8/30/2010 8:28:56 PM F Barley: Production, Improvement, and Uses 196 Reinbergs 1985). In South Australia and Victoria, a backcross strategy utilizing molecular markers has been used to introduce disease resistance genes and boron tolerance into the malting cultivars Sloop and Gairdner without signiicantly changing their malting attributes (Barr et al. 2000, 2001). Mass selection strategies are relatively inexpensive methods of early-generation screening (Anderson and Reinbergs 1985). Australian programs using this technique have included New South Wales (B. Read, pers. comm.) and Queensland (Poulsen et al. 1997). Recurrent selection is an effective breeding strategy used in self-pollinating crops to improve complex characteristics such as yield, grain quality, and partial disease resistance (Stuthman et al. 1996). It involves repeated cycles of intercrossing and selection in a restricted gene pool. Carver and Bruns (1993) concluded a review paper by stating that “A review of genetic gains (∆G) reported for grain yield and related traits since 1985 indicates that recurrent selection has been equally, if not more, effective than traditional breeding methods.” While ∆G in traditional pedigree programs was estimated at 1% per year, recurrent selection programs with at least two completed cycles achieved ∆G of 3.4% per annum. In barley, a modiication of the technique has been used in conjunction with a system of genetic male sterility to make signiicant yield improvements in North American feed barleys (Falk 1996, 2001). DH production through either anther culture (Clapham 1973, cited by Kasha and Reinbergs 1982) or isolated microspore culture (Kasha et al. 1992) is used in Australia. However, DH production from the microspore-based techniques is more attractive because of improved recovery of DH lines, dificulties in managing Hordeum bulbosum plants, and spontaneous chromosome doubling, eliminating the need to use colchicine (Broughton and Poulsen 1997). The principal reason breeding programs use DH systems is to save time (Logue et al. 1993). As DH lines are, by nature, homogeneous, the need for several generations of self-pollination to achieve the same ends is eliminated. It is antici- F pated that 2–4 years can be saved from cultivar development cycles in the Australian spring barley breeding programs through the use of DH techniques. For example, the DH cv. Tantangara was released in 8 years as opposed to the conventional breeding cycle of 10–12 years in the New South Wales Agriculture barley program (B.J. Read, pers. comm.). Interspeciic and intersubspeciic hybrids have been used on several occasions to bring new genes into cultivated barley. Hordeum vulgare subsp. spontaneum has been used as a source of disease resistance genes in many breeding programs (Feuerstein et al. 1990; Abbott et al. 1991, 1992; Chicaiza et al. 1996). Translocations between H. bulbosum and H. vulgare subsp. vulgare have also been used as a source of new genetic variations for barley breeding (Xu and Kasha 1992; Pickering et al. 1995, 1998, 2000). The scald-resistant variety Tantangara, bred by Barbara Read at NSW-DPI Wagga Wagga, has a resistance gene from a H. vulgare spontaneum source (CPI-71283), 59 which was developed by Dr. Tony Brown (Brown et al. 1993, 2000) as a Clipper backcross (BC3) line, AB6 (Brown et al. 1988). Disease resistance research and breeding in Australia Australia has diverse barley cropping environments, and consequently, the occurrence of barley diseases is also diverse (Boyd and Dube 1989; Murray and Brennan 2001). The soil-borne diseases, crown rot and common root rot, are widespread throughout Australia and have become recognized in recent years as a national problem (Murray and Brennan 2001). Scald and CCN (H. avenae Woll.) have been major pathological constraints to barley production in southern Australia for many years, and resistance has been virtually mandatory in any new cultivars for the region. However, net blotch and other stubble-borne diseases have become increasingly signiicant throughout the entire continent in recent years (Murray and Brennan 2001). There are ive rust diseases capable of infecting cultivated barley; leaf rust (P. hordei), stem rust (P. graminis f. spp.), and barley grass yellow rust Ullrich—Production, Improvement, and Uses c08.indd 196 8/30/2010 8:28:56 PM Barley Breeding History, Progress, Objectives, and Technology (BGYR) (Puccinia striiformis) are the major cereal rust pathogens of barley in Australia. Barley stripe rust (P. striiformis f. sp. hordei) and crown rust (Puccinia coronata Corda) have not been recorded in Australian barley crops. Stripe rust is considered to be a signiicant quarantine threat to the Australian industry and preemptive breeding strategies have been developed. The sexual stages of P. hordei have been reported on “Star of Bethlehem” (Ornithogalum umbellatum) growing on the Yorke Peninsula in South Australia (Wallwork et al. 1992). The apparent sexual recombination of virulence genes was described in this work, following isolation of single leaf rust pustules collected from the area and differential testing on appropriate barley genotypes. This potential for genetic recombination of virulence factors and the consequent development of complex new pathotypes is likely to have a signiicant impact on resistance breeding and on the deployment of resistance genes. The Australian National Cereal Rust Control Program (NCRCP) conducts annual surveys of rusts in wheat, barley, oats, and other cereals (McIntosh et al. 1995). The information is used to monitor the development of new pathotypes, recommend withdrawal of susceptible cultivars, and strategically deploy resistance genes for the most effective control of the diseases. Severe localized epidemics of barley leaf rust have been reported in Australia (Dill-Macky et al. 1989) associated with a predominance of leaf rustsusceptible cultivars. In Australia, leaf rust is a signiicant disease of barley in southern Queensland, in northern New South Wales, and in parts of South Australia, West Australia, and Tasmania (Murray and Brennan 2001). The sporadic nature of Australian leaf rust epidemics is due to climate variability and luctuation in the relative production areas of susceptible cultivars. The most recent epidemics of barley leaf rust in Queensland and in northern New South Wales occurred in 1978, 1983, 1984, and 60 1988 (•• Rees, pers. comm.). The 1988 epidemic resulted from the development of two new pathotypes (210P+ and 253P−) with virulence to the resistance gene in the predominant cultivar, Grimmett (Cotterill et al. 1994), causing substan- 197 tial yield losses of up to 30% (Dill-Macky et al. 1989). Two new and potentially damaging pathotypes of leaf rust (4610P+ and 5610P+) with virulence on most major cultivars were irst identiied in Tasmania during 1991 (Cotterill et al. 1992) 61 and in West Australia in 1997 associated with the varieties Triumph and Franklin (Loughman 1998). Park and Williams (2000) reported detection of the pathotypes in the northern region during 1999 and that they made up over 60% of the barley leaf rust isolates collected from across Australia in the annual rust survey of that season. The pathotypes are virulent on most Australian barley cultivars and have caused economic damage to crops in Western Australia and South Australia. Until 1999, Rph12 was effective against all pathotypes of leaf rust found in Queensland and Northern New South Wales. The irst Australian report of virulence to Rph12 was conirmed with pustules isolated from a Tasmanian crop of Franklin barley during 1991 (Cotterill et al. 1992). The new pathotype spread to South Australia by 1993 (H. Wallwork, pers. comm.). In 1998, a different pathotype, also with Rph12 virulence, was reported from Western Australia (Loughman 1998). The two pathotypes designated 4610P+ and 5610P+ by octal notation were identiied in South Queensland during 1999 (G. Platz, pers. comm.). Rph12, derived from European cv. Triumph, is present in barley cv. Tallon (Cotterill et al. 1994) and Lindwall (R.G. Rees, pers. comm.). Both of these are major commercial cultivars grown in the Queensland and northern New South Wales winter cropping areas, and thus, the northern region barley crop is at greater risk from the disease. The release of Gairdner and later Baudin, derived from either “Franklin sib” or Franklin and both of which carry the Rph12 gene, has further exacerbated the problem of having a longer-season, higher-rainfall barley crop with a high proportion of susceptibility. Backcrossing has been one of the more widely used resistance breeding strategies in barley. This is because the technique is simple and has a high probability of developing well-adapted cultivars. Backcross lines of Prior and Research were developed by the Waite Institute breeding program Ullrich—Production, Improvement, and Uses c08.indd 197 8/30/2010 8:28:56 PM F Barley: Production, Improvement, and Uses 198 during the 1940s and were used as parents (Pugsley and Vines 1946; Pugsley 1951). Later, Western Australian breeders used backcrossing to introduce net blotch resistance genes from unadapted material from Ethiopia, Manchuria, and Turkey into locally adapted genotypes (Boyd et al. 1981). CCN resistance is expressed as a dominant trait, and a limited backcrossing approach has been utilized in both South Australian and Victorian programs producing such varieties as Galleon, Chebec, Sloop-SA, and Sloop-Vic. Recurrent selection is another strategy that can be employed in barley to develop disease-resistant germplasm (Sharp 1985). However, as opposed to backcrossing, recurrent selection techniques increase general resistance levels through several cycles of recombination and selection in which genes of minor effect are accumulated. Major genes must not be present for recurrent selection to work effectively, as they mask the minor gene effects and interfere with the selection process. In barley, recurrent selection has been successfully used to improve resistance to leaf rust and powdery mildew (Parlevliet and van Ommeren 1988). The MSFRS system developed by Falk (1996) is now used routinely in the BBA-West program utilizing both genotypic and phenotypic screening methods. Genetic male steriles were irst used in composite crosses (CC XIV) to facilitate hybridization (Ramage 1987 loc. cit.). Ramage gave Eslick the credit for coining the term “male sterile facilitated recurrent selection” (Ramage 1975). It would seem that the best breeding strategy for long-term control of barley leaf rust would combine most of the systems described in the two previous sections because of the diversity of leaf rust pathotypes and the potential for sexual recombination of virulence genes in the pathogen. Australian barley pathologists and breeders have had to deal with the problem that the alternate host for barley leaf rust, Star of Bethlehem (O. umbellatum), is present on the southern end of Yorke Peninsula in South Australia (Wallwork et al. 1992) and is probably responsible for outbreaks of leaf rust in the region. F The nBMMP and AWCMMP In Australia, researchers participating in the Australian NBMMP (Langridge 1997; Langridge and Barr 2003, “Better Barley Faster: the Role of Marker Assisted Selection” loc. cit.) produced maps for 10 major populations and conducted marker discovery in 44 minor populations (Barr et al. 2001). The major populations were selected to represent Australian genotypes and key parent lines. They were Alexis × Sloop (DH/RIL) (Barr et al. 2003a), Amagi Nijo × WI-2585 (Pallotta et al. 2003), Barque*2 × H. vulgare subsp. spontaneum 71284-45 (advanced backcross quantitative trait locus [ABQTL]) (J. Eglinton, pers. comm.), Chebec x Harrington (DH) (Langridge et al. 1995; Barr et al. 2003b), Clipper × Sahara (Karakousis et al. 2003b), Galleon × Haruna Nijo (DH) (Langridge et al. 1995; Karakousis et al. 2003c), Mundah × Keel, (Long et al. 2003), Sloop × Halcyon (DH) (Read et al. 2003), Tallon × Kaputar (DH) (Cakir et al. 2003b), Tallon × Scarlett (DH) (D. Poulsen, unpublished data), VB9104 × Dash (DH) (D. Moody, unpublished data), VB9524 × ND11231 (DH) (Emebiri et al. 2003), and Baudin × AC-Metcalfe (DH) (R. Lance, pers. comm.). Maps of Chebec × Harrington and Galleon × Haruna Nijo have been published on the Graingenes Web site along with a Clipper × Sahara map (http://wheat. pw.usda.gov/ggpages/maps.shtml). One of the NBMMP objectives was to utilize and improve new technologies for Australian barley molecular geneticists. Simple sequence repeat (SSR) markers were introduced into the Australian barley research groups (Ablett et al. 2003; Karakousis et al. 2003a), and their application improved the eficiency and utility of markers in breeding programs. The availability of a substantial number of markers common across individual barley maps enabled the alignment of markers and the creation of consensus maps of the barley genome. Mapping data from six populations were combined to produce the irst consensus map of 587 markers (Langridge et al. 1995). The majority of the markers were obtained from the Steptoe × Morex, 62 63 64 65 Ullrich—Production, Improvement, and Uses c08.indd 198 8/30/2010 8:28:56 PM Barley Breeding History, Progress, Objectives, and Technology Igri × Franka, and Proctor × Nudinka maps but were linked together by common markers used to map three Australian populations: Clipper × Sahara 3771, Haruna Nijo × Galleon, and Chebec × Harrington. Overall, the consistency in gene order across the six original maps was high. Discrepancies only occurred where markers were tightly linked and were subject to the random error normal to mapping experiments. A second consensus map was constructed by Qi et al. (1996) from the Proctor × Nudinka, Igri × Franka, Steptoe × Morex, and Harrington × TR306 maps. The most recent barley consensus maps were constructed by Karakousis et al. (2001, 66 2003d) using RFLP, AFLP, and SSR markers. Among the early successes of the NBMMP were the mapping of genes and markers for resistance to CCN, a major soil-borne parasite of barley in southern production areas. Kretschmer et al. (1997) reported the association of a single dominant CCN resistance gene, Ha2, with lanking RFLP markers. The gene mapped to chromosome 2H. Molecular analysis of CCN-resistant Australian cultivars demonstrated that all possessed the Ha2 gene. A second gene for CCN resistance, designated Ha4, was mapped to chromosome 5H in Australian cv. Galleon (Barr et al. 1998). RFLP markers for both genes have been used for selection in the South Australian program (Barr et al. 2001). QTLs for genes associated with high malt extract were identiied and validated in populations between Australian and overseas barleys (Collins et al. 2003). QTLs controlling kernel discoloration were described by Li et al. (2003a). This information has been invaluable to select for a dificult to measure trait associated with weather damage at grain maturation and prior to harvest. Similarly, QTLs controlling preharvest sprouting have been identiied and implemented in breeding programs (Li et al. 2003b). Box et al. (1997) reported evaluation of markers with putative linkage to the gene for hulless grain (nud) on chromosome 7H, and marker assays were ultimately developed (A. Box, pers. comm.). Markers for the hulless trait enable the more eficient selection of progeny as heterozygotes from various crossing situations. 199 Further studies identiied QTLs affecting coleoptile length on 7H and 5H in Proctor × Nudinka and Galleon × Haruna Nijo, respectively (Box and Barr 2000). Four QTLs inluencing response to boron toxicity were mapped in Clipper × Sahara 3771 (Jefferies et al. 1999). Boron toxicity is a major problem for barley crops in low-rainfall areas in southern Australia. Two QTLs, located on chromosomes 4H and 6H, were associated with reduced boron uptake. The 4H QTLs also appeared to affect root length, dry matter production, and symptom expression. Leaf symptom expression was affected by another QTL on 2H, and suppression of root growth by high boron levels was inluenced by a QTL on 3H. In another series of molecular studies of nutrient–plant relationships, a region on chromosome 4HL was identiied as contributing to aluminum/acid soil tolerance in several genotypes (Raman et al. 2001, 2002, 2003). The Australian data matched a locus mapped elsewhere (Tang et al. 2000). Raman et al. (2001) identiied closely linked SSR markers for the trait, with the intent of developing an alternative to biological screening assays. Mel1, a gene affecting the eficiency of manganese uptake in barley, was mapped to the distal portion of chromosome 4HS (Pallotta et al. 2000, 2003). BSA of F2 plants from the cross Amagi 67 Nijo × WI-2585 was used to identify several closely linked RFLP markers for the trait, which was inherited from Amagi Nijo. Assessment of 95 genotypes used as parents in the South Australian breeding program indicated that two RFLP markers could be used to distinguish the presence of the Amagi Nijo locus in all cases. Consensus mapping was used to identify 62 SSR markers linked by less than 10 cM to 14 traits of interest to most Australian barley breeding programs (Karakousis et al. 2000). The traits included resistance to powdery mildew (mlo), net blotch (Rpt4), barley yellow dwarf virus (BYDV) (Yd2) and CCN (Ha4). The markers were validated with populations from the NBMMP and successfully predicted plant phenotypes. Further analysis with breeding lines from the South Australian Barley Improvement Ullrich—Production, Improvement, and Uses c08.indd 199 8/30/2010 8:28:56 PM F Barley: Production, Improvement, and Uses 200 F Program demonstrated that allelic variation was suficient for the majority of the markers to be used for routine screening. 68 Coventry et al. (2001, 2003a,b) examined QTLs associated with yield and yield components in the Australian mapping populations Alexis × Sloop, Chebec × Harrington, and Mundah × Keel. Several yield and yield component QTLs were coincident with chromosomal regions known to be associated with phenology and development, including the 2H Ppd-H1 photoperiod response gene and another locus on the same chromosome. The denso gene is present in the Alexis × Sloop population and was associated with major effects on grain size, yield, and plant height. The phenological development patterns of Australian barleys are an important aspect of adaptation to Mediterranean and East Coast environments. Boyd et al. (2003) enhanced the classical understanding by investigating the molecular genetic control of the basic vegetative period or earliness per se and photoperiod response as they apply to adaptation in the Australian context. Apart from the loci discussed above, the NBMMP mapped and developed markers for a signiicant number of agronomic, quality, and disease resistance characteristics. A mixed model approach to better improve the QTL analyses from multienvironment trials was investigated by biometricians (Verbyla et al. 2003). Quality research with respect to molecular approaches was reviewed by Fox et al. (2003). Furthermore, biometrics analyses of quality within breeding programs and through quality laboratories from a strategic point of view were studied by Cullis 69 et al. (2003). Other traits targeted by minor populations included grain staining, acid soil tolerance, preharvest sprouting, boron tolerance, grain size, and water sensitivity. Additional work conducted as part of the NBMMP has been the validation of markers for routine use in Australian breeding programs. For example, Collins et al. (2001) reported the validation of QTLs associated with the expression of malt extract, DP, α-amylase, and β-amylase in six representative breeding populations from the South Australian breeding program. The work clearly demonstrated that QTLs on chromosomes 1H, 2H, and 5H could be used to improve malting quality through the implementation of MAS. Mapping barley disease resistance genes in Australia As a demonstration of the scope of the NBMMP activities, disease resistance loci targeted in the major mapping populations are described by Williams (2003) and compared with other known genes for resistance. In a more recent phase of barley improvement in Australia, signiicant attention has been placed on the development of elite malting and feed barleys with multiple disease resistances pyramided from complex crosses. Minor mapping populations were also constructed by the NBMMP and were analyzed by BSA to identify markers linked to resistance to scald (Genger et al. 2003), powdery mildew (Paris et al. 2003), spot-type net blotch (Williams et al. 2003), net-type net blotch (Cakir et al., 2003; 70 Gupta et al. 2003; Raman et al. 2003), common root rot, stem rust, leaf rust (Park et al. 2003), crown rot, covered smut, spot blotch, and the exotic quarantine diseases, such as stripe or yellow rust (Cakir et al. 2003c). Scald (R. secalis) has also been a signiicant target for marker development as both the pathotype structure and the genetics of resistance are complex, making conventional selection techniques dificult. Rh, a gene conferring resistance to scald, was mapped to chromosome 3HL by BSA with RAPD markers in a set of near-isogenic 71 lines (Barua et al. 1993). It has been suggested that the 3HL locus, now designated Rrs1, is multiallelic as is scald resistance in several other unrelated genotypes and has been mapped to the same location (Williams et al. 2001). One of the alleles confers resistance to 22 of the 23 known Australian R. secalis pathotypes. The multiallelic nature of Rrs1 has signiicant strategic implications in designing crosses to combine scald resistance genes. Pathotype-speciic seedling resistance to nettype net blotch was mapped to chromosome 6H in a DH population derived from the Australian cross Tallon × Kaputar (Cakir et al. 2001, 2003a). Ullrich—Production, Improvement, and Uses c08.indd 200 8/30/2010 8:28:56 PM Barley Breeding History, Progress, Objectives, and Technology The resistance was inherited from Kaputar, a 72 reselection of the CIMMYT cv. Arupo. Further work is being carried out on the population to identify the genetic location of adult plant resistance to net-type net blotch inherited from Tallon. A dominant resistance gene to spot-type net blotch from Australian barley cv. Galleon was mapped to chromosome 7H and was designated Rpt4 (Williams et al. 1999). Flanking RFLP markers were identiied for the locus. Validation of the marker demonstrated greater than 90% accuracy in predicting the occurrence of resistance to spot-type net blotch. Karakousis et al. (2000) subsequently identiied SSR markers in the vicinity of Rpt4, which have since been used for selection in at least three Australian barley breeding programs. Other reports of marker development for barley disease resistance genes include identiication of speciic markers and/or QTL for resistance to BYDV (Collins et al. 1996; Paltridge et al. 1998). Stripe, leaf, and stem rust resistance genes have all been targets for marker development and genetic study in barley. In general, the development of markers has been seen as leading toward the implementation of better breeding strategies to develop long-term control of the diseases. The information obtained in these studies has contributed to the development of a greater depth of knowledge of the genetics and mechanisms of rust resistance. Resistance to South American stripe rust pathotypes was mapped to chromosomes 2H and 5H in Australian cv. Tallon (Cakir et al. 2001, 2003c). The data were obtained from a ield assessment of the NBMMP Tallon × Kaputar DH population at CIMMYT. Stripe rust resistance was also mapped in the NBMMP Arapiles × Franklin mapping population, with the resistance derived from Franklin (Cakir et al. 2003c). The Franklin resistance was also located on 2H and 5H. The data suggested that the two regions were common to both cultivars. It was suspected that both Tallon and Franklin had inherited the same resistance gene from Triumph, situated on chromosome 5H. The location of the QTL was not a perfect match for the 5H resis- 201 tance mapped by Chen et al. (1994), and further study was required to determine the relationship between the loci. The resistance QTL on 2H from Tallon and Franklin appeared to be located in the vicinity of a gene related to plant maturity (Cakir et al. 2003). It therefore appeared that differences in maturity may have inluenced the expression of resistance in the mapping populations at the time of observation. Several leaf rust resistance genes from the Rph series have been mapped and linked to molecular markers including Rph2, Rph9/12, and Rph16 (Borovkova et al. 1997a,b, 1998; Graner et al. 2000). Poulsen et al. (1995a) identiied an RAPD marker linked to the RphQ resistance gene from barley accession Q21861. RFLP and STS markers 73 were subsequently used by Borovkova et al. (1997a,b) to map RphQ to the centromeric region of chromosome 5H, in the same region as Rph2, and it was postulated on the basis of marker and phenotypic data that Rph2 and RphQ were allelic. Nine RAPD markers, two RFLP markers, CDO749 and Rrn2, and one STS marker, ITS1, were identiied as closely linked to RphQ 74 (Borovkova et al. 1997). Molecular genetic marker implementation in Australian barley breeding programs One of the major dificulties encountered in the implementation of marker-assisted breeding in Australian barley breeding programs has been the initial availability of only a limited number of suitable markers. Although RFLP markers were used fairly extensively for CCN resistance and boron tolerance by the Waite Institute program in the late 1990s (A. Barr, pers. comm), other Australian programs have taken longer to routinely adopt the technology. This has been partly due to time requirements of developing and validating suitable PCR markers. Most Australian 75 programs are not fully equipped for the largescale operation of RFLP systems. However, with the maturation of the NBMMP, the situation is changing and all of the programs are reaching the stage where routine marker implementation can occur. Ullrich—Production, Improvement, and Uses c08.indd 201 8/30/2010 8:28:57 PM F Barley: Production, Improvement, and Uses 202 The use of markers in Australian barley breeding programs was succinctly described in two review papers (Barr et al. 2000, 2001). The majority of those markers were developed and/or validated within the NBMMP strategic initiative. It was also reported that MAS had been used in Australian breeding programs to select for malt extract QTL on chromosomes 1H, 2H, and 5H (Barr et al. 2001). In a particularly elegant example, the South Australian program had used selection for these loci in conjunction with a “defect elimination” backcross breeding strategy to develop improved versions of the cultivars Sloop (Barr et al. 2000) and Gairdner (Barr et al. 2001). Markers were used to assist recovery of the recurrent parent genomes while tracking genes for BYDV resistance (yd2), spot-type net blotch resistance (Rpt4), and CCN resistance (Ha2). After the single genes had been introgressed, selected lines were intercrossed with the aim of developing versions of the cultivars possessing all of the target genes. However, the highest rate of usage of MAS in Australian barley breeding has been the enrichment of complex crosses by the South Australian program (Barr et al. 2000). By selecting F1 plants with markers linked to a range of speciic traits, the program is making more eficient use of material derived from three- and four-way crosses. Only progeny lines with desired combinations of loci are selected for ield assessment. F1 plants from complex crosses requiring additional crossing can also be identiied. The new malting variety Flagship from the University of Adelaide program, accredited in 2006, is a testimony to the eficiency and expediency of utilizing this approach. Marker use for the selection of QTL is also progressing in the Australian breeding programs. Collins et al. (2000, 2001) validated QTL related to malt extract, DP, α-amylase, and β-amylase and recommended markers for routine use in the southern region breeding programs. Validation of the markers in the northern and western region programs has also commenced. It is anticipated that the adoption of marker technology by the Australian barley breeding programs will rapidly increase over the next few F years, especially with increased identiication of PCR-based markers linked to traits of interest to the individual programs. REfEREnCEs ABB Grain. 2008. Available at http://www.abb.com.au/ or http://ezigrain.abb.com.au/home/home.asp. Abbott, D.C., J.J. Burdon, A.M. Jarosz, A.H.D. Brown, W.J. Muller, and B.J. Read. 1991. The relationship between seedling infection types to leaf scald in Clipper barley backcross lines. Aust. J. Agric. Res. 42:801–809. Abbott, D.C., A.H.D. Brown, and J.J. Burdon. 1992. Genes for scald resistance from wild barley (Hordeum vulgare ssp. spontaneum) and their linkage to isozyme markers. Euphytica 61:225–231. Ablett, G.A., A. Karakousis, L. Banbury, M. Cakir, T.A. Holton, P. Langridge, and R.J. Henry. 2003. Application of SSR markers in the construction of Australian barley genetic maps. Aust. J. Agric. Res. 54:1187–1195. Anderson, M.K. and E. Reinbergs. 1985. Barley breeding. pp. 231–268. In D.C. Rasmusson (ed.). Barley. American Society of Agronomy, Series: Agronomy Monograph No. 26. American Society of Agronomy, ••. Armitt, J. and H. Way. 1990. Breeding and assessment of new malting barleys in Australia. pp. 57–64. Proceedings of the 21st Convention of the Institute of Brewing, Australia and New Zealand Section, Auckland, New Zealand, 1990. Barley Australia. 2008. Available at http://www. barleyaustralia.com.au/. Barley Breeding Australia. 2008. Available at http:// www.barleybreedingaustralia.com.au/. Barr, A.R., K.J. Chalmers, A. Karakousis, J.M. Kretschmer, S. Manning, R.C.M. Lance, J. Lewis, S.P. Jeffries, and P. Langridge. 1998. RFLP mapping of a new cereal cyst nematode resistance locus in barley. Plant Breed. 117: 185–187. Barr, A.R., S.P. Jefferies, P. Warner, D.B. Moody, K.J. Chalmers, and P. Langridge. 2000. Marker assisted selection in theory and practice. pp. 167–178. Proceedings of the 8th International Barley Genetics Symposium, Adelaide, Australia, 2000. Barr, A., J. Eglinton, P. Langridge, P. Warner, and K. Chalmers. 2001. Marker assisted selection—where to now? In Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia, •• ••, ••. Available at http://www.regional.org.au/au/abts/2001/m2/barr. htm#TopOfPage. Barr, A.R., S.P. Jefferies, S. Broughton, K.J. Chalmers, J.M. Kretschmer, W.J.R. Boyd, H.M. Collins, S. Roumeliotis, S.J. Logue, S.J. Coventry, D.B. Moody, B.J. Read, D. Poulsen, R.C.M. Lance, G.J. Platz, R.F. Park, J.F. Panozzo, A. Karakousis, P. Lim, A.P. Verbyla, and P.J. Eckermann. 2003a. Mapping and QTL analysis of the barley population Alexis × Sloop. Aust. J. Agric. Res. 54: 1117–1123. Barr, A.R., A. Karakousis, R.C.M. Lance, S.J. Logue, S. Manning, K.J. Chalmers, J.M. Kretschmer, W.J.R. Boyd, 76 77 78 79 Ullrich—Production, Improvement, and Uses c08.indd 202 8/30/2010 8:28:57 PM Barley Breeding History, Progress, Objectives, and Technology 80 81 82 83 H.M. Collins, S. Roumeliotis, S.J. Coventry, D.B. Moody, B.J. Read, D. Poulsen, C.-D. Li, G.J. Platz, P.A. Inkerman, J.F. Panozzo, B.R. Cullis, A.B. Smith, P. Lim, and P. Langridge. 2003b. Mapping and QTL analysis of the barley population Chebec × Harrington. Aust. J. Agric. Res. 54:1125–1130. Barua, U.M., K.J. Chalmers, C.A. Hackett, W.T.B. Thomas, W. Powell, and R. Waugh. 1993. Identiication of RAPD markers linked to a Rhynchosporium secalis resistance locus in barley using near-isogenic lines and bulked segregant analysis. Heredity 71:177–184. Borovkova, I.G., Y. Jin, and B.J. Steffenson. 1997a. Mapping of the leaf rust resistance genes Rph9 and Rph12 in barley. Plant and Animal Genome V, poster abstract P152. Available at http://www.intl-pag.org/pag/5/abstracts/ p-5c-152.html. Borovkova, I.G., Y. Jin, B.J. Steffenson, A. Kilian, K.T. Blake, and A. Kleinhofs. 1997b. Identiication and mapping of a leaf rust resistance gene in barley line Q21861. Genome 40:236–241. Borovkova, I.G., Y. Jin, and B.J. Steffenson. 1998. Chromosomal location and genetic relationship of leaf rust resistance genes Rph9 and Rph12 in barley. Phytopathology 88:76–80. Bowman, J. and T. Blake. 1996. Barley feed quality for beef cattle. pp. 82–90. In G. Scoles and B. Rossnagel (eds.). Proceedings of the 5th International Oat Conference and 7th International Barley Genetics Symposium, Vol. 1. ••, Saskatoon, Saskatchewan, Canada. Box, A.J. and A.R. Barr. 2000. Identiication of molecular markers associated with improved coleoptile length in hulless barley. pp. 11–13. In S. Logue (ed.). Proceedings of the 8th International Barley Genetics Symposium, Vol. 3. ••, Adelaide, Australia. Box, A.J., A.R. Barr, and P. Langridge. 1997. The application of molecular markers to facilitate the rapid selection of hulless barley progeny. pp:56–59. In •• •• (ed.). Proceedings of the 8th Australian Barley Technical Symposium, Vol. 3. ••, Gold Coast, Australia. Boyd, W.J.R. and A. Dube (eds.). 1989. Barley diseases survey: importance, research status and support. A survey carried out in conjunction with the Second Barley Diseases Workshop, Adelaide, South Australia, October 26 and 27, 1989. Boyd, W.J.R., D. Janakiram, T.N. Khan, and P.A. Portman. 1981. Breeding for resistance to scald in barley. pp 118– 133. In •• •• (ed.). Barley diseases in Australasia: Proceedings of a Workshop held at Adelaide, October 17–19, 1979. ••, ••. Boyd, W.J.R., C.-D. Li, C.R. Grime, M. Cakir, S. Potipibool, L. Kaveeta, S. Men, M.R. Jalal Kamali, A.R. Barr, D.B. Moody, R.C.M. Lance, S.J. Logue, H. Raman, and B.J. Read. 2003. Conventional and molecular genetic analysis of factors contributing to variation in the timing of heading among spring barley (Hordeum vulgare L.) genotypes grown over a mild winter growing season. Aust. J. Agric. Res. 54:1277–1301. Broughton, S. and D.M.E. Poulsen. 1997. An overview of the Hordeum bulbosum method and its potential for Australian barley breeding programs. pp 2:9.11–2:9.19. Proceedings 203 of the 8th Australian Barley Technical Symposium, Gold Coast, Queensland, Australia, 1997. Brown, A.H.D., J. Munday, and R.N. Oram. 1988. Use of isozyme-marked segments from wild barley (Hordeum spontanaeum) in barley breeding. Plant Breed. 100: 280–288. Brown, A.H.D., D.C. Abbott, J.J. Burdon, B.J. Read, and D.B. Moody. 1993. Use of gene tags in developing barley with multiple scald resistances. pp 83–84. Proceedings of the 6th Australian Barley Technical Symposium, Launceston, Tasmania, 1993. Brown, A.H.D., J.J. Burdon, R.K. Genger, D.C. Abbott, J.S. Brown, G.B. Wildermuth, and P.M. Banks. 2000. Wild barley (Hordeum vulgare ssp. spontaneum) as a source of disease resistance for barley breeding. pp. 56–58. In S. Logue (ed.). Barley Genetics VIII: Proceedings of the 8th International Barley Genetics Symposium, Adelaide, South Australia, October 22–27, 2000, Vol. 1. ••, ••. Cakir, M., N. Galwey, D. Poulsen, G. Platz, R. Johnston, G. Ablett, C. Wellings, and H. Vivar. 2001. Mapping genes for resistance to net form net blotch and stripe rust in barley (Hordeum vulgare L.). Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia, •• ••, ••. Available at http://www.regional.org.au/ au/abts/2001/t3/cakir.htm#TopOfPage. Cakir, M., S. Gupta, G.J. Platz, G.A. Ablett, R. Loughman, L.C. Emebiri, D. Poulsen, C.-D. Li, R.C.M. Lance, N.W. Galwey, M.G.K. Jones, and R. Appels. 2003a. Mapping and validation of the genes for resistance to Pyrenophora teres f. teres in barley (Hordeum vulgare L.). Aust. J. Agric. Res. 54:1369–1377. Cakir, M., D. Poulsen, N.W. Galwey, G.A. Ablett, K.J. Chalmers, G.J. Platz, R.F. Park, R.C.M. Lance, J.F. Panozzo, B.J. Read, D.B. Moody, A.R. Barr, P. Johnston, C.-D. Li, W.J.R. Boyd, C.R. Grime, R. Appels, M.G.K. Jones, and P. Langridge. 2003b. Mapping and QTL analysis of the barley population Tallon × Kaputar. Aust. J. Agric. Res. 54:1155–1162. Cakir, M., M. Spackman, C.R. Wellings, N.W. Galwey, D.B. Moody, D. Poulsen, F.C. Ogbonnaya, and H. Vivar. 2003c. Molecular mapping as a tool for pre-emptive breeding for resistance to the exotic barley pathogen, Puccinia striiformis f. sp. hordei. Aust. J. Agric. Res. 54: 1351–1357. Carver, B.F. and R.F. Bruns. 1993. Emergence of alternative breeding methods for autogamous crops. pp. 43–56. Proc. of the 10th Australian Plant Breeding Conference, Gold Coast, Queensland, Australia, 1993. CBH Group. 2008. Harvest Handbook—an Important Information Guide to Assist Growers and Carters before, during and after Delivering Grain to CBH Group Receival Points—2007–08 Season. ••, ••. Chen, F.Q., D. Prehn, P.M. Hayes, D. Mulrooney, A. Corey, and H. Vivar. 1994. Mapping genes for resistance to barley stripe rust (Puccinia striiformis f. sp. hordei). Theor. Appl. Genet. 88:215–219. Chicaiza, O., J.D. Franckowiak, and B.J. Steffenson. 1996. New sources of resistance to leaf rust in barley. pp. 706– 708. In G. Scoles and B. Rossnagel (eds.). Proceedings of 84 85 86 87 Ullrich—Production, Improvement, and Uses c08.indd 203 8/30/2010 8:28:57 PM F Barley: Production, Improvement, and Uses 204 F the 5th International Oat Conference and 7th International Barley Genetics Symposium. Poster Sessions, Vol. 2. ••, 88 Saskatoon, Saskatchewan, Canada. Clapham, D. 1973. Haploid Hordeum plants from anthers in vitro. Planzenzuchtg 69:142–145. Cited by Kasha, K.J. and E. Reinbergs. 1982. Collins, H., S. Logue, S. Jefferies, and A. Barr. 2001. Validation of markers for malt extract, DP, alpha-amylase and beta-amylase. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia. Available at http://www.regional.org.au/au/abts/2001/ w2/collins.htm#TopOfPage. Collins, H.M., S.J. Logue, S.P. Jefferies, and A.R. Barr. 2000. Using QTL mapping to improve our understanding of malt extract. pp. 225–227. Proceedings of the 8th International Barley Genetics Symposium, Adelaide, 2000. Collins, H.M., J.F. Panozzo, S.J. Logue, S.P. Jefferies, and A.R. Barr. 2003. Mapping and validation of chromosome regions associated with high malt extract in barley (Hordeum vulgare L.). Aust. J. Agric. Res. 54:1223–1240. Collins, N.C., N.G. Paltridge, C.M. Ford, and R.H. Symons. 1996. The Yd2 gene for barley yellow dwarf virus resistance maps close to the centromere on the long arm of barley chromosome 3. Theor. Appl. Genet. 92:858–864. Cotterill, P.J., R.G. Rees, and W.A. Vertigan. 1992. Detection of Puccinia hordei virulent on the Pa9 and Triumph resistance genes in barley in Australia. Australas. Plant Pathol. 21;32–34. Cotterill, P.J., R.G. Rees, and G.J. Platz. 1994. Response of Australian cultivars of barley to leaf rust (Puccinia hordei). Australas. J. Exp. Agric. 34:783–788. Coventry, S., A. Barr, J. Eglinton, and G. McDonald. 2001. Characterisation of mapping parents and identiication of the genes involved in the yield and grain weight of barley (Hordeum vulgare L.) grown under Mediterranian environments. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia. Available at http:// www.regional.org.au/au/abts/2001/t3/coventry. htm#TopOfPage. Coventry, S.J., A.R. Barr, J.K. Eglinton, and G.K. McDonald. 2003a. The determinants and genome locations inluencing grain weight and size in barley (Hordeum 89 vulgare L.). Aust. J. Agric. Res. 54:1103–1115. Coventry, S.J., H.M. Collins, A.R. Barr, S.P. Jefferies, K.J. Chalmers, S.J. Logue, and P. Langridge. 2003b. Use of putative QTLs and structural genes in marker assisted selection for diastatic power in malting barley (Hordeum vulgare L.). Aust. J. Agric. Res. 54:1241–1250. Davies, R.A.H. 1993. GRDC grains research initiatives, particularly in malting barley. pp. 168–172. Proceedings of the 6th Australian Barley Technical Symposium, Launceston, Tasmania, 1993. Dill-Macky, R. 1992. The epidemiology and management of stem rust in barley in north-eastern Australia. PhD thesis, University of Queensland, Brisbane, Australia. Dill-Macky, R., R.G. Rees, R.P. Johnston, G.J. Platz, and A. Mayne. 1989. Stem and leaf rusts of barley. pp. 38–40. In •• •• (ed.). Queensland Wheat Research Institute Biennial Report 1984–1986, Queensland Department of Primary 90 Industries, Toowoomba, Australia. Ellis, S.E. 1983. Review of the Australian barley breeding programs—Victoria. pp. 12–15. Australian Barley Technical Symposium Report, Perth, Western Australia, 1983. E-malt.com. 2008. Available at http://e-malt.com/index. htm. Emebiri, L.C., D.B. Moody, J.F. Panozzo, K.J. Chalmers, J.M. Kretschmer, and G.A. Ablett. 2003. Identiication of QTLs associated with variations in grain protein concentration in two-row barley. Aust. J. Agric. Res. 54: 91 1211–1221. Falk, D.E. 1996. Use of recurrent introgressive population enrichment (RIPE) for utilisation of plant genetic resources in a barley breeding program. pp. 167–169. In A. Slinkard, G. Scoles, and B. Rossnagel (eds.). Proceedings of the 5th International Oat Conference and 7th International Barley Genetics Symposium. Poster Sessions, Vol. 1. ••, 92 Saskatoon, Saskatchewan, Canada. Falk, D.E. 2001. Recurrent introgression as a population enrichment (RIPE) method in barley. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia. Available at http://www.regional.org.au/au/ abts/2001/t3/falk.htm#TopOfPage. Feuerstein, U., A.H.D. Brown, and J.J. Burdon. 1990. Linkage of rust resistance genes from wild barley (Hordeum spontaneum) with isozyme markers. Plant Breed. 104: 318–324. Finlay, K.W. and G.N. Wilkinson. 1963. The analysis of adaptation in a plant-breeding program. Aust. J. Agric. Res. 14:742–754. Fox, G.P., J.F. Panozzo, C.D. Li, R.C.M. Lance, P.A. Inkerman, and R.J. Henry. 2003. Molecular basis of barley quality. Aust. J. Agric. Res. 54:1081–1101. French, R.J. and J.E. Schultz. 1984a. Water use eficiency of wheat in a Mediterranean-type environment. I. The relationship between yield, water use and climate. Aust. J. 93 Agric. Res. 35:743–764. French, R.J. and J.E. Schultz. 1984b. Water use eficiency of wheat in a Mediterranean-type environment. II. Some limitations to eficiency. Aust. J. Agric. Res. 35:765–775. Fukuda, K., S. Takahashi, and Y. Aida. 1999. Working together: a Japanese brewing company’s barley breeding strategy. pp. 2.14.1–2.14.8. Proceedings of the 9th Australian Barley Technical Symposium, Melbourne, Victoria, Australia, 1999. Genger, R.K., K.J. Williams, H. Raman, B.J. Read, H. Wallwork, J.J. Burdon, and A.H.D. Brown. 2003. Leaf scald resistance genes in Hordeum vulgare and Hordeum vulgare ssp. spontaneum: parallels between cultivated and wild barley. Aust. J. Agric. Res. 54:1335–1342. Gilmour, R.F. 1993. The Western Region barley industry development program. pp. 173–183. Proceedings of the 6th Australian Barley Technical Symposium, Launceston, Tasmania, 1993. Grainco. 2008. Speciications for Acceptance of Malting Barley. Grainco, Toowoomba, Queensland, Australia. Grains Research and Development Corporation (GRDC). 2001. Annual Report 2000-01. Available at http:// www.grdc.com.au/. Graner, A., W. Michalek, and S. Streng. 2000. Molecular mapping of genes conferring resistance to viral and fungal Ullrich—Production, Improvement, and Uses c08.indd 204 8/30/2010 8:28:57 PM Barley Breeding History, Progress, Objectives, and Technology 94 95 96 97 98 pathogens. pp. 45–52. In S. Logue (ed.). Barley Genetics VIII: Proceedings of the 8th International Barley Genetics Symposium, Adelaide, South Australia, October 22–27, 2000, Vol. 1. ••, Adelaide, South Australia. Gupta, S., R. Loughman, G.J. Platz, and R.C.M. Lance. 2003. Resistance in cultivated barleys to Pyrenophora teres f. teres and prospects of its utilisation in marker identiication and breeding. Aust. J. Agric. Res. 54: 1379–1386. Hawthorne, D. 1999. Role of the malting and brewing industry barley technical committee. pp. 2.33.1–2.33.5. Proceedings of the 9th Australian Barley Technical Symposium, Melbourne, Victoria, Australia, 1999. Healy, P. 2001. Malting and brewing industry barley technical committee—future directions. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia. Available at http://www.regional.org.au/au/ abts/2001/m2/healy.htm#TopOfPage. Hirsch, M. 1985. The southern Australian disease situation. pp. 91–95. Proceedings of the 2nd Australian Barley Technical Symposium, Toowoomba, Queensland, 1985. Jefferies, S.P., A.R. Barr, A.K.M. Islam, A.B. Frensham, A. Karakousis, J.M. Kretschmer, S. Manning, K.J. Chalmers, and P. Langridge. 1997. Molecular markers for boron tolerance in barley. pp. 63–65. In •• •• (ed.). Proceedings of the 8th Australian Barley Technical Symposium, Gold Coast, Australia, Vol. 3. ••, ••. Jefferies, S.P., A.R. Barr, A. Karakousis, J.M. Kretschmer, S. Manning, K.J. Chalmers, J.C. Nelson, A.K.M.R. Islam, and P. Langridge. 1999. Mapping of chromosome regions conferring boron toxicity tolerance in barley (Hordeum vulgare L.). Theor. Appl. Genet. 98:1293–1303. Johnston, R.P. 1974. Quality: the plant breeders dilemma. pp. 175–176. Proceedings of the 13th Convention of the Institute of Brewing (Australia & New Zealand Section), Surfers Paradise, Queensland, 1974. Johnston, R.P. 1983. Review of the Australian barley breeding programs—Queensland. pp. 3–5. In •• •• (ed.). Australian Barley Technical Symposium Report, CanningVale, Perth, WA, 1983. ••, ••. Johnston, R.P. 1985. Barley improvement in Queensland. pp. 27–28. Proceedings of the 2nd Australian Barley Technican Symposium, Toowoomba, Queensland, 1985. Joppich, G. 1985. Review of current barley research projects. pp. 6–11. Proceedings of the 2nd Australian Barley Technical Symposium, Toowoomba, 1985. Karakousis, A., K. Chalmers, A. Barr, and P. Langridge. 2000. Identiication of SSR markers for use in the Australian barley breeding programs. pp. 64–66. In S. Logue (ed.). Proceedings of the 8th International Barley Genetics Symposium, Adelaide, Australia, Vol. 3. ••, Adelaide, Australia. Karakousis, A., J.P. Gustafson, and K.J. Chalmers. 2001. ••. The Integration of SSR Markers into Consensus Maps of Australian Barley Mapping Populations Plant and Animal Genome IX Conference. Available at http://www. intl-pag.org/pag/9/abstracts/P3b_02.html. Karakousis, A., A.R. Barr, K.J. Chalmers, G.A. Ablett, T.A. Holton, R.J. Henry, P. Lim, and P. Langridge. 2003a. Potential of SSR markers for plant breeding and variety 205 identiication in Australian barley germplasm. Aust. J. Agric. Res. 54:1197–1210. Karakousis, A., A.R. Barr, J.M. Kretschmer, S. Manning, S.P. Jefferies, K.J. Chalmers, A.K.M. Islam, and P. Langridge. 2003b. Mapping and QTL analysis of the barley population Clipper × Sahara. Aust. J. Agric. Res. 54:1137–1140. Karakousis, A., A.R. Barr, J.M. Kretschmer, S. Manning, S.J. Logue, S. Roumeliotis, H.M. Collins, K.J. Chalmers, C.-D. Li, R.C.M. Lance, and P. Langridge. 2003c. Mapping and QTL analysis of the barley population Galleon × Haruna Nijo. Aust. J. Agric. Res. 54:1131– 1135. Karakousis, A., J.P. Gustafson, K.J. Chalmers, A.R. Barr, and P. Langridge. 2003d. A consensus map of barley integrating SSR, RFLP and AFLP markers. Aust. J. Agric. Res. 54:1173–1185. Kasha, K.J. and E. Reinbergs. 1982. Recent developments in the production and utilization of haploids in barley. pp. 655–665. In R.N.H. Whitehouse (ed.). Proceedings of the 4th International Barley Genetics Symposium, Edinburgh, 99 Scotland, 1981. ••, Edinburgh, Scotland. Kasha, K.J., U.-H. Cho, and A. Ziauddin. 1992. Application of microspore cultures. pp. 793–806. In L. Munck (ed.). Barley Genetics IV, Proceedings of the 6th International Barley Genetics Symposium, Helsingborg, Sweden, 1991. 100 Volume II: Barley Research Reviews 1981–1991. ••, ••. Kretschmer, J.M., K.J. Chalmers, S. Manning, A. Karakousis, A.R. Barr, A.K.M.R. Islam, S.J. Logue, Y.W. Choe, S.J. Barker, R.C.M. Lance, and P. Langridge. 1997. RFLP mapping of the Ha2 cereal cyst nematode resistance gene in barley. Theor. Appl. Genet. 94:1060–1064. Lance, R.C.M., J. Eglinton, J. Frankowiak, G. Platz, and M. VanGinkel. 2007. Barley breeding Australia—a new paradigm—working together for a new future. Proceedings of the 13th Australian Barley Technical Symposium, Fremantle, Western Australia, 2007. Available at http:// proceedings.com.au/abts2007/pdf/24_ABTS-07_ Lance.pdf. Langridge, P. 1997. A strategic initiative of the GRDC. pp. 2:10.1–2:10.4. Proceedings of the 8th Australian Barley Technical Symposium, Gold Coast, Queensland, September 7–12, 1997. Langridge, P. and A.R. Barr. 2003. Preface to “Better Barley Faster: the Role of Marker Assisted Selection,” coordinated by T.J. Higgins, G.R. Steed, J.C. Fegent, S. Banerjee. Aust. J. Agric. Res. 54:i–iv. Langridge, P., A. Karakousis, N. Collins, J. Kretschmer, and S. Manning. 1995. A consensus linkage map of barley. Mol. Breed. 1:389–395. Li, C.-D., R.C.M. Lance, H.M. Collins, A. Tarr, S. Roumeliotis, S. Harasymow, M. Cakir, G.P. Fox, C.R. Grime, S. Broughton, K.J. Young, H. Raman, A.R. Barr, D.B. Moody, and B.J. Read. 2003a. Quantitative trait loci controlling kernel discoloration in barley (Hordeum vulgare L.). Aust. J. Agric. Res. 54:1251–1259. Li, C.-D., A. Tarr, R.C.M. Lance, S. Harasymow, J. Uhlmann, S. Westcot, K.J. Young, C.R. Grime, M. Cakir, S. Broughton, and R. Appels. 2003b. A major QTL controlling seed dormancy and pre-harvest sprouting/grain Ullrich—Production, Improvement, and Uses c08.indd 205 8/30/2010 8:28:57 PM F Barley: Production, Improvement, and Uses 206 101 F α-amylase in two-rowed barley (Hordeum vulgare L.). Aust. J. Agric. Res. 54:1303–1313. Logue, S.J., L.C. Giles, R.C.M. Lance, and D.H.B. Sparrow. 1993. The application of anther culture technology to barley improvement. pp. 61–64. Proc. 6th Austr. Barley Tech. Symp., Launceston, Tasmania, 1993. Long, N.R., S.P. Jefferies, P. Warner, A. Karakousis, J.M. Kretschmer, C. Hunt, P. Lim, P.J. Eckermann, and A.R. Barr. 2003. Mapping and QTL analysis of the barley population Mundah × Keel. Aust. J. Agric. Res. 54: 1163–1171. Loughman, R. 1998. Western Region report. Australian Barley Diseases Newsletter, Issue No. 2, ISSN 1440 6519, p. 1. Lovett, J. 1997. Balancing the investment. pp. 2:1.3–2:1.9. Proceedings of the 8th Australian Barley Technical Symposium, Gold Coast, Queensland, Australia, 1997. Lovett, J.V. 2001. Barley research and development—21st century opportunities. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia. Available at http://www.regional.org.au/au/abts/2001/ m1/lovett.htm#TopOfPage. MacLeod, L. 2001. Servicing the Asian brewing market—an exporters perspective. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia. Available at http://www.regional.org.au/au/abts/2001/ w3/macleod.htm#TopOfPage. Malting and Brewing Industry Barley Technical Committee (MBIBTC). 1998. Industry Guidelines for Australian Malting Barley, Vol. 4. The Malting and Brewing Industry Barley Technical Committee, ••••. McIntosh, R.A., C.R. Wellings, and R.F. Park. 1995. Wheat Rusts: an Atlas of Resistance Genes. CSIRO Publications, East Melbourne, Victoria, Australia. Murray, G.M. and J.P. Brennan. 2001. Prioritising threats to the Australian grains industry. Technical Report on GRDC Project: DAN438. NSW Agriculture and the Grains Research and Development Corporation, 2001. Pallotta, M.A., R.D. Graham, P. Langridge, D.H.B. Sparrow, and S.J. Barker. 2000. RFLP mapping of manganese eficiency in barley. Theor. Appl. Genet. 101:1100–1108. Pallotta, M.A., S. Asayama, J.M. Reinheimer, P.A. Davies, A.R. Barr, S.P. Jefferies, K.J. Chalmers, J. Lewis, H.M. Collins, S. Roumeliotis, S.J. Logue, S.J. Coventry, R.C.M. Lance, A. Karakousis, P. Lim, A.P. Verbyla, and P.J. Eckermann. 2003. Mapping and QTL analysis of the barley population Amagi Nijo × WI2585. Aust. J. Agric. Res. 54:1141–1144. Paltridge, N.G., N.C. Collins, A. Bendahmane, and R.H. Symons. 1998. Development of YLM, a codominant PCR marker closely linked to the Yd2 gene for resistance to barley yellow dwarf virus. Theor. Appl. Genet. 96: 1170–1177. Paris, M., R.H. Potter, R.C.M. Lance, C.-D. Li, and M.G.K. Jones. 2003. Typing mlo alleles for powdery mildew resistance in barley by single nucleotide polymorphism analysis using MALDI-ToF mass spectrometry. Aust. J. Agric. Res. 54:1343–1349. Park, R. and K. Williams. 2000. 1999–2000 Cereal Rust Survey Annual Report. Cereal Rust Laboratory, Plant Breeding Institute, The University of Sydney, Cobbity, New South Wales, Australia. Park, R.F., D. Poulsen, A.R. Barr, M. Cakir, D.B. Moody, H. Raman, and B.J. Read. 2003. Mapping genes for resistance to Puccinia hordei in barley. Aust. J. Agric. Res. 54:1323–1333. Parlevliet, J.E. and A. van Ommeren. 1988. Accumulation of partial resistance in barley to barley leaf rust and powdery mildew through recurrant selection against susceptibility. Euphytica 37:261–274. Paynter, B.H. and N.A. Fettell. 2008. Cultural practices: focus on major barley producing regions—Australia. In S.E. Ullrich (ed.). Barley Subtitle: Production, Improvement, and Use. Washington State University, Pullman, WA. Pickering, R.A., A.M. Hill, M. Michel, and G.M. Timmerman-Vaughan. 1995. The transfer of a powdery mildew resistance gene from Hordeum bulbosum L. to barley (H. vulgare L.) chromosome 2 (2I). Theor. Appl. Genet. 91:1288–1292. Pickering, R.A., B.J. Steffenson, A.M. Hill, and I. Borovkova. 1998. Association of leaf rust and powdery mildew resistance in a recombinant derived from a Hordeum vulgare × Hordeum bulbosum hybrid. Plant Breed. 117: 83–84. Pickering, R.A., S. Malyshev, G. Kunzel, P.A. Johnston, V. Korzun, M. Menke, and I. Schubert. 2000. Locating introgressions of Hordeum bulbosum chromatin within the H. vulgare genome. Theor. Appl. Genet. 100:27–31. Platz, G. 2001. The onset and effectivness of adult plant resistance (APR) in Tallon barley. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, ACT, 2001. Available at http://www.regional.org.au/au/abts/ 2001/t1/platz.htm#TopOfPage. Portman, P. 1983. Review of the Australian barley breeding programs—Western Australia. pp. 15–16. Australian Barley Technical Symposium Report, Perth, Western Australia, 1983. Portman, P. 1985. Barley in Western Australia. pp. 12–13. Proceedings of the 2nd Australian Barley Technical Symposium, Toowoomba, 1985. Poulsen, D.M.E. 2001. A tremendous innings: Paul Johnston’s legacy to the barley industry. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, ACT, 2001. Available at http://www.regional.org.au/au/ abts/2001/w2/poulsen.htm#TopOfPage. Poulsen, D.M.E., R.J. Henry, R.P. Johnston, J.A.G. Irwin, and R.G. Rees. 1995a. The use of bulk segregant analysis to identify an RAPD marker linked to leaf rust resistance in barley. Theor. Appl. Genet. 91:270–273. Poulsen, D.M.E., P.A. Inkerman, and R.P. Johnston. 1995b. Breeding better barley varieties for northern Australia. pp. 246–250. In Y.A. Williams and C.W. Wrigley (eds.). Proceedings of the 45th Australian Cereal Chemistry 102 Conference, Adelaide, South Australia, 1995. ••, ••. Poulsen, D.M.E., D. Butler, J.M. Sturgess, R.L. Fromm, M.J. Laufer, and R. Mole. 1997. Improving grain size distribution in segregating breeding material. pp. 3:78– 3:79. Proceedings of the 8th Australian Barley Technical 103 Symposium, Gold Coast, Australia. Ullrich—Production, Improvement, and Uses c08.indd 206 8/30/2010 8:28:57 PM Barley Breeding History, Progress, Objectives, and Technology Poulsen, D.M.E., R.P. Johnston, G.J. Platz, G. Fox, A. Kelly, J.M. Sturgess, R.L. Fromm, M.J. Laufer, P.A. Inkerman, and D. Butler. 1999. Effects of foliar diseases on Northern Region grain production in the 1998 winter cropping season. pp. 2.20.1–2.20.5. Proceedings of the 9th Australian Barley Technical Symposium, Melbourne, Victoria, Australia, 1999. Poulsen, D.M.E., G.J. Platz, B.J. Steffenson, R. Dill-Macky, J.A.G. Irwin, R.J. Henry, and R.P. Johnston. 2000. Breeding disease resistant parent lines using yield trial evaluation, in-vivo disease screening and marker assisted selection. pp. 33–35. In S. Logue (ed.). Barley Genetics VIII: Proceedings of the 8th International Barley Genetics Symposium, Adelaide, South Australia, October 22–27, 104 2000, Vol. 3. ••, Adelaide, South Australia. Powell, G. 1997. Quality requirements for Australian malting barley. pp. 2:2.2(a)–2:2.2(h). Proceedings of the 8th Australian Barley Technical Symposium, Gold Coast, Queensland, Australia, 1997. Pugsley, A.T. 1951. Genetics and plant breeding. p. 14. Report of the Waite Agricultural Research Institute 1950–51. Pugsley, A.T. and A. Vines. 1946. Breeding Australian barleys resistant to covered smut. J. Austr. Inst. Agric. Sci. 12:44. Purse, G.S. and D. McNee. 1985. National barley research. pp. 2–5. Proceedings of the 2nd Australian Barley Technical Symposium, Toowoomba, 1985. Qi, X., P. Stam, and P. Lindhout. 1996. Comparison and integration of four barley genetic maps. Genome 39: 379–394. Ramage, R.T. 1975. Techniques for producing hybrid barley. Barley Newsl. 18:62–65. Ramage, R.T. 1987. A history of barley breeding methods. Plant Breed. Rev. 5:95–138. Raman, H., S. Moroni, R. Raman, A. Karakousis, B. Read, K. Sato, and B.J. Scott. 2001. A genomic region associated with aluminium tolerance in barley. Proceedings of the 10th Australian Barley Technical Symposium, Canberra, Australia. Available at http://www.regional.org.au/au/ abts/2001/t3/raman.htm#TopOfPage. Raman, H., J.S. Moroni, K. Sato, B.J. Read, and B.J. Scott. 2002. Identiication of AFLP and microsatellite markers linked with an aluminium tolerance gene in barley (Hordeum vulgare L.). Theor. Appl. Genet. 105:458–464. Raman, H., G.J. Platz, K.J. Chalmers, R. Raman, B.J. Read, 105 A.R. Barr, and D.B. Moody. 2003. Mapping of genomic regions associated with net form of net blotch resistance in barley. Aust. J. Agric. Res. 54:1359–1367. Read, B. 1983. Review of the Australian barley breeding programs—New South Wales. pp. 2–3. Australian Barley Technical Symposium Report, Perth, Western Australia, 1983. Read, B.J. 1985. Review of barley breeding in New South Wales. pp. 16–17. Proceedings of the 2nd Australian Barley Technical Symposium, Toowoomba, Queensland, 1985. Read, B.J., H. Raman, G. McMichael, K.J. Chalmers, G.A. Ablett, G.J. Platz, R. Raman, R.K. Genger, W.J.R. Boyd, C.-D. Li, C.R. Grime, R.F. Park, H. Wallwork, R. 207 Prangnell, and R.C.M. Lance. 2003. Mapping and QTL analysis of the barley population Sloop x Halcyon. Aust. J. Agric. Res. 54:1145–1153. Reading, P. 2007. Varieties for a vibrant future: maximising impact of investment in barley breeing. Proceedings of the 13th Australian Barley Technical Symposium, Fremantle, Western Australia, 2007. Available at http://proceedings. com.au/abts2007/pdf/24_ABTS-07_Reading.pdf. Rees, R.G. 1985. Northern Australian disease situation. pp. 96–98. Proceedings of the 2nd Australian Barley Technical Symposium, Toowoomba, Queensland, 1985. Rees, R.G., R.P. Johnston, and T.J. Moore. 1981. Foliar and head diseases of barley in Queensland. pp. 58–61. Barley diseases in Australasia: Proceedings of a Workshop held at Adelaide, October 17–19, 1979. Rees, R.G., W.M. Strong, and T.J. Neale. 1999. The effects of foliar diseases on the production of wheat and barley in the Northern Region in 1998. Report to the Northern Panel of the Grains Research and Development Corporation. Riggs, T.J., P.R. Hanson, and N.D. Start. 1982. Modiication of the pedigree method in spring barley breeding by incorporating bulk selection in F4. pp. 138–141. In R.N.H. Whitehouse (ed.). Proceedings of the 4th International Barley Genetics Symposium, Edinburgh, Scotland, 1981. ••, ••. Sewell, R. 1997. Barley Quality Objectives Group Report to “The Australian Barley Technical Symposium.” pp. 2:3.1–2:3.3. Proceedings of the 8th Australian Barley Technical Symposium, Gold Coast, Queensland, Australia, 1997. Sharp, E.L. 1985. Breeding for pest resistance. pp. 313–333. In D.C. Rasmusson (ed.). Barley. American Society of Agronomy, Series: Agronomy Monograph No. 26. American Society of Agronomy, ••. Smith, A.N. 1987. Barley Research Council: priorities for research and development. pp. 1–6. Proceedings of the 3rd Australian Barley Technical Symposium, Wagga Wagga, NSW, 1987. Sparrow, D.H.B. 1983. Review of the Australian barley breeding programs—South Australia. pp. 5–10. Australian Barley Technical Symposium Report, Perth, Western Australia, 1983. Sparrow, D.H.B. 1984. Recent and projected advances in the improvement of malting barleys for Australian conditions. pp. 59–66. Proceedings of the 18th Convention of the Institute of Brewing (Australia and New Zealand Section), Adelaide, South Australia, 1984. Sparrow, D.H.B. and J.B. Doolette. 1975. Barley. pp. 431– 480. In A. Lazenby and E.M. Matheson (eds.). Australian Field Crops: Wheat and Other Temperate Cereals, Vol. 1, 2nd ed. Angus and Robertson Publishers, Australia. Spiel, G. 1999. Current and future trends in barley quality requirements. pp. 2.13.1–2.13.6. Proceedings of the 9th Australian Barley Technical Symposium, Melbourne, Victoria, Australia, September 12–16, 1999. Stuart, J. 1997. Advancements in malting barley marketing. pp. 2:12.1–2:12.6. Proceedings of the 8th Australian Barley Technical Symposium, Gold Coast, Queensland, Australia, 1997. 106 107 Ullrich—Production, Improvement, and Uses c08.indd 207 8/30/2010 8:28:57 PM F Barley: Production, Improvement, and Uses 208 108 Stuthman, D.D., G. Sosa-Dominguez, and D.L. De Koeyer. 1996. Recurrent selection in autogamous crops—Past, current and future. pp. 240–246. In G. Scoles and B. Rossnagel (eds.). Proceedings of the 5th International Oat Conference and 7th International Barley Genetics Symposium, Saskatoon, Saskatchewan, Canada. Invited Papers, Vol. 1. ••, Saskatoon, Saskatchewan, Canada. Tang, Y., M.E. Sorrells, L.V. Kochian, and D.F. Garvin. 2000. Identiication of RFLP markers linked to the barley aluminium tolerance gene Alp. Crop Sci. 40:778–782. Verbyla, A.P., P.J. Eckermann, R. Thompson, and B.R. Cullis. 2003. The analysis of quantitative trait loci in multienvironment trials using a multiplicative mixed model. Aust. J. Agric. Res. 54:1395–1408. Vertigan, W. 1983. Review of the Australian barley breeding programs—Tasmania. pp. 10–11. Australian Barley Technical Symposium Report, Perth, Western Australia, 1983. Wallwork, H., P. Preece, and P.J. Cotterill. 1992. Puccinia hordei on barley and Ornithogalum umbellatum in South Australia. Australas. Plant Pathol. 21:95–97. Williams, K.J. 2003. The molecular genetics of disease resistance in barley. Aust. J. Agric. Res. 54:1065–1079. Williams, K., P. Bogacki, L. Scott, A. Karakousis, and H. Wallwork. 2001. Mapping of a gene for scald resistance in barley line “B87/14” and validation of microsatellite and RFLP markers for marker-assisted selection. Plant Breed. 120:301–304. Williams, K.J., A. Lichon, P. Gianquitto, J.M. Kretschmer, A. Karakousis, S. Manning, P. Langridge, and H. Wallwork. 1999. Identiication and mapping of a gene conferring resistance to the spot form of net blotch (Pyrenophora teres f. maculata) in barley. Theor. Appl. Genet. 99:323–327. Williams, K.J., G.J. Platz, A.R. Barr, J. Cheong, K. Willsmore, M. Cakir, and H. Wallwork. 2003. A comparison of the genetics of seedling and adult plant resistance to the spot form of net blotch (Pyrenophora teres f. maculata). Aust. J. Agric. Res. 54:1387–1394. Xu, J. and K.J. Kasha. 1992. Transfer of a dominant gene for powdery mildew resistance and DNA from Hordeum bulbosum into cultivated barley (H. vulgare). Theor. Appl. Genet. 84:771–777. NEAR EAST, NORTH AND EAST AFRICA, AND LATIN AMERICA Salvatore Ceccarelli, Stefania Grando, and Flavio Capettini nEAR EAsT Barley is grown on about 8 million hectares in 16 different countries in the Near East. However, 98% of the area is in six countries, namely, Turkey, Iran, Syria, Iraq, Afghanistan, and F Pakistan (http://faostat.fao.org/, searched in 109 2008). There is a large variability in the types of barley grown in this area. Two-row types predominate in Syria, Turkey, the northern part of Iraq, and the rainfed areas of Iran, while six-row types predominate in Pakistan and Afghanistan and in the irrigated areas of Iraq and Iran. In most of the dry areas of Syria and Iraq, and in some of the rainfed areas of Northern Iran, farmers have a strong preference for black-seeded types. Most of the barley grown in the areas is either winter or facultative and with a strong photoperiod sensitivity. With the exception of the irrigated areas, barley is predominantly grown in the semiarid and arid areas where rainfall is too low and erratic to grow wheat. In the Near East, the predominant use of barley is as animal feed. However, it is also used as food in Iraq and Iran. syria Formal agricultural research in Syria began in the early 1940s with the establishment of experiment farms at Deir Elhajar and Kharabo, close to Damascus. After independence, in 1946, various Ministry of Agriculture and Agrarian Reform (MAAR) directorates (such as horticulture, forestry, animal resources, and plant protection) began to conduct limited agricultural research. In 1964, the Directorate of Agricultural Scientiic Research (DASR) was established, taking on 110 responsibility for research activities under MAAR. In 1982, DASR formed a technical committee to develop new varieties of cereals, legumes, and forage crops and to coordinate research activities with the Department of Agricultural and Scientiic Research (DASR), the International Center for Agricultural Research in the Dry Areas (ICARDA), and the Arab Center for Studies of the Arid Zones and Dry Lands (ACSAD). This could be considered as the beginning of the barley breeding program, with training programs conducted on crossing techniques and breeding methods. In 2002, the nine existing agricultural research entities under MAAR were merged to form the General Commission for Scientiic and Agricultural Research (GCSAR), which is now Ullrich—Production, Improvement, and Uses c08.indd 208 8/30/2010 8:28:57 PM Barley Breeding History, Progress, Objectives, and Technology responsible for all the breeding programs (wheat, barley, chickpea, lentil, faba bean, cotton, etc.). The breeding methods used vary from the introduction of international nurseries from international centers such as ICARDA and from other countries and their direct release as cultivars (examples in barley are “Furat 1” and “Furat 2”released directly from ICARDA’s international nurseries); the introduction of segregating populations from ICARDA; the execution of crosses followed by classical pedigree and the induction of mutations (wheat, barley, chickpea, and soya beans). Plant breeding programs give special importance to traits such as drought resistance, earliness, disease resistance, and high yield. The barley breeding work is conducted at the stations and centers of GCSAR, and neither foreign companies nor the private sector conducts plant breeding activities in Syria. The inal stage of the breeding program is the on-farm trials, which are used to eventually recommend varieties for release. As a result of breeding work since the early 1970s, seven barley varieties have been released, but their rate of adoption is very low partly because of their lack of adaptation to the farmers’ real conditions and partly because of the limited capacity of the General Organization of Seed Multiplication (GOSM), which is only able to produce 10% of the barley seed required for planting, while the corresponding igure in the case of wheat is 60% (Al-Ahmad et al. 1999). Jordan Formal agricultural research in Jordan began in the early 1950s with the creation of the irst agricultural research station in the Jordan Valley. During that decade, various other research stations were established throughout the country. Research was initially carried out by the technical divisions of the Ministry of Agriculture, which were transferred to the Department of Scientiic 111 Agricultural Research when it was created in 1958, and then in 1970, the department was merged with the Ministry’s extension unit to form the Department of Scientiic Research and Agricultural Extension. The Ministry of 209 Agriculture was restructured in the mid-1980s, resulting in the creation of the National Center for Agricultural Research and Technology Transfer (NCARTT). In 2007, NCARTT was renamed as the National Center for Agricultural Research and Extension (NCARE). The Center developed over the years, in part through the National Agricultural Development Project (NADP), which was cofunded by the United States Agency for International Development (USAID) and the Government of Jordan. Agricultural research within the higher education sector began in the early 1970s with the establishment of the Faculty of Agriculture (1972) and the Marine Science Station (1974) at the University of Jordan (UoJ). Even if research in barley breeding is occasionally conducted at the university, NCARE has the main responsibility for breeding and releasing varieties (Taimeh and Sunna 1999). The barley breeding program in Jordan is entirely based on introduced germplasm mostly from ICARDA and ACSAD. The material is tested in two research stations for 4 years before being tested in on-farm trials and eventually recommended for release. In addition to the local landrace (two-row), the most popular improved variety has been Rum (six-row). Recently, the program has released three additional varieties, namely, Athroh (six-row), Yarmouk (Esp/18084L//Harmal, two-row), and Muta’a (Roho/A. Abiad/6250, two-row). During the last 4 years, the barley breeding program in Jordan has gradually been converted into a participatory program following the methodology described in Ceccarelli and Grando (2007). iran Agricultural research in Iran dates back to 1925 when the Razi Institute, the “father” of the current Razi Serum and Vaccine Research and Production Institute (RSVRI), began its research activity in the district of Karaj, about 35 km west of Tehran. In 1926, the irst agricultural college, afiliated with the Ministry of Agriculture, was founded in Karaj. Important changes occurred in Ullrich—Production, Improvement, and Uses c08.indd 209 8/30/2010 8:28:57 PM F Barley: Production, Improvement, and Uses 210 the 1950s and 1960s. New colleges of agriculture were established and institutes were created with technical support from international agencies, such as the United Nation’s Food and Agriculture Organization (FAO). Among these, the Seed and Plant Improvement Institute (SPII) was established in Karaj in 1959 to conduct research and seed multiplication on the main crops (cereals, oil crops, cotton, rice, horticultural crops, forages, etc.). In 1991, the Dryland Agricultural Research Institute (DARI) was established in Maragheh (Azerbaijan province) to relieve SPII from its heavy responsibilities and to implement extensive research on some of the major commodities and noncommodity items of economic importance. In particular, DARI conducts basic research on mechanisms of abiotic (cold and heat) tolerance from the physiological, genetic, and agronomic points of view, and on developing strategies to overcome these stresses and to increase productivity. Barley breeding started in Iran around 1930 with the exploitation of local germplasm. This was followed in the 1950s by the beginning of hybridization of local germplasm with exotic germplasm and the extensive use of the pedigree method, bulking in F6 and multilocation testing in about 22 stations throughout the country. The inal testing is done in the Uniform Regional Yield Trials, which are speciically targeted to the warm, cold, and rainfed regions of the country. 112 After the establishment of DARI, SPII kept the responsibility for barley breeding in the irrigated areas of the country, while DARI focused on the rainfed areas (Roozitalab 1996). Both breeding programs (SPII and DARI) collaborate actively with ICARDA: they receive, test, and use both international nurseries as well as special nurseries speciically targeted to the growing environments in Iran. About 80% of the barley germplasm is either a direct introduction from ICARDA or is derived from crosses with the ICARDA germplasm. iraq In the 1920s, a Directorate General of Agriculture, afiliated with the Ministry of Economics and F Transport, started agricultural research activities and established the irst experimental stations at Abu Ghraib, near Baghdad and at Neinevah, near Mosul. In the 1940s, agricultural research activities were conducted by the Directorate General for Agricultural Research and Extension (DGARE) until in 1958 the Directorate General for Agricultural Research and Projects (DGAREJ) was established. During the 1970s, research activities were expanded as several specialized research stations and centers were established including the Iraqi Atomic Energy Commission (IAEC). In 1980, the State Board for Applied Agricultural Research (SBAAR) was established, which in 1987 was renamed as the State Board for Agricultural Research and Water Resources (SBARWS). In 1990, SBARWS was terminated and replaced by the State Board for Agricultural 113 Research (SBAR) and CWSR. Barley breeding in Iraq used to be divided into two subprograms: one addressing the irrigated areas around Baghdad and based in Abu Ghraib, and the second addressing the rainfed areas of the North, which, from an agroecological viewpoint, are very similar to the areas in Northeast Syria. The irst subprogram, based on six-row types mostly bred for dual purpose as cattle feed, fully exploited the segregating populations received from ICARDA and released successful cultivars such as IPA 7, IPA 9, and IPA 265. The second subprogram, although addressing an area where farmers’ preferences are for two-row black-seeded barley, attempted the introduction of six-row types. The major success of the program was the release of Rihane-03, a straight introduction from ICARDA, which in the late 1990s was cultivated on about 250,000 ha. Currently, due to the war since 2003, Iraq is conducting a limited amount of variety trials using material introduced from ICARDA. Yemen Agricultural research in Yemen dates back to the 1940s during the rule of the British Colonial Government. El-Kod Research Station was established in 1955 about 50 km from Aden, and Ullrich—Production, Improvement, and Uses c08.indd 210 8/30/2010 8:28:57 PM Barley Breeding History, Progress, Objectives, and Technology Seiyun Research Center was established in Wadi Hadramout in 1972 to cover the mid-altitude region of South Yemen. Research activities were later developed in most parts of the country, especially through numerous projects supported by 114 UNDP/FAO and IDA. They were introduced in North Yemen starting in 1970, later developing into a central research station in Taiz in 1978. In 1980, the Ministry of Agriculture and Agrarian Reform of South Yemen created the Department of Research and Extension (DRE), based at Aden, which was transformed in 1986 to the Directorate of Research and Extension (Ministry of Agriculture and Fisheries 1989). At the same time, the Agricultural Research Authority (ARA) was established in Dhamar in North Yemen for conducting research and applied studies to improve agricultural production. In 1990, after the uniication of North and South Yemen, the Agricultural Research and Extension Authority (AREA) was formed by merging DRE and ARA and their respective research centers and stations. As in the case of other countries in the region, Yemen’s barley breeding program is based on introductions from international centers, particularly ICARDA and ACSAD, and more recently from the International Atomic Energy Agency (IAEA). Most of the breeding work takes place at the research station of Al Erra, just outside the capital city Sana’a. Two varieties, introduced through ICARDA nurseries, Beecher and Arivat, were released in 1986. In 1999, ICARDA and AREA conducted a 2-year project introducing participatory plant breeding in Yemen: two barley varieties were adopted by farmers at the end of the project, and currently, AREA is using participatory methodology to evaluate barley germplasm obtained by mutation breeding from IAEA. nORTH AfRiCA Cereal cultivation in North Africa dates back to ancient times. It seems that it preceded the Phoenician colonization, which took place about the twelfth century BC. The founding of Utica at the delta of the Médjerda valley, particularly 211 suited to wheat cultivation, had the only objective of supplying food for the long sea route from Tyrus to Guadalquivir. Carthage eventually replaced Utica as the export harbor near the present site of Tunis. When the Roman Empire, after destroying Carthage, spread over North Africa, cereal cultivation was already established in the plateau of Setif (Algeria) and in Numidia, which was later occupied by the Romans because of the soil fertility and the abundance of cereals they could provide. “The soil of Africa,” wrote Pliny, the Elder in the irst century of our era, “has been given as gift from Ceres: oil and wine were almost refused; all the glory of the country is in the harvest.” Today, barley is second only to wheat in the ive North African countries where it is grown on about 3.5 million hectares (average of 2002–2006) of which about 60% is in Morocco (http:// faostat.fao.org/). It is predominantly grown in the semiarid and arid areas where rainfall is too low and erratic to grow wheat. These include the Northwest coast of Egypt, where high humidity allows growing a barley crop with about 100 mm annual rainfall, and in some provinces of Western Algeria. In the latter, barley is an established crop because of high demand for livestock feed. Contrary to the Near East, the majority of the barley landraces and improved varieties grown in North Africa are six-row with very few exceptions. The release of superior two-row cultivars has not been followed by any signiicant adoption because of religious reasons (two-row types are associated with beer production). As in the Near East, the main use of barley is as animal feed. However, the use of barley as human food is much more widespread than in the Near East, and in fact, Morocco has the world’s largest per capita human consumption of barley. In recent times, the consumption of barley as human food has increased also in Algeria and Egypt (Grando and Gomez Macpherson 2005). 115 The history of barley breeding in North Africa varies among the countries, but, as already described in the case of the Near East, in all countries is much more recent than for wheat. As a consequence, and for many years, barley and Ullrich—Production, Improvement, and Uses c08.indd 211 8/30/2010 8:28:58 PM F Barley: Production, Improvement, and Uses 212 wheat breeding were done by the same scientist, in the same research stations, and with the same philosophies and methodologies. This situation did not consider that the two crops are grown in distinct agroecologies by farms differing in size, by farmers differing in wealth with access to differing amounts of information. Tunisia F The history of barley breeding in Tunisia is emblematic of most of North Africa and the Near East. In Tunisia, cereal breeding started with wheat at the beginning of the last century in the then Service Botanique et Agronomique de Tunisie (SBAT). Wheat breeding evolved from the utilization of the variability existing within local varieties of durum wheat, followed by mass selection until 1930, when the irst crosses were made. It was about at this time that bread wheat was introduced from France and Algeria. Later in the 1970s, the introduction of germplasm from Centro Internacional de Mejoramiento de Maíz y Trigo (CIMMYT) occurred, when several varieties were released, both using direct introduction and selection from segregating populations. The introduction of these varieties was accompanied by the wider use of inputs such as fertilizers and chemical weed control. The barley breeding programs are much more recent as the local varieties were used until 1950 when “Martin” and “Cérès” were introduced in the North of the country. Problems with seed production of these two varieties did not allow replacing the local germplasm in the center and the south of the country. The irst serious attempt to start a barley breeding program was made in 1973 (•• Harrabi, 116 pers. comm.) with crosses and selection of early material for the semiarid environments. The materials obtained were tested in central Tunisia and in the high plateau of the Northwest. With the establishment of ICARDA in 1976, Tunisia, where ICARDA had a regional ofice, was used as a testing site for ICARDA’s barley breeding program, thus receiving more genetic material than other countries. A large number of lines were screened, and already in 1984, the irst promising lines, such as ER/Apm, Roho, and WI 2198 (from the Waite Institute in Adelaide, South Australia) were identiied and eventually released in 1985 with the names of Faiz, Roho, and Taj, respectively. The collaboration with ICARDA has continued over the years with the exchange of germplasm (both inished lines and segregating populations) and the release of varieties based on those nurseries. Algeria Wheat (bread and durum) and barley are the main winter cereals in Algeria. Among the wheats, it is possible to ind small amounts of Triticum turgidum, Triticum dicoccoides, and Triticum polonicum in the local landraces of wheat and Triticum spelta particularly in the oases. As in Tunisia, farmers have initially grown local landraces. Cereal breeding started in 1905 by Prof. L. Ducellier, who produced the irst pure lines of durum wheat. After his premature death, his work was continued by his students at the Station Centrale d’Essais de Semences et d’Amélioration des Plantes de Grande Culture de Maison-Carrée and by other botanists as Trabut, Bœuf, and Laumont. The history of breeding can be divided into 117 three periods: 1. 2. The irst period (1900–1962) was characterized by the utilization of the existing material within local landraces with collections and characterization of durum wheat. Laumont and Ducellier started from 1930 the production of varieties selected by mass selection procedure and then the irst crosses were made; these crosses aimed at higher productivity and better quality. The second period (1962–1990) was characterized by the introduction of germplasm from Spain, Italy, Greece, Syria, Russia, and the United States through European projects, FAO, CIMMYT, and then ICARDA. Several varieties were released either by direct introductions or selection from segregating populations. These varieties were characterized by the need for more inputs Ullrich—Production, Improvement, and Uses c08.indd 212 8/30/2010 8:28:58 PM Barley Breeding History, Progress, Objectives, and Technology (fertilizers and weed control) than landraces. During this second stage, a limited crossing program also started. This irst stage was followed by the introduction of durum wheats and by hybridization between the local landraces and the introduced varieties. Already at that time, the Station Centrale de Maison-Carrée could count on a network of a dozen regional experiment stations corresponding to the main cereal regions of Algeria. The improvement of bread wheat, which started being grown in Algeria only in 1930, followed the same route as durum wheat. Similarly, in the case of barley, the initial cultivation of mixtures introduced from the Near East was followed by pedigree selection, which led to the development of locally adapted cultivars such as Saida 183 and Tichedrette, the latter particularly adapted to the mountainous areas. The majority of cultivated barley has white kernels; however, farmers remember cultivating black types considered resistant to drought. Malting barley did not become important in Algeria because of its susceptibility to shattering and because of the indifference of the local industry. As a result of the collaboration with ICARDA, several varieties were released, but the two locally adapted types are still the most widely grown. At the moment, Algeria’s barley breeding program is almost entirely based on ICARDA’s germplasm and in the last 3 years has started being organized in a participatory way (Ceccarelli and Grando 2007). Morocco The beginning of barley breeding in Morocco dates back to 1920 and was based on the improvement of local landraces and on the introduction of two-row foreign varieties. Among hundreds of cultivars from Europe, the United States, and Australia, about a dozen were selected including Chevalier, Hannchen, Combesse, Guldkorn, Princesse, and Prior (Grillot 1939). Selection within the local populations initially identiied two lines, 077 and 071, later named 213 Rabat 77 and Merzaga 71, respectively (Saidi et al. 2005). The introduction of two- and six-row varieties characterized barley breeding until 1970 and led to the release of cultivars such as Arig 8, Tamelalt, Asni, and Azilal. Starting from 1980, the introduction of collections of barley from the United States was the beginning of another phase of barley breeding with the objective of improving earliness and harvest index. The germplasm generated in this period was used for multilocation testing to identify widely adapted lines (Amri 1993). Selection was conducted in the research stations for disease resistance and yield. A dozen varieties were released, but despite their performance, they were grown on no more than 5% of the barley growing area. The poor adaptation to farmers’ conditions was considered to be the cause of the lack of adoption (Saade 1994). This imposed a change of strategy, which consists of breeding for wide adaptation in the case of favorable environments and for speciic adaptation in the case of marginal environments. In the latter, the barley breeding program in Morocco is now also using participatory plant breeding. Libya Agricultural research in Libya dates back to the early part of the twentieth century during the Italian colonial era, when the “Centro Sperimentale Agrario e Zootecnico della Libia” at Sidi El Masri near Tripoli was established to serve the Italian agricultural settlers. In the early 1950s, the Ministry of Agriculture was established and agricultural research became afiliated with the Directorate of Plant and Animal Production, with major changes in goals and organization. In 1981, the General People’s Committee (Minister’s Cabinet) initiated the National Authority for Scientiic Research (NASR) to formulate and supervise the national research policy, to ill in gaps in research not tackled by any existing research institutes and centers, and technically to coordinate research carried out at research centers. New research centers were established under the umbrella of NASR, including the Agricultural Research Ullrich—Production, Improvement, and Uses c08.indd 213 8/30/2010 8:28:58 PM F Barley: Production, Improvement, and Uses 214 Center (ARC), where most of the plant breeding is conducted. The Center has the following four 118 “Regional AR Centers” (RARC), supported by 13 experimental stations (Ben-Mahmoud 1966): 1. 2. 3. 4. The RARC for the western area (which encompasses the largest part of agriculture and of the population) located at Tajura, 20 km east of Tripoli, has six research stations and three experimental sites and focuses mainly on fully irrigated and supplementaryirrigated crops such as vegetables, cereals, fruits, and forages, with some emphasis on rainfed agriculture as well. The RARC for the eastern area, located at Al-Marj, 1100 km east of Tripoli, covers the areas from Sirte to the Egyptian borders (four stations)and focuses on rainfed agriculture (cereals, legumes, and fruits), forestry, and range. The RARC for the central area, located in Musrata, 200 km east of Tripoli, covers the areas from Khomes to Sirte (two stations). Its 10 researchers are working mainly on salinity research. The RARC located at Sebha (700 km south of Tripoli) covers the southern area of the country (one station) and works in the areas of hot/dry climate and fully irrigated crops (cereals, legumes, forages, and palm trees). The breeding programs are mostly based on testing germplasm introduced from international organizations: in the case of barley, the main supplier of germplasm has been traditionally ICARDA, which, as described below, in the last 10 years has begun distributing very early germplasm particularly suitable to the short growing season of most areas where barley is grown in Libya. Eight varieties have been released between 1992 and 2005. Egypt The irst school of agriculture in Egypt was established in 1869 and the irst directorate of agriculture in 1875.During the nineteenth century, agricultural research was carried out by the F Egyptian Royal Society, and as early as 1897, a number of experimental farms were established at various locations. In 1910, the Agricultural Authority was established with the responsibility for conducting research and producing seed, extending methods in crop production to farmers, especially soil analysis and use of fertilizers, pest control, and production of scientiic and technical publications. The Ministry of Agriculture was established in 1913. In 1957, research departments were formed, and one of the irst was the Department of Plant Breeding (Rivera and Elkalla 1997). The Ministry of Agriculture has undergone several reforms in the past decades. It has grown from only 7 departments in 1913 to 28 in 1950, to 194 in 1963, and to 92 these days dealing with various aspects of agricultural production. Early 119 on among the major departments were Agriculture, Horticulture, Plant Protection, Soil, Animal Production, Veterinary Laboratories, and Seed Production. These research departments were reorganized in 1971 into one research body within the Ministry of Agriculture and Land Reclamation named the General Authority for Agricultural Research, which was later (1983), renamed the ARC, and evolved as the major institution for agricultural research and extension in Egypt until today and has the major responsibility for crop breeding. Barley breeding in Egypt has a very long history, which started more than a century ago. This has attracted the attention of international organizations for the Egyptian genetic resources, especially in relation to adaptation to abiotic stresses such as drought, salinity, and poor soil fertility. Many landraces and local varieties have been collected from the desert areas of the Northwest Coast, Siwa, and Sinai. These collections are maintained in gene banks and utilized by different barley breeding programs for their valuable attributes. Until the beginning of the last century, farmers relied on their own local varieties developed by selecting superior plants from existing landraces. Each tribe had its own local variety (known as Bedouin variety) selected and maintained by the members of the tribe. Ullrich—Production, Improvement, and Uses c08.indd 214 8/30/2010 8:28:58 PM Barley Breeding History, Progress, Objectives, and Technology The irst organized efforts in barley breeding started with the establishment of the Department of Plant Breeding at Bahteem Agricultural Research Station in 1898 as a part of the Sultanic Agricultural Organization (Royal Agricultural Society). Until the 1940s, barley breeding relied mostly on the collection and selection from local varieties and landraces known as “Baladi varieties.” These efforts resulted in the selection of several high-yielding and disease-resistant varieties, such as Baladi 16, Bahteem 52, and Giza 24. Starting in the 1940s, barley breeders started to introduce germplasm from other countries, such as Abyssinia 12, Hungaria 1, and Palestine. Selection from this introduced germplasm resulted in the release of two new varieties, Giza 68 and Giza 73, in 1948. In 1956, a new extremely early barley variety (Hybrid 100) was developed, which is also known as Saharawi 100. This variety has been extensively used in the breeding program for earliness until now. It was the irst barley variety developed through crossing between Baladi-16 and Atsel. Another important variety was California Mariout, used side by side with Sahrawi 100 in the rainfed areas of Egypt. Barley breeders continued to cross local varieties and introductions to produce new varieties with better adaptation to low rainfall areas. For example, the variety Giza 117 was produced in 1959 as a selection from the cross Baladi 16 × Palestine 10, and the variety Giza 119 and Giza 121, produced in 1973 and 1980, respectively, as selections from the cross Baladi 16 × Gem. Another wave of barley varieties was developed in 1980s, such as Giza 121, CC-89, Giza 123, and Giza 124. The last two varieties replaced all the old varieties because of their high yield potential, especially in saline soils and under heat stress conditions, respectively. Barley breeding activities focus on screening of barley genotypes for drought and salinity tolerance and for disease and aphid resistance as well as heat tolerance. The germplasm is based on introductions from ICARDA and on local breeding material developed from local crosses. The program addresses three major environments, that is, the rainfed areas of the Northwest 215 Coast and Sinai, the new reclaimed land (irrigated) and saline soils, and the old irrigated areas. The program follows a traditional pedigree method with yield testing conducted on station in micro and macro yield trials in several locations. Seed multiplication is conducted at Sakha research Station under irrigation. In collaboration with the DRC, ICARDA has 120 conducted for the last 8 years a participatory barley breeding program in the Northwest Coast: this involves eight villages and has so far generated ive varieties, which are being multiplied by farmers (by law, they cannot be oficially multiplied), and this is so far the main factor limiting their wider adoption. EAsT AfRiCA The two most important countries for barley cultivation in East Africa are Ethiopia and Eritrea, two of poorest countries in the world. In both countries, agriculture is almost the only source of living for the majority (85%) of the population. Since Eritrea gained independence from Ethiopia in 1991, the history of barley breeding was common to the two countries before that date. Barley is one of the most important staple food crops and has a signiicant role in the diet of many millions of people. Cultivation of slightly more than 1 million hectares of barley occurs in the highlands of the two countries. The farmers of both countries grow barley because it has several advantages over other cereals: (i) barley can be grown in marginal areas where the choice of other cereals is limited; (ii) it offers the farmer an earlier crop harvest than most cereals, providing relief of food shortages, which frequently occur during the long rainy season; (iii) it has better stability of production over other cereals; (iv) it is a dependable food crop as it is grown in different seasons and production systems; (v) it is the preferred crop for the preparation of traditional drinks and beer; and (vi) its straw is a good source of feed and bedding for animals and thatching of roofs. Barley is believed to have been cultivated in Ethiopia and Eritrea as early as 3000 BC. (Gamst 1969). Both countries, but Ethiopia in particular, Ullrich—Production, Improvement, and Uses c08.indd 215 8/30/2010 8:28:58 PM F Barley: Production, Improvement, and Uses 216 have diverse climates, soils, topography, social environments, vegetation cover, and livestock. The long history of barley cultivation and the diverse agroecological and cultural practices have resulted in a large number of landraces and traditional agricultural practices. All the different types of barley are grown: hulled, hulless, sixrow, two-row, irregular forms, dense, lax, hooded, long awn, short awn, and rough and smooth awn (Asfaw 1988). In addition, barley landraces vary in their characteristics such as maturity, seed color, seed size, seedling vigor, straw strength, and in disease and insect pest resistance. Landraces are also grown in mixtures, often with other crops; the best known of these mixtures, called hanfets, is a barley–wheat mixture very popular in Tigray, the Northern part of Ethiopia, and in the highlands of Eritrea (Woldeamlak et al. 2001, 2008). Ethiopia and Eritrea are two of the centers of diversity of barley, but also of other important crops such as durum wheat, and therefore they have a very long history of barley breeding if we also include the millennia of selection conducted by farmers. Ethiopia In Ethiopia, barley research started in the early 1950s by the former Alemaya College of Agriculture and Mechanical Arts at its experimental station in Debre Zeit, with the evaluation of landraces and introduced nurseries. After the establishment of the Institute of Agricultural Research (IAR) in 1966, barley research was transferred from Debre Zeit to Holetta Research Center. After the restructuring of agricultural research with the establishment of a federal center (Ethiopian Agricultural Research Organization [EARO]) and of agricultural regional centers, barley research has been coordinated from the Holetta Research Center in collaboration with the regional research centers (at Kulumsa, Ambo, Sheno, Adet, Sirinka, Mekele, and Sinana), and other Ethiopian institutions. International collaborations include FAO, the United States Department of Agriculture (USDA), and the F Swedish Agency for Research and Education Cooperation (SAREC). Since the mid-1970s, the collaboration with ICARDA has been valuable for the exchange of information and germplasm and for capacity building. Between 1970 and 1990, nearly 14,000 local landraces were evaluated. Most of the entries were found susceptible to scald (caused by Rhynchosporium secalis Oud.), net blotch (caused by Helminthosporium teres Sacc.), spot blotch (caused by Helminthosporium sativum Pum.), leaf rust (caused by Puccinia hordei Otth.), and lodging. However, from this work, some outstanding varieties such as Shege, Misratch, and Abay were released in late 1990s (Lakew et al. 1997; Yitbarek et al. 1998). Between 1966 and 2001, nearly 30,000 exotic entries were evaluated. Most of them were found to be highly susceptible to scald, net and spot blotch, and barley shoot ly (Delia lavibasis) and had poor plant vigor and small grains. About 6% were selected for further study. In the early 1970s, the major contributors of germplasm were the FAO Near East Regional Program, the USDA, and the Arid Land Agriculture Development (ALAD), and from the mid-1970s, ICARDA. Germplasm has been also received from Brazil, Colombia, the former Czechoslovakia, Egypt, India, Kenya, Peru, Sweden, and the former Republic of Yugoslavia (Gebre et al. 1996). From these efforts, one hulled variety, AHOR 880/61, was released, and some other elite lines are still being used in the national crossing program as sources of genes for desirable agronomic traits such as grain quality and stiff straw and for disease and insect pest resistance. Between 1974 and 2001, over 1600 crosses (single, double, and three-way) were made between exotic and local germplasm and among local germplasm to improve their resistance to lodging and to major diseases such as scald, net blotch, spot blotch, and leaf rust. F2 progenies have been evaluated at a number of locations in the country. Only one outstanding hulled barley variety, HB 42, has been identiied from this program. Currently, some regional centers have started programs of participatory barley breeding. Ullrich—Production, Improvement, and Uses c08.indd 216 8/30/2010 8:28:58 PM Barley Breeding History, Progress, Objectives, and Technology Eritrea In Eritrea, barley research started with a germplasm collection conducted by the staff of the Department of Agricultural Research and Human Resource Development (DARHRD) in 1997 (shortly after independence) and with the repatriation of local germplasm held in the Ethiopian Gene Bank. Most of this work was funded by the government of Denmark. The barley breeding program started formally in 1998 with the evaluation of the landraces collected in 1997 and with targeted crosses made at ICARDA. In 1999, the irst ield trials were conducted in three villages, and three potential varieties were identiied. After an interruption caused by the war with Ethiopia, the barley breeding program started again and, since 2004, was funded by a project of the Challenge Program on Water 121 and Food of the CGIAR. In this second phase, after the systematic evaluation of all the landraces available in the gene bank, and of a number of experimental hanfets, the National Agricultural Research Institute (NARI), the new denomination of DARHRD, has started the ield testing of new breeding material derived from crosses made at ICARDA between local landraces and exotic germplasm. LATin AMERiCA Barley was introduced in the Americas by Columbus as early as his second trip to the New World. Barley was irst planted in 1493 by the Spaniards who stayed at Isabela, Puerto Rico and from there it was introduced to Mexico and the United States. It is hard to document the irst barley and wheat crops in South America, but it most probably was in the Fort of Sancti Spiritus in 1527, in the present day province of Santa Fe in Argentina (Arias 1995). From there, it spread to the Andes and the countries that share that mountain range. The irst scientiic works in barley and wheat in South America were carried out starting in 1912 in Uruguay by Drs. Alberto Boerger and Enrique Klein, selecting pure lines 217 resistant to leaf rust (gene Pa7) from populations being used by farmers. Barley production areas in Latin America can be divided in two main groups, based on ecological regions and end use: (i) the Andean region, with Bolivia, Ecuador, Colombia, and Peru, where the main uses of barley are for food, feed, and forage; and (ii) the countries of the Southern Cone of South America, with Argentina, Brazil, Chile, and Uruguay, plus México, where almost exclusively barley is made into malt for beer production. The production in the irst group is stable, whereas in the second, the production is increasing due to the higher demand of malt because of the rise in consumption of beer and malt derivatives in the continent and worldwide. After the establishment of the irst barley research program in the subcontinent in La Estanzuela, Uruguay, research has been carried out at different times in different countries in the region. Barley breeding at public institutions has always been an appendix to wheat breeding and has always counted on less economic and human resources. Despite this, an important volume of research and selection has been done in the Latin American countries where barley is cultivated at commercial scale (Arias 1995). At present, several public and private long-term, well-established, and successful breeding programs are working in Argentina, Brazil, Chile, Ecuador, México, Peru, and Uruguay. As the main varieties released were for malting, very few local genetic resources have been used in crossings because they were forage and feed barleys. Almost all cultivated barleys have spring habit, except in Chile, where some winter or facultative barleys have been released. Almost all cultivars are covered, except in Ecuador and Peru, where a few hulless varieties have been released. In the Andean countries, epidemics of stripe rust (caused by Puccinia striiformis f. sp. hordei) introduced from Europe in the late 1970s affected all varieties planted at that time, severely decreasing the production in all the highlands (Dubin and Stubbs 1985). All the native cultivars were highly susceptible, and the local breeding programs had to produce resistant cultivars in a relatively short period of time. At present, all released Ullrich—Production, Improvement, and Uses c08.indd 217 8/30/2010 8:28:58 PM F Barley: Production, Improvement, and Uses 218 cultivars must show resistance to this disease in the Andean countries and in Mexico. ACKnOWLEDGMEnTs The authors acknowledge the information on the history of barley breeding in their respective countries received from Dr. A.F. El-Sayed (Egypt), Dr. M. Noaman (Egypt), Dr. S. Saidi (Morocco), Mr. A. Shreidi (Libya), Dr. A. Benbelkacem (Algeria), and Mr. M. Maatougui (Algeria). REfEREnCEs 122 123 Al-Ahmad, H., W. El Taweel, and G. Some. 1999. The national agricultural research system of Syria. In J. Casas, M. Solh, and H. Hafez (eds.). The National Agricultural Research Systems in the West Asia and North Africa Region. ICARDA, FAO, AARINENA, and CIHEAM, Aleppo, Syria. Amri, A. 1993. Comparison of performances of barley, wheat and critical varieties. pp. 62–70. Project report IFAD/ ICARDA/Maghreb, Settat, January 1993. Arias, G. 1995. Mejoramiento Genético Y Producción De Cebada Cervecera En América Del Sur. Technical Cooperation Network for Food Crop Production. Plant Production and Protection Division, Regional Ofice of the FAO for Latin America and the Caribbean, Food and Agriculture Organization—FAO, ••. Asfaw, Z. 1988. Variation in the morphology of the spike within Ethiopian barley, Hordeum vulgare L. (Poaceae). Acta Agric. Scand. 38:277–288. Ben-Mahmoud, K.R. 1966. Institutions of Agricultural Education in Libya. Paper presented in the Workshop of Agricultural Education in the Arab Maghrib Union Countries, ••, May 22–23, 1996. Ceccarelli, S. and S. Grando. 2007. Decentralizedparticipatory plant breeding: an example of demand driven research. Euphytica 155:349–360. Dubin, H.J. and R.W. Stubbs. 1985. Epidemic spread of barley stripe rust in South America. Plant Dis. 70: 141–144. Gamst, C.F. 1969. Qemant: a Pagan-Hebraic Peasantry of Ethiopia. Holt, Rinehart and Winston, New York. Gebre, H., B. Lakew, F. Fufa, B. Bekele, A. Assefa, and T. Getachew. 1996. Food barley breeding. pp. 9–23. In H. Gebre and J. van Leur (eds.). Barley Research in Ethiopia: Past Works and Future Prospects: Proceedings, First Barley Research Review Workshop, IAR/ICARDA, 124 October 16–19, 1993, Addis Ababa, Ethiopia. ••, ••. Grando, S. and H. Gomez Macpherson (eds.). 2005. Food barley: importance, uses and local knowledge. In •• •• (ed.). 125 Proceedings: International Workshop on Food Barley Improvement, Hammamet, Tunisia, January 14–17, 2002. ICARDA, Aleppo, Syria. Grillot, G. 1939. Les meilleures variétés d’orge. Extrait de la Terre Marocaine N° 116. Lakew, B., S. Yitbarek, F. Alemayehu, H. Gebre, S. Grando, J.A.G. van Leur, and S. Ceccarelli. 1997. Exploiting the diversity of barley landraces in Ethiopia. Genet. Resour. Crop Evol. 44(2):109–116. Ministry of Agriculture and Fisheries. 1989. Agricultural research in the Yemen Arab Republic, Vol. 1: Problems, research activities, and evaluation of future programmes. Proposals for the third ive year plan 1987–1991. Taiz, 1989. Rivera, W.M. and M.A. Elkalla. 1997. Restructuring agricultural extension in the Arab Republic of Egypt. Eur. J. Agr. 126 Educ. Ext. 3(4):251–260. Roozitalab, M.H. 1996. Operational Partnership Framework for WANA-NARS Collaboration in Global Agricultural Research System. AREEO, Ministry of Agriculture, Tehran. Saade, E.M. 1994. Constraints to the adoption of new barley varieties in Morocco. p. 88. Report Project INRA/DPVMARA/ICARDA, Rabat. Saidi, S., A. Jilal, A. Amri, S. Grando, and S. Ceccarelli. 2005. Amelioration Genetique de l’Orge au Maroc. pp. 97–138. In F.A. Andaloussi and A. Chahbar (eds.). La Création Variétale A L’INRA: Methodologie, Acquis et Perspectives. Edition INRA, Rabat, Maroc. Taimeh, A. and S. Sunna. 1999. The national agricultural research system of Jordan. In J. Casas, M. Solh, and H. Hafez (eds.). The National Agricultural Research Systems in the West Asia and North Africa Regions. ICARDA, FAO, AARINENA, and CIHEAM, Aleppo, Syria. Woldeamlak, A., L. Bastiaans, and P.C. Struik. 2001. Competition and niche differentiation in barley (Hordeum vulgare) and wheat (Triticum aestivum) mixtures under rainfed conditions in the Central Highlands of Eritrea. Neth. J. Agric. Res. 49:95–112. Woldeamlak, A., S. Grando, M. Maatougui, and S. Ceccarelli. 2008. Hanfets, a barley and wheat mixture in Eritrea: yield, stability and farmer preferences. Field Crops Res. doi: 10.1016/j.fcr.2008.06.007. Yitbarek, S., B. Lakew, F. Alemayehu, S. Grando, J.A.G. van Leur, and S. Ceccarelli. 1998. Variation in Ethiopian barley landrace populations for resistance to barley leaf scald and net blotch. Plant Breed. 117:419–423. F Ullrich—Production, Improvement, and Uses c08.indd 218 8/30/2010 8:28:58 PM AUTHOR QUERY FORM Dear Author During the preparation of your manuscript for publication, the questions listed below have arisen. Please attend to these matters and return this form with your proof. Many thanks for your assistance. Query References 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. Query Remarks AUTHOR: Please conirm if all heading levels in this chapter are correct. AUTHOR: Please conirm if BaYMV should be deined as “barley yellow mosaic virus.” Please conirm also if this abbreviation is the same as BYMV. If so, please conirm which of these abbreviations should be used in the chapter. AUTHOR: Fischbeck 1992 has not been included in the reference list. Please supply full publication details. AUTHOR: Barley Malt News 2009 has not been found in the reference list. Please provide full reference details. AUTHOR: Please check this Web site address and conirm that it is correct. (Please note that it is the responsibility of the author[s] to ensure that all URLs given in this chapter are correct and usable.) AUTHOR: Please conirm if the phrase “actual German spring barley … superior malting quality” is correct. AUTHOR: Please conirm if the deinition of QTL is correct. AUTHOR: Please deine CAPS. AUTHOR: Please conirm f PCR should be deined as “polymerase chain reaction.” AUTHOR: Please deine GM. AUTHOR: Please deine EMS. AUTHOR: Please deine TILLING. AUTHOR: Anonymous (2009): This entry has been styled as a book chapter. Is this correct? If so, please provide the chapter title and the editors’ surnames and initials. AUTHOR: Behn et al. 2004 has not been cited in the text. Please indicate where it should be cited or delete it from the reference list. F Ullrich—Production, Improvement, and Uses c08.indd 1 8/30/2010 8:28:58 PM 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. F AUTHOR: Friedt and Rasmussen. (2004): Please provide the article title and the date of the proceeding (month, day, and year). AUTHOR: Friedt et al. (2000): This reference has been styled as a book chapter. Is this correct? If so, please provide the editors’ surnames and initials, the publisher name, and the city of publication. AUTHOR: Habekuss et al. (2009): Please check if the abbreviated journal title is correct. AUTHOR: Please provide the volume number and page range for Humbroich et al. (2009) if available. AUTHOR: Schmalenbach et al. (2009) has not been cited in the text. 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AUTHOR: Roelfs and Huerta-Espino 1994 has been changed to Roelfs and Huerto-Espino 1994 so that it matches the reference list. Please conirm that this is correct. AUTHOR: Please check “to uptake suficient water.” Do you mean “to take up suficient water”? AUTHOR: Please conirm if the sentence “Finally, nonmalting or feed barley … quality testing is not required” is correct. Ullrich—Production, Improvement, and Uses c08.indd 2 8/30/2010 8:28:58 PM 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. AUTHOR: Please conirm if PCR should be deined as “polymerase chain reaction.” AUTHOR: Please conirm if the phrase “malt quality is compared to the checked cultivars” is correct. AUTHOR: Please check “The composition of these yield trials … by a test coordinator.” Do you mean “The composition of these yield trials, including checks, is approved by the committee annually and the tests are administered by a test coordinator”? AUTHOR: Please provide the abbreviated journal title, the volume number, and the page range for Chen (2004). AUTHOR: Please update the publication year for Horsley et al. (2008). AUTHOR: Jorgensen (1992) has not been cited in the text. Please indicate where it should be cited or delete it from the reference list. AUTHOR: Mornhinweg et al. (1999): Please conirm if the page range is correct. AUTHOR: Please provide the editors’ initials for Prom et al. (1996). AUTHOR: Weaver (1950) has not been cited in the text. Please indicate where it should be cited or delete it from the reference list. AUTHOR: Sparrow and Doolette 1987 has not been included in the reference list. Please supply full publication details or delete these citations from the text. AUTHOR: Please conirm if “The areas of greatest production are … the Darling Downs of Queensland” is correct. AUTHOR” “mt” in all instances has been changed to “MT.” Please conirm that this is correct. AUTHOR: Please conirm if South Australia, Western Australia, Victoria, New South Wales, Queensland, and Tasmania should be abbreviated throughout the text as SA, WA, Vic, NSW, Qld, and Tas, respectively. AUTHOR: “Northern Region,” “Southern Region,” and “Western Region,” respectively, have been lowercased throughout the text. Please conirm if this is correct. AUTHOR: AAB Grain 2008 has been changed to ABB Grain 2008 so that it matches the reference list. Please conirm that this is correct. AUTHOR: Please conirm if the phrase “The result of the signiicant investment … grain handling organizations and authorities” is correct. F Ullrich—Production, Improvement, and Uses c08.indd 3 8/30/2010 8:28:58 PM 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. F AUTHOR: Please conirm if “Germplasm from the NSWDPI, Victorian and Tasmanian breeding programs is being progressed” is correct. AUTHOR: Please deine HACCP. AUTHOR: Please check the Web site addresses in the chapter and conirm that they are correct. (Please note that it is the responsibility of the author[s] to ensure that all URLs given in this chapter are correct and usable.) AUTHOR: Barr and Kneipp 1995 has not been included in the reference list. Please supply full publication details. AUTHOR: Please check if “Yield, lodging resistance, … stem breakage” should be changed into a sentence. AUTHOR: Poulsen, unpublished data: Please provide the initial(s) of Poulsen. AUTHOR: Poulsen et al. (1995): Please indicate whether the year is 1995a or 1995b. AUTHOR: Please conirm if “*” is correct or if it should be changed to “×” in all instances. AUTHOR: Franckowiak, pers. comm.: Please provide the initial(s) of Franckowiak. AUTHOR: Please deine NBIP. AUTHOR: Dill Macky, 1992 has been changed to Dill-Macky, 1992 so that it matches the reference list. Please conirm that this is correct. AUTHOR: Harlan et al. 1922 has not been included in the reference list. Please supply full publication details. AUTHOR: H. v. spontaneum has been changed to H. vulgare spontaneum. Is this correct? AUTHOR: Rees, pers. comm.: Please provide the initial(s) of Rees. AUTHOR: Cotterill, 1992b has been changed to Cotterill et al. 1992 so that it matches the reference list. Please conirm that this is correct. AUTHOR: Please deine RIL. AUTHOR: Please conirm that the full form of QTL is correct. AUTHOR: Please conirm if the deinition of SSR is correct. AUTHOR: Please conirm if the phrase “enabled the alignment of markers and the creation of consensus maps of the barley genome” is correct. AUTHOR: Please deine RFLP and AFLP. AUTHOR: Please deine BSA. Ullrich—Production, Improvement, and Uses c08.indd 4 8/30/2010 8:28:58 PM 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. AUTHOR: Coventry et al. (2003): “a” and “b” have been inserted after the year of publication. Please conirm that this is correct. AUTHOR: Cullis et al. (2003) has not been included in the reference list. Please supply full publication details. AUTHOR: Cakir et al. 2003: Please indicate in all instances in the text whether the years are 2003a, 2003b, or 2003c. AUTHOR: Please deine RAPD. AUTHOR: Please deine CIMMYT. AUTHOR: Please deine STS. AUTHOR: Borovkova et al. 1997: Please indicate whether the year is 1997a or 1997b. AUTHOR: Please conirm if PCR should be deined as “polymerase chain reaction.” AUTHOR: Anderson and Reinbergs (1985): Please conirm if the book title and the publisher names are correct. Also, please provide the publisher location. AUTHOR: Barley Australia (2008) has not been cited in the text. Please indicate where it should be cited or delete it from the reference list. AUTHOR: Barley Breeding Australia (2008) has not been cited in the text. Please indicate where it should be cited or delete it from the reference list. AUTHOR: Barr et al. (2001): Please conirm if the initial of Barr is correct and please provide the month, day, and year of the proceeding. AUTHOR: Bowman and Blake (1996): Please conirm if the book title and the publisher location are correct. Also, please provide the publisher name. AUTHOR: Box and Barr (2000): Please conirm if the book title and the publisher location are correct. Also, please provide the publisher name. AUTHOR: Box et al. (1997): This reference entry has been styled as a book chapter. Is this correct? If so, please provide the editor name(s) and initial(s) and the publisher name. Please conirm also if the book title and the publisher location are correct. AUTHOR: Boyd et al. (1981): This reference entry has been styled as a book chapter. Is this correct? If so, please provide the editor name(s) and initial(s), the publisher name, and the publisher location. AUTHOR: Broughton and Poulsen (1997): Please conirm if the page range is correct. F Ullrich—Production, Improvement, and Uses c08.indd 5 8/30/2010 8:28:58 PM 85. 86. 87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. F AUTHOR: Brown et al. (2000): Please provide the publisher and its location. AUTHOR: Cakir et al. (2001): Please provide the month, day, and year of the proceeding. AUTHOR: CBH Group (2008): Please conirm that this reference entry is a book. If so, please provide the publisher and its location. AUTHOR: Chicaiza et al. (1996): Please conirm that this is a book chapter. If so, please provide the publisher name and please conirm that the book title and the publisher location are correct. AUTHOR: Coventry et al. (2003): Two reference entries share the same author and year of publication. Hence “a” and “b,” respectively, have been added to the years to distinguish these entries in the text. AUTHOR: Please provide the editor name(s) and initial(s) for Dill-Macky et al. (1989). AUTHOR: Emebiri et al. (2003): Please conirm that the year of publication is correct. AUTHOR: Falk (2001): Please conirm that this is a book chapter. If so, please provide the publisher name and please conirm that the book title and the publisher location are correct. AUTHOR: French and Schultz (1984a,b): Please conirm if the article titles are correct. AUTHOR: Graner et al. (2000): Please conirm if the book title and the publisher location are correct and please provide the publisher name. AUTHOR: Jefferies et al. (1997): Please provide the editor name(s) and initial(s), the publisher name, and the publisher location. AUTHOR: Johnston (1983): Please provide the editor name(s) and initial(s), the publisher name, and the publisher location. AUTHOR: Karakousis et al. (2000): Please provide the publisher name. Also, please conirm if the book title and the publisher location are correct. AUTHOR: Please provide the article title for Karakousis et al. (2001). AUTHOR: Kasha and Reinbergs (1982): Please provide the publisher name. Also, please conirm if the book title and the publisher location are correct. AUTHOR: Kasha et al. (1992): Please provide the publisher name and the publisher location. Ullrich—Production, Improvement, and Uses c08.indd 6 8/30/2010 8:28:59 PM 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. AUTHOR: Malting and Brewing Industry Barley Technical Committee (MBIBTC) (1998): Please provide the publisher location and please conirm if the publisher name is correct. AUTHOR: Poulsen et al. (1995b): Please provide the publisher name and the publisher location. AUTHOR: Poulsen et al. (1997): Please conirm if the page range is correct. AUTHOR: Poulsen et al. (2000) has not been cited in the text. Please indicate where it should be cited or delete it from the reference list. Also, please provide the publisher name and conirm if the publisher location is correct. AUTHOR: Raman et al. (2003): Please conirm if the publication year is correct. AUTHOR: Riggs et al. (1982): Please provide the publisher name and the publisher location. AUTHOR: Please provide the city location for Sharp (1985). AUTHOR: Stuthman et al. (1996): Please provide the publisher name and please conirm if the publisher location is correct. AUTHOR: Please check this Web site address and conirm that it is correct. (Please note that it is the responsibility of the author[s] to ensure that all URLs given in this chapter are correct and usable.) AUTHOR: Please note that two deinitions of DASR have been found in the text. Please conirm which of these should be used in the chapter. AUTHOR: Please conirm if “Department of Scientiic Agricultural Research” is correct or if it should be changed to “Department of Agricultural Scientiic Research.” AUTHOR: SPPI has been changed to SPII. Is this correct? AUTHOR: Please deine CWSR. AUTHOR: Please deine UNDP and IDA. AUTHOR: Grando and Gomez-Macpherson, 2005 has been changed to Grando and Gomez Macpherson 2005 so that it matches the reference list. Please conirm that this is correct. AUTHOR: Harrabi, pers. comm.: Please provide the initial(s) of Harrabi. F Ullrich—Production, Improvement, and Uses c08.indd 7 8/30/2010 8:28:59 PM 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. F AUTHOR: Please conirm if “three periods” in the phrase “The history of breeding can be divided into three periods” should be changed to “two periods” as only two periods were described in the text. AUTHOR: Please spell out AR in “Regional AR Centers.” AUTHOR: Please check the phrase “and to 92 these days dealing with various aspects of agricultural production.” Should this be changed to “and to 92 at present, dealing with various aspects of agricultural production”? AUTHOR: Please deine DRC. AUTHOR: Please deine CGIAR. AUTHOR: Please provide the publisher location for Arias (1995). AUTHOR: Ben-Mahmoud (1966): Please provide the place (city and country) where the workshop was held. AUTHOR: Gebre et al. (1996): Please provide the publisher and its location. AUTHOR: Grando and Gomez Macpherson (2005): Please provide the editor name(s) and initial(s). AUTHOR: Please conirm that the abbreviated journal title is correct in Rivera and Elkalla (1997). AUTHOR: Please deine DUS. AUTHOR: Fig. 8.1.4: “Author” has been changed to “Source.” Is this okay? AUTHOR: Please provide suitable legends for Figs. 8.3.1 and 8.3.2. AUTHOR: Fig. 8.3.2: The deinitions of A, Y, Q, and D have been deleted from the igure and moved to the igure legend. Is this okay? AUTHOR: Table 8.1.2: Please conirm if the area harvested in Malta (400F) is correct. AUTHOR: Table 8.1.2: Please conirm if the sentence “Of the total global annual seed … (29%) in the EU (27)” is correct. AUTHOR: Table 8.1.3: n.d. has been deined as “not detected.” Is this correct? AUTHOR: Table 8.1.4: Please conirm if Pyrenophora teres is correct. AUTHOR: Table 8.2.2: The symbols †, ‡, and § have been changed to superscript a, b, and c, respectively, as per book style for table footnotes. Is this correct? Please provide the meaning of these linking letters. Ullrich—Production, Improvement, and Uses c08.indd 8 8/30/2010 8:28:59 PM 136. 137. 138. 139. AUTHOR: Table 8.2.2: Please conirm if “db” should be deined as “dry basis.” AUTHOR: Table 8.3.1: Please conirm if the igure “000 t” is correct. AUTHOR: Table 8.3.2: Please conirm if the igure “’000” in the table is correct. AUTHOR: Table 8.3.4: Please spell out the genus name in “P. sp.” in this table. F UBA 08 Ullrich—Production, Improvement, and Uses c08.indd 9 8/30/2010 8:28:59 PM