Contents

• Introduction
• Breeding
• Pathology
• Cytogenetics
• Biotechnology
• Physiology
• Agronomy
• Animal Nutrition
• Malting and Food Barley
• Summary
• Acknowledgments
• References
• Appendix 1

Introduction

Barley (Hordeum vulgare L.) has been grown in Eastern Canada for almost four hundred years. First grown in Port Royal in 1606 (Hamilton 1955), barley is now grown in all six provinces of Eastern Canada. Eastern Canada produced 1,000,700 tonnes of barley grains and 438,700 tonnes of mixed grains (mainly mixtures of barley and oats) in 1998 (Statistics Canada 1998). Despite this, Eastern Canada, on average, purchased 364,400 tonnes of barley annually from Western Canada under the Feed Freight Assistance Program (Canada Grains Council 1997). Barley is used mainly for livestock feed. Almost all of it is consumed by dairy cattle (44%), hogs (38%) and beef cattle (16%) (Statistics Canada 1995). Canada Malting purchases malting barley from Western Canada for its Montreal and Thunder Bay plants, which have the capacity to process annually 81,700 and 144,700 tonnes of barley, respectively (Canada Malting 1995).

Barley in Eastern Canada is of spring type; some winter barley is also grown, exclusively in southern Ontario. Six-row barley predominates because of its high yield (Jui et al. 1997). Some six-row varieties, however, are high in fibre (Kong et al. 1995) and thus hog producers prefer to buy two-row varieties (Anonymous 1997). Until now, only covered barley is grown commercially in Eastern Canada. This will change in the future with the impending release of the first hulless barley variety (AB168-11) for Eastern Canada. Barley research is conducted by federal and provincial governments, universities, as well as the private sector. This article provides an overview of the current research activities on barley across Eastern Canada.

Breeding

(1). Breeding Programs. Agriculture and Agri-Food Canada has consolidated its previous barley breeding efforts in one single program at the Eastern Cereal and Oilseed Research Centre (ECORC). The goals of the ECORC's program are to develop barley varieties (including both two-row and six-row, and both covered and hulless) for Eastern Canada and to develop early maturing varieties for northern Ontario, northern Quebec and Newfoundland. The University of Guelph has been funded by the Ontario Ministry of Agriculture, Food, and Rural Affairs to develop six-row barley varieties for Ontario. Besides these two public breeding programs, six private companies in the region are actively involved with barley breeding. W. G. Thompson and Sons Limited aims at feed barley varieties (both two-row and six-row) for Eastern Canada, and two-row (for malt) and six-row (for malt and feed) varieties for Western Canada. Thompson also runs the only winter barley breeding program in Canada to develop six-row varieties for Ontario and north-eastern United States. Similarly, Semico Inc. runs a strong private breeding program to develop six-row varieties for Eastern Canada. Coopérative Fédérée de Québec is breeding six-row barley. Prograin Inc. conducts its six-row barley breeding in partnership with Laval University. C & M Seeds works on six-row and two-row barley breeding in collaboration with the University of Guelph and a German company, respectively. It is also searching for malting barley and hulless barley. Minas Seed Co-op Limited is introducing European varieties/lines to Canada.

The barley breeding programs in Eastern Canada have been very successful. The yields of barley varieties released from 1935 to 1988 increase at a rate of about 0.03 t/ha/yr (Bulman et al. 1993). Twenty-nine of the 76 barley varieties on the 1990 List of Registered Varieties originated in Eastern Canada (Kong et al.1994). Some of the eastern varieties yield well in both Eastern and Western Canada (Kong et al. 1994). Since 1990, Eastern Canada has registered 32 spring barley and 2 winter barley varieties (Appendix 1).

(2) Breeding Objectives. The above breeding programs aim at improving specific traits of barley. The important traits for spring barley for Eastern Canada are: high grain yield, disease resistance, lodging resistance, resistance to neck break, good threshability, high protein, high test weight, low fibre, early maturity, high straw yield, tolerance to water/heat stress, and tolerance to low pH. For winter barley, winter survival, barley yellow dwarf virus (BYDV), and scald resistance are the major challenges.

(3) Breeding Methods. A variety of breeding methods are used by barley breeders in Eastern Canada. The most common one is the modified bulk method. Guelph, however, is using male sterile facilitated recurrent selection to conduct its breeding program (Kannenberg and Falk 1995). Kasha and Kao (1970) developed the bulbosum method for producing doubled-haploid (DH) lines. Since then, Eastern Canada has used it to develop 20 barley varieties (Table 1). Ouédraogo et al. (1998) developed an amino acid enriched FHG medium for barley anther culture. The medium is now used routinely by Laval to produce DH lines. In vitro selection for resistance to biotic and abiotic stresses is also used by Laval to develop new barley lines (St-Pie`rre et al. 1998). Recently, Kasha et al. (1997) perfected the isolated microspore culture system for barley, and ECORC is attempting to use it to produce DH lines for breeding barley. A callus derived line (RE80) recently received support for registration from the Ontario Cereal Crops Committee.

Pathology

(1). Diseases. Barley suffers many diseases in Eastern Canada, with the most important ones being net blotch, spot blotch/common root rot, scald, powdery mildew, fusarium headblight, BYDV, leaf rust, stem rust, loose smut, covered and semi-loose smuts, and ergot. Net blotch could reduce yield by 15% (Martin 1985) and increase kernel discoloration (Edney et al. 1998). The epidemiology of Pyrenophora teres is being studied by the Crops and Livestock Research Centre (CLRC). Pathotypes of P. teres in Eastern Canada remain unknown. It is believed that they are similar to the isolate WRS102. The most prevalent isolates of Erysiphe graminis are those virulent to genotypes with resistance genes Ml-h, Ml-p, Ml-a1, and/or Ml-k (Louter 1991). OAC Kippen possesses three resistance genes (Ml-a1, Ml-h, and Ml-g), but it became susceptible within a few years after it was released. Falak (1994) attributed this to an increase of isolates that are virulent to genotypes with the Ml-g gene. Many two-row varieties, however, show durable resistance to powdery mildew. Some callus derived lines were found to be resistant to powdery mildew (Choo et al. 1998). Fusarium infection is higher in barley than wheat in Quebec (15% vs. 11%) (Couture 1982) and in Prince Edward Island (65% vs. 52%) (Sturz and Johnston 1985). The most frequently isolated fusaria from barley ear tissues are F. graminearum (30%), F. poae (25%), F. avenaceum (16%), and F. culmorum (8%) (Sturz and Johnston 1985). Despite high infection levels, fusaria seem to have very little effect on barley yield (Sturz and Johnston 1985). Effects of fusarium infection on the DON content, however, remain to be investigated. Some barley seeds or products from Quebec in 1992-94 contained up to 6 mg/kg of DON and 24 mg/kg of ochratoxin A (Canadian Food Inspection Agency, Mycotoxin Databank). Mycotoxins at these levels would have deleterious effects on animal health and productivity (Friend and Trenholm 1988). The impact of weather conditions on the airborne spore load of Fusarium sp. is being studied by CLRC.

(2) Control. Currently, resistant varieties to major diseases are very limited. AC Kings is resistant to WRS102 of P. teres, whereas ACCA was found to be resistant to BYDV (Yd2) and to isolate 1493 of Rhynchosporium secalis. Both AC Sterling and Morrison are resistant to powdery mildew. Winthrop is resistant to ergot. ECORC and CLRC are screening barley accessions for fusarium headblight and DON content. CLRC and New Liskeard are evaluating seed treatments and/or foliar fungicides for controlling diseases for both covered and hulless barley. The Nova Scotia Agricultural College (NSAC) is also studying seed treatments. The Soils and Crops Research and Development Centre (SCRDC) is searching for tolerance to root rot, BYDV, and waterlogging.

Cytogenetics

Xu and Kasha (1992) transferred a dominant gene for powdery mildew resistance from H. Bulbosum to cultivated barley. At ECORC, resistance to QCC of stem rust was transferred from H. bulbosum to cultivated barley. Populations are being developed for inheritance studies and gene tagging. The Hordeum-Secale amphyploid and Hordeum-Elymus hybrids will be backcrossed to barley in order to transfer tolerance to biotic and abiotic stresses to barley (Fedak and Petroski 1996). The meiotic behaviour of these interspecific hybrids are under investigation. H. spontaneum accessions are being screened for salinity tolerance. Laval will continue its work on interspecific hybridization and novel germplasm development.

Biotechnology

(1) Variety Identification. Barley varieties were successfully distinguished on the basis of random amplified polymorphic DNA (RAPD) (Tinker et al. 1993; Baum et al. 1998), restriction fragment length polymorphism (RFLP) (Molnar and McKay 1995), and simple sequence repeats (SSRs) (William et al. 1997). Baum et al. (1998) published an identification key for all the 65 registered six-row barley varieties in Canada. The transferability of SSRs across species and their stability over time are being investigated by Guelph. It is now possible to use DNA extracted from single seeds for variety identification (William et al. 1997; Baum et al. 1998).

(2) Molecular Systematics. Baum et al. (1997) used RAPDs to study molecular diversity in H. spontaneum. Molnar et al. (1992) used RFLPs to analyze the phylogenetic relationships in the genus Hordeum. Baum and Johnson (1994; 1996; 1998) studied the 5S rRNA gene thoroughly in many Hordeum species. Use of molecular markers to study the phylogenetic relationship among the Hordeum species will be continued at ECORC. The extent of synergy between barley and oat will be investigated.

(3) QTL Analysis. QTL analysis for the Harrington/TR306 cross led by Kasha was completed (Tinker et al. 1996; Mather et al. 1997; Spaner et al. 1998). The genome mapping project is currently led by Mather with an objective of testing strategies for using molecular markers in the development of barley lines with good adaptation and malting quality. Studies on QTL mapping, QTL verification, and marker-assisted selection for QTL are being conducted at McGill. Two DH populations have been created by ECORC to identify QTLs affecting agronomic traits, physiological traits, disease resistance, and feed quality of barley. Single seed descent lines are being developed for the purpose of tagging fusarium resistance gene(s). Attempts to tag net blotch resistance genes are in progress at ECORC using RAPD and sequence characterized amplified regions. Three independent loci that are resistant to loose smuts have been tagged; further work is in progress at Laval to perform interval mapping in order to more precisely locate these loci and to assess their respective contributions to resistance.

(4) Transformation. Transgenic barley plants have been produced by bombardment of isolated microspores (Yao et al. 1997; Yao and Kasha 1997). Selection of transgenic plants was achieved using the herbicide Basta and the antibiotic geneticin in the medium and by visual selection using the green fluorescent protein. Efforts are continuing at Guelph to improve the frequencies of transgenic plants using both bombardment and Agrobacterium methods.

Physiology

Research on intensive cereal management has been conducted at McGill. Post-anthesis application of ethephon increases yield (Ma and Smith 1992). Higher rates of nitrogen fertilizer applied at seeding or at awn emergence increase grain protein concentration (Bulman and Smith 1993). Grain protein accumulation seems to be more limited by the ability of roots to take up N from the soil than by the seed to take up N from the rest of the plant (Foroutan-pour et al. 1997). The final conclusion of their research is that the short growing season in Eastern Canada does not allow much time for barley to respond to really high levels of N fertilizer and the application of ethephon. The feasibility of using physiological traits as selection criteria for high yield and using canopy reflectance for early prediction of grain yield are being evaluated. Barley is being used by Laval and NSAC as a indicator plant to estimate the uptake of nutrients.

Agronomy

A multi-discipline, multi-institute project on mixed grains is in progress to identify the best varietal combinations of barley and oat for commercial production in Eastern Canada and to assess the benefits of mixed grains over pure stands. In a rotation study, Johnston (1997) investigated the effects of management practices on the forage yield and quality of barley and on subsequent alfalfa establishment. CLRC is studying the effects of seeding rate, variety, and management practices on the total production of both barley and red clover and to assess the usefulness of barley as a cover crop after potatoes in the control of soil erosion. Three rotation systems for corn were compared by the St-Hyacinthe Research Station. Results showed that corn in the second year yielded more in the barley-corn or soybean-corn system than in the corn-corn system and that soybean did not supply as much as barley did for corn that followed because of nitrate leaching on soybean. Recently, the Atlantic Cool Climate Crop Research Centre (ACCCRC) initiated an active research program to develop sustainable feed grain production and utilization systems for cool climates. CLRC is screening barley varieties for tolerance to low pH for use in potato rotation. A study is conducted by Emo (Ontario) to assess the effects of paper mill biosolids on barley if they are used as soil conditioner.

Animal Nutrition

(1) Grains. The chemical composition of 76 Canadian barleys has been studied (Kong et al. 1995; Narasimhalu et al. 1995). Six-row breeding materials from ECORC are now screened for high protein and low fibre. ECORC is developing Grainspec calibration for estimating the protein content of hulless barley. The Atlantic Veterinary College wants to find ways of improving the feeding value of barley for pigs. For pigs, application of xylanase improves digestibility and availability of nutrients in barley and hulless barley based diets. Shur-Gain's research also focuses on anti-nutritional or negative nutritional factors in barley for monogastric species. Its goal is to develop better formulation methods which counter these negative factors and identify feeding situations where these factors are of concern.

(2) Straw and Silage. Two-row barley yields more straw than six-row barley (Narasimhalu et al. 1998). Two-row barley straw is higher in rumen degradability than six-row barley straw (Salgado et al. 1995). Preliminary results from Guelph showed that production performance of dry pregnant beef cows is not affected with diets containing up to 75% of barley straw from two-row varieties and that there was no difference in production performance between control (alfalfa haylage) and mix diet (25% alfalfa haylage and 75% barley straw). Eventually, Guelph wants to develop an environmentally friendly approach for improving the utilization of straw for ruminants. Silage barley was evaluated by New Liskeard for yield and quality. The first forage barley variety (Summerville) was registered in Canada by W. G. Thompson and Sons Limited.

Malting and Food Barley

Several scientists have collaborated with Canada Malting in identifying malting barley varieties for Eastern Canada. Two malting barley varieties (B1602 and Robust) show promise in Eastern Canada. In food barley research, the Southern Crop Protection and Food Research Centre (SCPFRC) aims at developing novel techniques for processing barley into bran and flour fractions; exploring the use of bran and flour fractions in food and non-food applications; and evaluating the content, molecular weight distributions and rheological properties of ß-glucans.

Summary

In summary, the goals of the barley research in Eastern Canada are: (1) to develop new technologies in order to speed up the breeding process, to increase selection precision for economically important traits, to facilitate gene transfer, and to protect intellectual property rights; (2) to better understand the behaviour of pathogens and to devise environmentally and economically sustainable approaches for controlling diseases; (3) to explore the genetic diversity in barley and its related species, and to transfer the resistance/tolerance genes from wild species to barley; (4) to develop sustainable cultural practices for barley production and to find ways of optimizing barley production under unfavourable environments; (5) to identify value-added products for feed and for food, and to reduce/neutralize anti- nutritional factors in livestock feed; and ultimately (6) to develop superior barley varieties for feed, food, and malting purposes.

Acknowledgements

The author is grateful to the following scientists for written input regarding their respective barley research which was provided during the preparation of this manuscript: C. Azar (Coop Fédérée), B. R. Baum (ECORC), F. Belzile (Laval), C. D. Caldwell (NSAC), F.-P. Chalifour (Laval), J. Collin (Laval), H. Chrétien (Prograin), P. E. Colucci (Guelph), A. Comeau (SCRDC), S. Cui (SCPFRC), L. M. Dwyer (ECORC), G. Fedak (ECORC), J. Frégeau-Reid (ECORC), S. Guertin (St-Hyacinthe), K. M. Ho (ECORC), J. Johnston (New Liskeard), K. J. Kasha (Guelph), B. L. Ma (ECORC), R. A. Martin (CLRC), D. E. Mather (McGill), E. McMillan (Shur-Gain), S. J. Molnar (ECORC), J. Rowsell (New Liskeard), L. Shugar (W. G. Thompson), D. Smith (McGill), D. Spaner (ACCCRC), E. Sparry (C & M Seeds), C. A. St-Pierre (Laval), and T. Van Lunen (AVC).

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Appendix 1.
Eastern Canada barley varieties registered from 1991 to 1998.

1. Agriculture and Agri-Food Canada:
   AC Alma, AC Burman, AC Hamilton, AC Legend, AC Nadia, AC Stephen, AC Westech, AC Kings, AC Parkhill, AC Queens, AC Sirius, AC Sterling, and Almonte.
2. University of Guelph:
   Codac.
3. W. G. Thompson and Sons Limited:
   Beluga, Brucefield, Grant, Sandrine, TB891-6, Sommerville (forage), Belmore, Breslau, Sunderland, McDiarmid (winter), and McGregor (winter).
4. Semico Inc.:
   Bella, Gamine, Myriam, and Serena.
5. C & M Seeds:
   Formosa, Frin, and Viking.
6. Laval University:
   ACCA
7. McGill University:
   Labelle.