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Agronomic characteristics, malt quality, and disease resistance of barley germplasm lines with partial Fusarium head blight resistance.

FUSARIUM HEAD BLIGHT has caused devastating losses in both the yield and quality of barley produced in the northern Great Plains from 1993 to 2003. From 1993 to 1997, total losses due to FHB in the upper Midwest of the United States exceeded $200 million (U.S. GAO, 1999). Nganje et al. (2001) estimated losses of $136 million in the same region from 1998 to 2000. The greatest losses were due to reductions in yield and grain quality. Much of the reduced grain quality was due to the accumulation of the mycotoxin DON produced by the pathogen, F. graminearum. Depending on the purchaser of the grain, barley samples with DON concentrations as low as 0.6 [micro]g [g.sup.-1] have been rejected. DON has been found to carry through malting and brewing into finished beer (Schwarz et al., 1995). Even beer with low DON levels poses a marketing problem for brewing companies because nobody wants to consume a "toxin" and DON has been associated with beer gushing (Schwarz et al., 1995). Beer gushing or overfoaming is the most easily identifiable consumer complaint associated with DON content.

Chemical and cultural management strategies for FHB in barley have been unsuccessful; thus, development of improved cultivars with genetic resistance to the disease offers the greatest potential for controlling this disease. The inheritance of FHB resistance is not well understood; however, some progress has been made in breeding for resistance to FHB in barley. Thirty-five six-rowed and two-rowed barley germplasm lines have been identified with partial resistance to FHB (Prom et al., 1996). Most of these accessions originate from eastern Asia and have the two-rowed spike morphology. Based on our initial evaluations of these lines, they appeared to be unadapted for growth in northern Great Plains of the United States and the western Prairie Provinces of Canada because of late maturity and tall, weak straw.

Many of these germplasm lines are being used as sources of genes for FHB resistance and low DON accumulation; yet, little is known about their agronomic and malt quality characteristics and their response to other foliar pathogens. Information in these areas is needed so barley breeders can make informed decisions when working with segregating populations developed from crosses to these resistant germplasm lines. The objective of this study was to compare the agronomic characteristics, malt quality, and reaction to foliar pathogens of 35 barley germplasm lines with partial FHB resistance to current cultivars grown in the northern Great Plains.

MATERIALS AND METHODS

Plant Materials

Forty barley germplasm lines (Table 1), 35 with partial resistance to FHB (B. Steffenson, personal communication, 1996; Prom et al., 1996), were used in this study. Susceptible checks used in the study for comparison purposes included the two-rowed adapted cultivars Logan and Conlon and the adapted six-rowed cultivars Foster, Morex, and Stander. Conlon, Foster, and Morex are on the "2004 List of Recommended Malting Barley Varieties" by the American Malting Barley Association (Milwaukee, WI) when grown in the northern Great Plains. Logan and Stander are high-yielding feed barley cultivars currently grown in North and South Dakota.

Agronomic and Fusarium Head Blight Evaluations

Trials for the agronomic comparisons were sown on 15 May 1998 and 4 May 2000 at Fargo, ND, in a fine, montmorillonitic, frigid Typic Haploboroll soil; on 18 May 1999 and 26 April 2000 at Osnabrock, ND, in a fine, montmorillonitic Udic Natriboroll soil; and on 27 April 2000 at Langdon, North Dakota in a fine, montmorillonitic Udic Natriboroll soil. Entries were assigned to experimental units using a randomized complete block design (RCBD) with two replicates. The experimental units consisted of three 3.38-m rows of barley spaced 0.83 m apart. To obtain a valid estimate of the agronomic potential of these germplasm lines under disease-free conditions, they were protected against FHB infection and other foliar diseases with foliar applications of the fungicide benomyl (0.5 kg a.i. [ha.sup.-1]), beginning one week after heading and once a week for three consecutive weeks. The applications were done with a tractor-mounted sprayer using water at 159 L [ha.sup.-1] at 207 kPa and a ground speed of 5.6 km [h.sup.-1].

Morphological and agronomic data were collected throughout the growing season. Days to heading were recorded as the number of days after 31 May when 50% of the spikes were fully emerged from the boot. Between maturity and harvest, data on plant height (stem plus spike-excluding awns), lodging (1 = no lodging, 9 = severe lodging), and spike row type (vrs1vrs1Int-cInt-c = sessile six-rowed, Vrs1Vrs1int-cint-c = normal two-rowed, and Vrs1Vrs1Int-cInt-c = hybrid two-rowed) were recorded.

At maturity, plots were harvested using a plot combine. Grain samples were dried in a forced dryer to approximately 100 g [kg.sup.-1] moisture, de-awned, and cleaned. Yield of clean grain was recorded as megagrams per hectare (Mg [ha.sup.-1). Data were also recorded on test weight (kg h[L.sup.-1]), kernel plumpness (g [kg.sup.-1]), and grain protein concentration (g [kg.sup.-l]). Grain protein concentration was determined on a dry matter basis by near infrared reflectance measurement using a Tecator Infratec 1226 grain analyzer (Perstorp Analytycal Inc., Silver Spring, MD) and expressed in milligrams per kilogram (mg [kg.sup.-1]). Kernel plumpness was determined according to the method specified by the American Society of Brewing Chemists (ASBC, 1992). Kernels retained on a sieve with 0.2 by 1.9 cm slotted openings were considered plump. Kernel brightness was determined using a modification of the ASBC standard method, Barley-9 (ASBC, 1992), using the L-value obtained from a Pacific Scientific XL-800 series Gardner colorimeter with XL-845 circumferential sensor (Perstorp Analytical Inc., Silver Spring, MD).

Fusarium Head Blight and Deoxynivalenol Evaluations

The 40 barley germplasm lines also were grown in FHB-epidemic nurseries at Fargo and Langdon in 1997 and at Fargo, Langdon, and Osnabrock in 1998. The soil types at each location are the same as previously described. Experimental units consisted of one 1-m row. Germplasm lines were assigned to experimental units using a RCBD, and two replicates of each line were included at each location. Germplasm lines were inoculated four times with F. graminearum, beginning one week before heading, and once a week for four consecutive weeks using the grain-spawn method described by Urrea et al. (2002).

Disease ratings were recorded at the soft dough stage (Zadoks 85) of development. Fifteen spikes within each row were harvested at random, and the number of infected kernels per spike was counted. The percentage of FHB severity was calculated by dividing the total number of infected kernels by the total of kernels and multiplying by 100. Deoxynivalenol content ([micro]g [g.sup.-1]) was determined using the methodology described by Schwarz et al. (1995). The threshold for DON detection by the method we used was 0.5 [micro]g [g.sup.-1].

Foliar Disease Evaluations

The 40 germplasm lines were also grown in the greenhouse during fall 1997. Their responses to the wheat stem rust pathogen (Puccinia graminis Pers.: Pers. f. sp. tritici), leaf rust pathogen (Puccinia hordei G. Otth), net blotch pathogen (Pyrenophora teres Drechs.), spot blotch pathogen [Cochliobolus sativus (Ito & Kuribayashi) Drechs. ex Dastur], and the powdery mildew pathogen (Blumeria graminis DC. f. sp. hordei Em. Marchal) were determined. Three to five seeds of each germplasm line were sown in 3.8 by 21 cm Ray Leach Conetainers (Stuewe & Sons, Corvallis, OR) with #1 Sunshine mix (3:1 peat moss/perlite) (Sun Gro, Bellevue, WA). Osmocote (14-14-14) (Scotts, Marysville, OH) was added to each cone at a rate of 1 g [cone.sup.-1]. Additional water-soluble fertilizer was added as needed. Initial growing conditions in the greenhouse were 22 [+ or -] 2[degrees]C with supplemental lighting (430-W Agrosun bulbs, Hydrofarm Inc., Petaluma, CA) for 14 h [day.sup.-1]. After inoculation with the respective pathogen at the first leaf stage for wheat stem rust and leaf rust and the second leaf stage for spot and net blotch, plants were placed in inoculation chambers at 20[degrees]C with a relative humidity near 100% and 16 h in the dark. Afterward, chamber doors were opened, and plants were allowed to dry for 4 h. Then, plants were returned to the greenhouse at 23 [+ or -] 2[degrees]C and 16-h photoperiod. For powdery mildew, plants were inoculated at the first leaf stage and returned back to the greenhouse as previously described.

A 0.7 mL rust suspension (3.5 mg urediniospores and 0.65 mL lightweight mineral oil) of P. graminis f. sp. tritici (pathotypes Pgt-MCC and Pgt-QCC) or P. hordei (race 8) was applied using a rust inoculator pressurized by an air pump (20 kPa). Inocula of P. teres (isolate NB89-19) and C. sativus (isolate SB85F) were applied to plants using an atomizer pressurized (Model 15, DeVilbiss Inc., Somerset, PA) by an air pump set at 60 kPa. One milliliter of the conidial suspension (5-8 conidia [mL.sup.-1]) was applied per plant. Inoculum of B. graminis f. sp. hordei was applied by shaking heavily infected barley plants over the test entries. Disease ratings were assigned 1 wk after inoculation with each pathogen.

Infection types of wheat stem rust (Miller and Lambert, 1955) and leaf rust (Levine and Cherewick, 1952) were scored using a 0 to 4 rating scale, where a rating of 0 to 2 was indicative of low compatibility and 3 to 4 of high compatibility. Infection response caused by net blotch was scored using a 1 to 9 rating scale, where a rating of 1 to 4 was indicative of low compatibility, 5 was an intermediate compatibility, and 6 to 9 was indicative of high compatibility (Tekaus, 1985). Infection response caused by spot blotch was evaluated using a 1 to 9 rating scale, where a rating of 1 to 3 was indicative of a low compatibility, 4 to 5 was indicative of an intermediate compatibility, and 6 to 9 was indicative of high host-parasite compatibility (Fetch and Steffenson, 1999). Infection response caused by powdery mildew was evaluated using a 0 to 4 rating scale, where a rating of 0 to 2 was indicative of low compatibility and 3 to 4 was indicative of high compatibility (Torp et al., 1978).

Malt Quality Evaluations

In the experiments performed at Fargo in 1998 and Osnabrock in 2000, grain samples were collected from each experimental unit for malting in the Barley Quality Laboratory in the Department of Plant Sciences, North Dakota State University. Samples were malted using the methods described in Karababa et al. (1993). Data collected for each malt sample were moisture (g [kg.sup.-1]), diastatic power (DP) ([degrees]ASBC), [alpha]-amylase activity (20[degrees]C DU), wort viscosity (cP), wort protein (g [kg.sup.-1]), fine- and coarse-grind extract (g [kg.sup.-1]), and malt [beta]-glucan content (g [kg.sup.-1]). Moisture content was determined by heating 10-g samples of ground malt in a semi-automatic Brabender moisture tester (Karababa et al., 1993) at 130[degrees]C for 30 min. Diastatic power and [alpha]-amylase activity of malt samples were determined as described by Technican Industrial Method No. 424-76A (Bran and Luebbee, Inc., Tarrytown, NY) (Banasik, 1971). Wort viscosity at 20[degrees]C was determined according to ASBC wort method 13 (ASBC, 1992). Wort protein was determined according to the UV spectroscopic method of Pyler (Pyler, 1981). Fine- and coarse-malt extract were determined using a modification of ASBC Malt Method 4 (ASBC, 1992). In the modification, 20 g of malt was used rather than the 50 g described in the official method. Malt [beta]-glucan content was determined according to the enzymatic method of McCleary and Nurthen (1986).

Statistical Analyses

Combined analyses of variance across environments were done using the PROC GLM procedure of SAS (Cary, NC) for the agronomic, FHB severity, DON accumulation, and malt quality data. In the combined analyses, environments were considered a random effect and germplasm lines a fixed effect. Thus, the environment x germplasm line mean square was used as the denominator of the F-test for the germplasm line source of variation, and the experimental error was used as the denominator of the F-test for the environment x germplasm line source of variation. F-tests were considered significant at P [less than or equal to] 0.05. Mean separation was done using an F-protected LSD at P [less than or equal to] 0.05.

RESULTS AND DISCUSSION

The 35 germplasm lines with partial FHB resistance evaluated in this study represent all the germplasm lines available to the barley improvement projects at North Dakota State University in 1996. The environment x germplasm line interaction was nonsignificant for all agronomic, malt quality, and foliar disease traits but plump kernels. Upon further investigation, the significance of this interaction was due to differences in magnitude between means from the different environments and not due to a "true" interaction. Hence, all discussion for the traits evaluated is based on means averaged across environments.

Based on the analysis of FHB severity and DON data (data not presented), 22 of the 35 germplasm lines previously reported to have FHB resistance had disease severities that were unacceptable for breeding purposes. Germplasm lines with mean FHB severity greater than 12.5% were deemed susceptible. The two-rowed cultivars judged susceptible were Maja, Balder, Kombainesis, Primus II, Isaria, Francks Hohenloher, Horny Peseky, Gobernadora, Shyri, Aleli, Fuji Nijo, Daisen Gold, Zao Shu 3, Mimai 114, and Misato Golden. The experimental lines F102-61, F103-53, F101-78, and F103-105 also were deemed susceptible to FHB. The partial resistant six-rowed germplasm lines judged susceptible were Glenn, Hazen, and ND15967. To address the objectives of this study, discussions are limited to comparisons of two-rowed barley germplasm lines with partial FHB-resistance to Conlon and Logan, and six-rowed barley germplasm lines with partial FHB-resistance to Morex, Stander, and Foster.

Two-rowed Barley Comparisons

The two-rowed barley germplasm lines Xiao Shan Er Leng Da Mai, Zhedar 1, Dai Shan Er Leng Da Mai, Messidor, CIho 4196, Svanhals, Xiao Shan Ci Mang Er Leng Da Mai, Zhedar 2, Kyoto Nakate, CIho 7595, and Imperial were partially resistant to FHB (Table 2). Some of these germplasm lines actually are reselections from resistant germplasm lines. For example, Kyoto Nakate and Messidor are selections from Svanhals (Table 1). Kyoto Nakate originates from Japan and Messidor from France. FHB severity of this group of germplasm lines ranged from 4.8 to 9.6%, and DON concentration ranged from 2.3 to 6.0 [micro]g [g.sup.-1] (Table 2). The FHB severities and DON concentrations of the germplasm lines within this class were not significantly different, but they were significantly lower than the North Dakota cultivars Logan and Conlon. Logan and Conlon had FHB severities and DON concentrations of 24.9% and 12.8 [micro]g [g.sup.-1] and 27.1% and 8.0 [micro]g [g.sup.-1], respectively.

Days to heading of the partially resistant germplasm lines on average were 8.4 and 11.9 d later than Logan and Conlon, respectively (Table 2). The significantly later maturity of the partially resistant germplasm lines as compared to the checks would be undesirable to growers. Growers prefer the relatively early maturity of barley because it allows them to harvest it before their other crops are mature. CIho 7595 was similar in height to Logan and taller than Conlon. The remaining partially resistant germplasm lines were all taller than Logan and Conlon (Table 2). Short plants often resist lodging better than tall plants, and plants that lodge are more likely to have greater levels of FHB and other foliar diseases. The partially resistant germplasm lines lodged more than Conlon and Logan (Table 2). There was no difference in resistance to lodging between Conlon and Logan. The lower FHB severity in the lodged resistant germplasm lines shows the importance of having genetic resistance. Conlon and Logan yielded significantly greater than all partially FHB-resistant germplasm lines (Table 2). Test weight of these germplasm lines was generally less than that of Conlon and Logan. Only Imperial and PI161970 had test weight similar to those of the checks.

All FHB-resistant two-rowed germplasm lines were susceptible to leaf rust, and most were susceptible to wheat stem rust, net blotch, spot blotch, and powdery mildew (Table 3). Messidor was resistant to pathotype Pgt-MCC of wheat stem rust and CIho 7595 was resistant to net blotch, spot blotch, and powdery mildew.

None of the partially FHB-resistant two-rowed germplasm lines had acceptable malt quality for the traits measured (Table 4). In general, kernel plumpness, grain protein concentration, and malt extract were the traits most severely impacted in the resistant germplasm lines. These germplasm lines had adequate levels of wort protein, diastatic power, and [alpha]-amylase activity

Six-rowed Barley Comparisons

Six-rowed barley germplasm lines CIho 16128 and Chevron were partially resistant to FHB (Table 5). CIho 16128 is a Chevron-derived germplasm line developed in the United States, and Chevron originates from Switzerland (Table 1). FHB severity of the two partially resistant germplasm lines ranged from 9.1 to 12.5%, and no DON was detected. Foster, Morex, and Stander had FHB severities and DON concentrations of 34.6% and 32.2 [micro]g [g.sup.-1], 39.9% and 32.2 [micro]g [g.sup.-1], and 41.1% and 28.9 [micro]g [g.sup.-1], respectively (Table 5). The FHB severities and DON concentrations of the two partially resistant germplasm lines were not significantly different, and they had less FHB than Foster, Morex, and Stander.

All partially FHB-resistant six-rowed barley germplasm lines headed later and were taller than the six-rowed checks (Table 5). On average, the partially resistant germplasm lines headed 6.1 d later and were 15.3 cm taller than the susceptible checks. Chevron and CIho 16128 were more susceptible to lodging and yielded less than the susceptible checks; however, test weight of the partially resistant germplasm lines and susceptible checks was similar.

All partially FHB-resistant six-rowed germplasm lines were susceptible to leaf rust, pathotype Pgt-QCC of wheat stem rust, spot blotch, net blotch, and powdery mildew (Table 6). Chevron and CIho 16128 were resistant to wheat stem rust pathotype Pgt-MCC. None of the partially FHB resistant germplasm lines had acceptable malt quality for all traits. Chevron and CIho 16128 had inadequate levels of kernel plumpness, malt extract, and a-amylase activity; excessive grain protein; and adequate levels of wort protein and diastatic power (Table 7).

Choice of Parents to Use for Introducing Fusarium Head Blight Resistance

Until we know if the partially resistant germplasm lines have different genes for FHB resistance, it would be difficult to recommend one germplasm line over another as a source of FHB resistance because of the deficiencies they have in agronomic traits, malt quality, and foliar disease resistance. Learning more about the genetics of FHB resistance in each of the partially resistant germplasm lines would take many years; however, a more timely method of determining if the germplasm lines may have similar FHB-resistance genes would be to determine via molecular markers the genetic diversity between the lines. The genetic diversity between germplasm lines that may have similar FHB-resistance genes would likely be less than that of lines with different genes.

Urrea (2000) reported on the genetic relationships among the germplasm lines evaluated in this report using cluster analysis of genetic distance based on RAPD marker data. He found that all of the partially resistant two-rowed germplasm lines identified in this report appeared in the same cluster, except Imperial and CIho 7595. These two germplasm lines appeared together in a different cluster and had the highest FHB severity and DON accumulation of the partially resistant two-rowed germplasm lines (Table 2). Urrea (2000) found that Zhedar 1, Zhedar 2, Dai Shan Er Leng, and Svanhals were very similar based on his genetic diversity evaluation. Finally, he found the six-rowed germplasm lines Chevron and CIho 16198 were genetically similar and appeared in a different cluster than the two-rowed germplasm lines. The genetic similarity between Chevron and CIho 16198 is not unexpected since the pedigree of CIho 16198 is 'Atsel'/ Chevron. Research is continuing on the determination of the genetic diversity among the resistant germplasm lines described in this study and additional resistant accessions identified by Scholz et al. (1999) using SSR and RFLP markers.

CONCLUSIONS

In general, germplasm lines with partial FHB resistance were taller, headed and matured later, and yielded less than cultivars adapted to the northern Great Plains of the United States. The partially resistant germplasm lines also tended to have unacceptable grain protein concentration, kernel plumpness, malt extract, and are susceptible to most other fungal pathogens that attack barley in this region. Fusarium head blight severity of the most resistant two-rowed germplasm lines was generally lower than that of the partially resistant six-rowed germplasm lines; however, DON content of these same germplasm lines was similar.

Improvements in malt quality, agronomic traits, and foliar disease resistance of germplasm lines derived from crosses to the accessions with partial FHB resistance have been made; yet, further improvements are needed before FHB-resistant cultivars will be acceptable to pro ducers and the malting and brewing industry. The germplasm lines discussed in this report that are progeny from crosses to germplasm lines with partial FHB resistance have gone through at least two cycles of breeding. Our experience in working with unadapted germplasm tells us that at least four cycles of breeding will be necessary to develop FHB-resistant cultivars that are acceptable to growers and the malting and brewing industries.
Table 1. Barley germplasm lines evaluated for resistance to
F. graminearum in North Dakota trials from 1997 to 2000.

                                  PI
Germplasm line                  number    Source

Aleli ([dagger])                  --      National Small Grains
                                            Collection
Balder ([dagger])               181149    National Small Grains
                                            Collection
CIho 4196 ([dagger])            64275     K. Takeda, 1995
Conlon ([double dagger])        597789    J. Franckowiak, 1997
Daisen Gold ([dagger])            --      B. Steffenson, 1997
Dai Shan Er Leng                  --      B. Steffenson, 1997
  Da Mai ([dagger])
Xiao Shan Ci Mang                 --      B. Steffenson, 1997
  Er Leng Da Mai ([dagger])
F101-78                           --      R. Horsley, 1997
F102-61                           --      R. Horsley, 1997
F103-53                           --      R. Horsley, 1997
F103-61                           --      R. Horsley, 1997
F103-105                          --      R. Horsley, 1997
Francks Hohenloher ([dagger])   467513    National Small Grains
                                            Collection
Fuji Nijo ([dagger])            383928    K. Takeda, 1995
Gobernadora ([dagger])            --      J. Franckowiak, 1997
Horny Peseky ([dagger])           --      K. Takeda, 1995
Imperial ([dagger])             61340     National Small Grains
                                            Collection
Isaria ([dagger])               321800    National Small Grains
                                            Collection
Kombainesis ([dagger])            --      K. Takeda, 1995
Kyoto Nakate ([dagger])           --      K. Takeda, 1995
Logan ([double dagger])           --      J. Franckowiak, 1997
Maja ([dagger])                 184884    K. Takeda, 1995
Messidor ([dagger])             174473    National Small Grains
                                            Collection
Mimai 114 ([dagger])            584962    National Small Grains
                                            Collection
Misato Golden ([dagger])          --      B. Steffenson, 1997
CIho 7595 ([dagger])            161970    National Small Grains
                                            Collection
Xiao Shan Er Leng               566203    National Small Grains
  Da Mai ([dagger])                         Collection
Primus II ([dagger])              --      National Small Grains
                                            Collection
Svanhals ([dagger])              5474     K. Takeda, 1995
Shyri ([dagger])                  --      J. Franckowiak, 1997
Zao Shu 3 ([dagger])            466772    National Small Grains
                                            Collection
Zhedar 1 ([dagger])               --      B. Steffenson, 1997
Zhedar 2 ([dagger])               --      B. Steffenson, 1997
Chevron                          38061    R. Horsley, 1997
CIho 16128 ([dagger])             --      B. Steffenson, 1997
Glenn                             --      R. Horsley, 1997
Hazen                           483238    R. Horsley, 1997
ND15967                           --      R. Horsley, 1997
Foster ([double dagger])        592758    R. Horsley, 1997
Morex ([double dagger])           --      R. Horsley, 1997
Stander ([double dagger])       564743    R. Horsley, 1997

Germplasm line                      Pedigree information

Aleli ([dagger])                Unknown
Balder ([dagger])               Gull/Swedish landrace//Maja
CIho 4196 ([dagger])            Unknown
Conlon ([double dagger])        Bowman*2/Birgittamatt//ND10232
Daisen Gold ([dagger])          Unknown
Dai Shan Er Leng                Unknown
  Da Mai ([dagger])
Xiao Shan Ci Mang               Unknown
  Er Leng Da Mai ([dagger])
F101-78                         Gobernadora/Foster//ND9712
F102-61                         Zhedar 1/Foster//ND9712
F103-53                         Zhedar 2/Foster//ND9712
F103-61                         Zhedar 2/Foster//ND9712
F103-105                        Zhedar 2/Foster//ND9712
Francks Hohenloher ([dagger])   Imperia/Bethge II
Fuji Nijo ([dagger])            Plumage Archer/Nirasaki Wase 1
Gobernadora ([dagger])          OC640/Mari//Pioneer/3/Maris Concord
Horny Peseky ([dagger])         Unknown
Imperial ([dagger])             Mutant Imperial, CIho3197
Isaria ([dagger])               Bavaria/Danubia
Kombainesis ([dagger])          Unknown
Kyoto Nakate ([dagger])         Selection from Svanhals
Logan ([double dagger])         ND7085/ND4994-15//ND7556
Maja ([dagger])                 Blinder/Gull
Messidor ([dagger])             Selection from Svanhals
Mimai 114 ([dagger])            Unknown
Misato Golden ([dagger])        Unknown
CIho 7595 ([dagger])            Bavaria/Danubia
Xiao Shan Er Leng               Unknown
  Da Mai ([dagger])
Primus II ([dagger])            Bulked from Primus
Svanhals ([dagger])             Selection from Besterhon Diamant
Shyri ([dagger])                Lignee 640/Kober//Feran 78
Zao Shu 3 ([dagger])            Unknown
Zhedar 1 ([dagger])             Unknown
Zhedar 2 ([dagger])             Unknown
Chevron                         Unknown
CIho 16128 ([dagger])           Atsel/Chevron
Glenn                           Br5755-3/Trophy//NDB138
Hazen                           Glenn/4/Nordic//Dickson/Frophy/3/Azure
ND15967                         Stander/3/ND9712//Foster/Pl 452421
Foster ([double dagger])        Robust/3/ND5570//Glenn/Karl
Morex ([double dagger])         Cree/Bonanza
Stander ([double dagger])       Excel//Robust/Bumper

                                                 Row
Germplasm line                    Origin       number

Aleli ([dagger])                Switzerland       2
Balder ([dagger])               Sweden            2
CIho 4196 ([dagger])            China             2
Conlon ([double dagger])        ND, USA           2
Daisen Gold ([dagger])          Japan             2
Dai Shan Er Leng                China             2
  Da Mai ([dagger])
Xiao Shan Ci Mang               China             2
  Er Leng Da Mai ([dagger])
F101-78                         ND, USA           2
F102-61                         ND, USA           2
F103-53                         ND, USA           2
F103-61                         ND, USA           2
F103-105                        ND, USA           2
Francks Hohenloher ([dagger])   Germany           2
Fuji Nijo ([dagger])            Kirin, Japan      2
Gobernadora ([dagger])          Mexico            2
Horny Peseky ([dagger])         China             2
Imperial ([dagger])             Japan             2
Isaria ([dagger])               Germany           2
Kombainesis ([dagger])          Unknown           2
Kyoto Nakate ([dagger])         Japan             2
Logan ([double dagger])         ND, USA           2
Maja ([dagger])                 Denmark           2
Messidor ([dagger])             France            2
Mimai 114 ([dagger])            China             2
Misato Golden ([dagger])        Japan             2
CIho 7595 ([dagger])            Germany           2
Xiao Shan Er Leng               China             2
  Da Mai ([dagger])
Primus II ([dagger])            SD, USA           2
Svanhals ([dagger])             Sweden            2
Shyri ([dagger])                Ecuador           2
Zao Shu 3 ([dagger])            China             2
Zhedar 1 ([dagger])             China             2
Zhedar 2 ([dagger])             China             2
Chevron                         Switzerland       6
CIho 16128 ([dagger])           MD, USA           6
Glenn                           ND, USA           6
Hazen                           ND, USA           6
ND15967                         ND, USA           6
Foster ([double dagger])        ND, USA           6
Morex ([double dagger])         MN, USA           6
Stander ([double dagger])       MN, USA           6

([dagger]) Barley germplasm lines reported to possess partial
resistance to Fusarium head blight (Steffenson, personal
communication, 1996; Prom et al., 1996).

([double dagger]) Barley cultivars adapted for production in the
northern Great Plains of the United States.

Table 2. Mean Fusarium head blight (FHB) severity, ([dagger])
deoxynivalenol (DON) concentration, ([double dagger]) and agronomic
performance ([section]) of selected two-rowed barley germplasm lines
grown in North Dakota trials from 1997 to 2000.

                                      FHB                     Days to
Germplasm line                     severity       DON         heading

                                                [micro]      days after
                                       %       [g.sup.-1]      31 May

Xiao Shan Er Leng Da Mai              4.8         3.8           33.1
Zhedar 1                              5.9         2.9           33.3
Dai Shan Er Leng Da Mai               6.1         2.5           32.9
Messidor                              6.2         2.3           33.5
CIho 4196                             6.6         2.7           33.6
Svanhals                              6.7         2.9           32.7
Xiao Shan Ci Mang Er Leng Da Mai      6.9         3.8           33.3
Zhedar 2                              7.4         5.0           33.6
Kyoto Nakate                          7.9         4.3           32.2
CIho 7595                             9.6         4.9           33.2
Imperial                              9.6         6.0           33.3
Logan                                24.9        12.8           27.8
Conlon                               27.1         8.0           24.3
LSD (0.05)                            5.1         5.4            1.7

                                     Plant
Germplasm line                      height    Lodging       Yield

                                      cm       (1-9)     Mg [ha.sup.-1]

Xiao Shan Er Leng Da Mai            100.0       6.8           2.9
Zhedar 1                             96.9       6.6           2.9
Dai Shan Er Leng Da Mai              99.3       7.2           2.6
Messidor                             98.6       6.7           3.0
CIho 4196                            95.9       7.4           2.6
Svanhals                             97.7       6.9           3.0
Xiao Shan Ci Mang Er Leng Da Mai     96.2       7.4           2.7
Zhedar 2                             99.5       5.9           3.1
Kyoto Nakate                         98.5       6.4           2.7
CIho 7595                            90.8       5.5           3.1
Imperial                             97.2       4.4           3.3
Logan                                83.5       1.7           4.6
Conlon                               85.6       2.6           4.5
LSD (0.05)                            5.6       1.1           0.7

                                        Test
Germplasm line                         weight

                                   kg [hL.sup.-1]

Xiao Shan Er Leng Da Mai                60.6
Zhedar 1                                62.0
Dai Shan Er Leng Da Mai                 60.6
Messidor                                59.6
CIho 4196                               60.7
Svanhals                                59.2
Xiao Shan Ci Mang Er Leng Da Mai        61.8
Zhedar 2                                61.1
Kyoto Nakate                            60.5
CIho 7595                               66.1
Imperial                                64.7
Logan                                   65.9
Conlon                                  66.4
LSD (0.05)                               2.2

([dagger]) Fusarium head blight data were collected on germplasm lines
grown in FHB nurseries in 1997 and 1998 at Fargo, Langdon, and
Osnabruck, ND.

([double dagger]) Deoxynivalenol data were collected on grain grown in
FHB nurseries in 1997 at Fargo, Langdon, and Osnabrock, ND; and in 1998
at Langdon and Osnabrock, ND.

([section]) Agronomic performance of selected germplasm lines grown in
yield trials in 1998 at Fargo, ND; in 1999 at Osnabrock, ND; and in
2000 at Fargo, Langdon, and Osnabrock, ND. Germplasm lines were not
inoculated with F. graminearum and were protected from FHB and foliar
diseases using multiple applications of benomyl.

Table 3. Greenhouse seedling response of selected barley germplasm
lines to different foliar pathogens. Germplasm lines are
ordered from most resistant to most susceptible to Fusarium
head blight.

                       Wheat stem
                     rust ([dagger])

                          MCC              QCC            Leaf rust
Germplasm line        ([section])     ([paragraph])   race 8 ([dagger])

Xiao Shan Er Leng     2,3              3,2,3          3,[3.sup.-]
  Da Mai
Zhedar1               2,[3.sup.-]      3,[sup.-]2     3
Dai Shan Er Leng     [3.sup.-],2,3     2,[3.sup.-]    3,[3.sup.+]
  Da Mai
Messidor              2,1,[3.sup.-]    2,1           [3.sup.-],3
CIho 4196            [3.sup.-],2       2             [3.sup.-]
Svanhals             [3.sup.-],2      [3.sup.-],2    [3.sup.-],3
Man Shan Ci Mang      3                2,[3.sup.-]   [3.sup.-],2
  Er Leng Da Mai
Zhedar 2              3,[3.sup.-]     [3.sup.-]2,3   [3.sup.-],2
Kyoto Nakate         [3.sup.-],2,1    [3.sup.-],2     3,[3.sup.+]
CIho 7595             3                3,[sup.-]2    [3.sup.-],2
Imperial              3,[3.sup.-]     [3.sup.-],2     3,[3.sup.+]
Logan                 0,1             2,[3.sup.-]    [3.sup.-],2
Conlon                1,0,[2.sup.-]   [3.sup.-],2    [3.sup.-]

                          Spot             Net
                         blotch          blotch
                        ([double        ([double           Powdery
Germplasm line          dagger])        dagger])      mildew ([dagger])

Xiao Shan Er Leng          8,7             8,9                3
  Da Mai
Zhedar1                    7,6             8,9                3
Dai Shan Er Leng           5,6             8,9               34
  Da Mai
Messidor                   7,8             7,6               34
CIho 4196                  7,6             8,9               34
Svanhals                   6,5             8,9               34
Man Shan Ci Mang           7,6             8,7                3
  Er Leng Da Mai
Zhedar 2                   7,8             7,8                3
Kyoto Nakate               6,7             8,7                3
CIho 7595                  3,2             3,2               12
Imperial                   5,4             8,9               34
Logan                      3,4             2,3                1
Conlon                     5,4             2,3                0

([dagger]) Germplasm lines exhibiting infection types 0 to 2 were
classified as resistant, and those exhibiting infection types 3 and
4 were classified as susceptible.

([double dagger]) Germplasm lines exhibiting infection types 1 to 3
were classified as resistant, 4 and 5 as intermediate, and 6 to 9 as
susceptible.

([section]) Pathotype Pgt-MCC.

([paragraph]) Pathotype Pgt-QCC.

Table 4. Malt quality of selected two-rowed barley germplasm lines
grown at Fargo, ND, in 1998 and Osnabrock, ND, in 2000.

Germplasm           Plump       Grain       Malt         Wort
line ([dagger])    kernels     protein     extract      protein

                      %                 g [kg.sup.-1]

Xiao Shan Er        59.0         150         759          510
  Ling Da Mai
Zhedar 1            63.0         151         757          510
Dai Shan Er         62.1         152         763          500
  Leng Da Mai
Messidor            60.2         138         769          490
CIho 4196           55.4         149         766          490
Svanhals            57.4         150         761          510
Mao Shan Ci         56.9         147         763          500
  Mang Er
  Leng Da Mai
Zhedar 2            59.6         150         757          510
CIho 7595           62.4         147         771          460
Kyoto Nakate        63.2         152         755          510
Imperial            74.5         139         772          450
Logan               79.1         126         803          480
Conlon              85.5         125         812          520
LSD (0.05)          13.3          13.4        16.2         64

Germplasm                Diastatic             [alpha]-Amylase
line ([dagger])            power                  activity

                     [degrees] ASBC
                    ([double dagger])   20[degrees] DU ([section])

Xiao Shan Er               107                     50.3
  Ling Da Mai
Zhedar 1                   113                     53.1
Dai Shan Er                109                     48.2
  Leng Da Mai
Messidor                   106                     54.5
CIho 4196                  113                     49.1
Svanhals                   109                     51.8
Mao Shan Ci                111                     50.1
  Mang Er
  Leng Da Mai
Zhedar 2                   114                     49.4
CIho 7595                  111                     50.5
Kyoto Nakate               116                     50.3
Imperial                   122                     48.0
Logan                      121                     53.0
Conlon                     104                     56.0
LSD (0.05)                  16.2                    8.14

([dagger]) Germplasm lines are ranked from most resistant to most
susceptible to Fusarium head blight.

([double dagger]) [degrees]ASBC = Degrees American Society of Brewing
Chemist.

([section]) 20[degrees]DU = Dextrinizing units at 20[degrees]DU.

Table 5. Mean Fusarium head blight (FHB) severity, ([dagger])
deoxynivalenol (DON) concentration, ([double dagger]) and agronomic
performance ([section]) of selected six-rowed barley germplasm lines
grown in North Dakota from 1997 to 2000.

Germplasm         FHB                     Days to       Plant
line           severity        DON        heading      height

                            [micro]g    days after
                   %       [g.sup.-1]     31 May         cm

CIho 16128        9.1          0.0         32.9         108.5
Chevron          12.5          0.0         33.7         103.3
Foster           34.6         17.5         27.3          88.8
Morex            39.9         32.2         26.3          90.9
Stander          41.1         28.9         28.0          88.0
LSD (0.05)        5.9         23.7          1.4           7.2

Germplasm                                       Test
line            Lodging        Yield           weight

                 (1-9)     Mg [ha.sup.-1]   kg [hL.sup.-1]

CIho 16128        6.1           2.8              58.5
Chevron           6.3           3.0              60.1
Foster            3.3           4.4              55.8
Morex             3.3           4.2              55.9
Stander           1.9           4.6              58.1
LSD (0.05)        1.4           0.9               2.7

([dagger]) Fusarium head blight data were collected on germplasm lines
grown in FHB nurseries in 1997 and 1998 at Fargo, Langdon, and
Osnabrock, ND.

([double dagger]) Deoxynivalenol data were collected on grain grown in
FHB nurseries in 1997 at Fargo, Langdon, and Osnabrock, ND; and in 1998
at Langdon and Osnabrock, ND.

([section]) Agronomic performance of selected germplasm lines grown in
yield trials in 1998 at Fargo, ND; in 1999 at Osnabrock, ND; and in
2000 at Fargo, Langdon, and Osnabrock, ND. Germplasm lines were not
inoculated with F. graminearum and were protected from FHB and foliar
diseases using multiple applications of benomyl.

Table 6. Greenhouse seedling response of selected barley germplasm
lines to different foliar pathogens. Germplasm lines are ordered
from most resistant to most susceptible to Fusarium head blight.

                       Wheat stem
                     rush ([dagger])
Germplasm                                                Leaf rust
line          MCC ([section])   QCC ([paragraph])    race 8 ([dagger])

CIho 16128    1                 3                   [3.sup.-],3
Chevron       1                 3                   [3.sup.-],3
Foster        0,[1.sup.-]       [3.sup.-],2         [3.sup.-],2
Morex         0,[1.sup.-]       [3.sup.-],2         [3.sup.-],3
Slander       1,0               2,[3.sup.-]         [3.sup.-]

                   Spot                Net
Germplasm     blotch ([double    blotch ([double        Powdery
line             dagger])           dagger])        mildew ([dagger])

CIho 16128          5,6                5,6              3 ([section])
Chevron             6,7                8,7              3
Foster              4,3                7,8              3
Morex               5,6                7,8              3
Slander             4,3                7,8              3

([dagger]) Germplasm lines exhibiting infection types 0 to 2 were
classified as resistant, and those exhibiting infection types 3 and 4
were classified as susceptible.

([double dagger]) Germplasm lines exhibiting infection types 1 to 3
were classified as resistant, 4 and 5 as intermediate, and 6 to 9 as
susceptible.

([section]) Pathotype Pgt-MCC.

([paragraph]) Pathotype Pgt-QCC.

Table 7. Malt quality of selected six-rowed barley germplasm
lines ([dagger]) grown at Fargo, ND, in 1998 and Osnabrock, ND,
in 2000.

Germplasm          Plump      Grain      Malt       Wort
line ([dagger])   kernels    protein    extract    protein

                    %                 g [kg.sup.-1]

CIho 16128         14.6       164        733        530
Chevron            15.5       167        725        520
Foster             76.4       122        796        560
Morex              64.8       132        793        560
Stander            79.5       130        799        590
LSD (0.05)          7.8        11.4       19.1       81

Germplasm              Diastatic           [alpha]-Amylase
line ([dagger])          power                 activity

                    [degrees]ASBC     20[degrees] DU ([section])
                  ([double dagger])

CIho 16128               174                    53.0
Chevron                  171                    50.9
Foster                   151                    63.1
Morex                    143                    68.1
Stander                  135                    63.7
LSD (0.05)                35.2                  13.5

([dagger]) Germplasm lines are ranked from most resistant to most
susceptible to Fusarium head blight.

([double dagegr] [degress]ASBC Degrees American Society of Brewing
Chemist.

([section]) 20[degrees]DU = Dextrinizing units at 20[degrees]DU.


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Carlos A. Urrea, Richard D. Horsley, * Brian J. Steffenson, and Paul B. Schwarz

C.A. Urrea, CIMMYT, Mexico; R.D. Horsley and P.B. Schwarz, Dep. of Plant Sciences, North Dakota State Univ., Fargo, ND 58105-5051; B.J. Steffenson, Dep. of Plant Pathology, Univ. of Minnesota, 495 Borlaug Hall, 1991 Upper Buford Circle, St. Paul, MN, 55108. Received 24 Nov. 2003. * Corresponding author (richard.horsley@ndsu. nodak.edu).
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Author:Urrea, Carlos A.; Horsley, Richard D.; Steffenson, Brian J.; Schwarz, Paul B.
Publication:Crop Science
Date:Jul 1, 2005
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