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a Dep. of Plant Sciences, North Dakota State Univ., P.O. Box 5051, Fargo, ND 58105
b Dep. of Plant Pathology, North Dakota State Univ., P.O. Box 5051, Fargo, ND 58105
c USDA-ARS, Fargo, ND
* Corresponding author (mohamed.mergoum{at}ndsu.nodak.edu).
ABSTRACT
Faller (Reg. No. CV-1026, PI 648350) hard red spring wheat (HRSW) (Triticum aestivum L.) was developed at North Dakota State University (NDSU) and released by the North Dakota Agricultural Experiment Station (NDAES). Faller was derived from the ND2857/ND2814 cross made at NDSU in fall 1997. ND2857 (ND2709/ND688) is a hard red spring experimental line with good resistance to Fusarium head blight (FHB) (caused by Fusarium graminearum Schwabe [telomorph Gibberella zeae (Schwein.) Petch]) originating from ND2709, a line known to possess the Fhb1 quantitative trait locus derived from Sumai3 (PI 481542). Sumai3, a spring wheat from China, is arguably the most widely used source of resistance to FHB in the world. Both ND2709 and ND688 are HRSW experimental lines developed by the NDSU breeding program. ND2814 (Kitt [PI 518818]/Amidon [PI 527682]/3/Grandin [PI 531005]/Stoa S [PI 520297]) is an HRSW line developed by the NDSU HRSW breeding program. Kitt is an HRSW cultivar released in 1975 by the Minnesota Agricultural Experiment Station and the USDA-ARS, while Amidon, Grandin, and Stoa are HRSW cultivars released by NDAES in 1988, 1989, 1984, respectively. Faller was produced from a bulk of one purified F4:5 plot selected in 2001 at Christchurch, New Zealand. Faller was released because it combines very high yield with good end-use quality and resistance to FHB and leaf diseases.
Abbreviations: AYT, advanced yield trial EYT, elite yield trial FHB, Fusarium head blight HRSW, hard red spring wheat IYT, intermediary yield trial NDAES, North Dakota Agricultural Experiment Station ND HRSW-VT, North Dakota Hard Red Spring Wheat Variety Trials NDSU, North Dakota State University NZ, New Zealand PYT, preliminary yield trial SNB, Septoria nodorum blotch STB, Septoria tritici blotch URN, Uniform Regional Nursery
Faller (Reg. No. CV-1026, PI 648350) hard red spring wheat (HRSW) (Triticum aestivum L.) was tested as experimental line ND805, developed at North Dakota State University (NDSU), and released by the North Dakota Agricultural Experiment Station (NDAES). In addition to researchers at NDSU, USDA-ARS researchers at Fargo, ND, contributed by testing Faller for stem rust (caused by Puccinia graminis Pers.:Pers. f. sp. tritici Eriks. & E. Henn) resistance. Faller was released because it combines very high yield potential with good end-use quality and resistance to both Fusarium head blight (FHB) (caused by Fusarium graminearum Schwabe [telomorph Gibberella zeae (Schwein.) Petch]) and leaf diseases. The name Faller was chosen as recognition to the contribution of the late James Faller, a former technician in the HRSW breeding program for almost three decades.
Faller was selected from the ND2857/ND2814 cross made at NDSU in fall 1997. ND2857 (ND2709/ND688) is a hard red spring experimental line with good resistance to FHB originating from ND2709, a line derived from a cross involving Sumai3 (PI 481542). Sumai3, a spring wheat from China, is arguably the most commonly used source of resistance to FHB in the world. Both ND2709 and ND688 are HRSW experimental lines developed by the NDSU breeding program. ND2814 (Kitt [PI 518818]/Amidon [PI 527682]/3/Grandin [PI 531005]/Stoa S [PI 520297]) is a HRSW line developed by the NDSU HRSW breeding program. Kitt is a HRSW cultivar released in 1975 by the Minnesota Agricultural Experiment Station and the USDA-ARS, while Amidon, Grandin and Stoa are HRSW cultivars released by the NDAES in 1988, 1989, 1984, respectively.
Methods
Early Generation Development
Faller was developed using a modified bulk breeding procedure. The F1 seeds from the cross leading to Faller were grown in the field at Prosper, ND, in spring 1998, and the F2 population was grown in the New Zealand (NZ) off-season nursery during winter 1998–1999. From the F2 population, 100 spikes were harvested, threshed individually, and advanced to obtain F3 seed in summer 1999. Subsequently, 10 spikes selected from each selected F3 row were threshed individually and sown in F4 head-row plots in the observation nursery and in hill-plots in the FHB nursery at Prosper, ND, in summer 2000. The FHB nursery was inoculated with the FHB pathogen using the spray-inoculation method (Rudd et al., 2001) and overhead mist irrigation to enhance disease development. Ten spikes from each hill showing less than 10% FHB disease severity (Stack and Frohberg, 1997) were harvested, threshed individually, and advanced as F5 head row plots at the NZ off-season nursery during winter 2000–2001. Selected rows from NZ were harvested in bulk and planted in a F4:6 preliminary yield trial (PYT) in 2001. Selections in the F3, F4, and F5 generations were based on reaction to FHB and leaf diseases, particularly leaf rust (caused by P. triticina Eriks.), and agronomic merits including plant vigor, height, and earliness. In NZ selection was based mainly on visual uniformity, lack of grain shattering, plant height, and lodging resistance.
Line Selection and Evaluation
Faller was evaluated for agronomic and quality traits in the intermediary yield trial (IYT) in 2002, advanced yield trial (AYT) in 2003, and elite yield trial (EYT) in 2004. Subsequently, Faller was tested in the North Dakota HRSW Variety Trials (ND HRSW-VT) in 2005 and 2006. Faller was also tested in the HRSW Uniform Regional Nursery (URN) in 2005 and 2006. All yield trials were arranged in a randomized complete block design with two replicates for IYT and four replicates for AYT and EYT. The experimental unit consisted of seven rows, 3 m long, and 30 cm apart. While IYT and AYTs were grown at four locations (Casselton, Minot, Carrington, and Prosper, ND), EYTs were grown in three additional locations; Hettinger, Langdon, and Williston, ND. The ND HRSW-VT is a statewide trial conducted at seven locations across North Dakota in an RCBD with four replicates. The plot size consisted of eight rows, 10 m long, and 30 cm apart. The URN was conducted during 2005 and 2006 in 37 location-years across the states of North Dakota, Minnesota, South Dakota, Nebraska, Montana, Wyoming, Washington, and Manitoba, Canada. These trials were laid out in an RCBD with three replicates. Depending on location, the experimental unit consisted of six to eight rows, 3 m long, and 30 cm apart.
Faller was tested for its reaction to different races of tan spot [caused by Pyrenophora tritici-repentis (Died.) Drechs], Septoria tritici blotch (STB) [caused by Mycosphaerella graminicola (Fückl) J. Schröt. in Cohn], Septoria nodorum blotch (SNB) [caused by Stagonospora nodorum (Berk.) Castellani & E.G. Germano], leaf rust, stem rust, and FHB in greenhouse and field tests during 2001 to 2006. The SNB, STB, and tan spot diseases are major components of the leaf spotting complex of wheat in North America. A complex of these diseases occurs in nature; hence, managing leaf spot diseases is difficult. Resistant cultivars are the most effective and economical means of controlling leaf spot.
Tan spot of wheat can cause two phenotypically distinct and independent symptoms: tan necrosis and extensive chlorosis (Lamari and Bernier, 1989). Currently, eight races of tan spot have been identified (Lamari et al., 2003). From 2003 to 2006, Faller was tested for resistance to four races of tan spot in six greenhouse trials. The races tested included all the virulent races observed in North America (race 1, the most prevalent tan spot race in North Dakota, and races 2, 3, and 5). The greenhouse trials were arranged as an RCBD with three replicates and three plants per replicate as the experimental unit. The 1 to 5 lesion-type scale developed by Lamari and Bernier (1989) was used, where 1 = resistant with small, dark brown to black spots without any surrounding chlorosis or tan necrosis, and 5 = susceptible with dark brown or black centers that may or may not be distinguishable. Most lesions consisted of coalescing chlorotic or tan necrotic zones.
Faller was also evaluated for reaction to SNB based on a 1 to 5 lesion-type scale developed by Feng et al. (2004), where 1 = pinpoint dark brown lesions without chlorosis, 2 = small lesions with very little necrosis or chlorosis, 3 = chlorotic or necrotic lesions completely surrounded by a chlorotic ring, 4 = lesions completely surrounded by chlorotic zones, some of the lesions coalescing, and 5 = extensive chlorosis and large necrotic lesions. Ratings of 1 to 2 were considered resistant while those with 3 to 5 were classified as susceptible. Faller was also evaluated for STB based on a 0 to 5 scale developed by McCartney et al. (2002), where 0 = immune, 1 = highly resistant, 2 = resistant, 3 = moderately susceptible, 4 = susceptible, and 5 = highly susceptible. Ratings of 0 to 2 were considered resistant while those with 3 to 5 were considered susceptible. Evaluation of Faller for SNB and STB was made on the basis of eight greenhouse experiments arranged in an RCBD with three replicates, and three plants per replicate as the experimental unit.
Screening of Faller for FHB was conducted from 2003 to 2006 in 12 FHB nurseries under both field (eight location-years) and greenhouse (four experiments) conditions. The field FHB nursery was laid out in an RCBD with four replicates and inoculated with the FHB pathogen using the spray-inoculation method described by Rudd et al. (2001) with overhead mist irrigation to enhance disease development. Entries were assigned to a hill plot consisting of eight to ten plants. Similarly, the greenhouse experiments were arranged in a RCBD with three replicates. The entries were assigned to a 0.5 m row plot.
The reaction type of Faller to the prevalent races of leaf and stem rusts was done on the basis of six field tests (RCBD, four replicates, and 1 m row-plot per replicate) and eight greenhouse tests (RCBD, three replicates, and four plants per replicate) from 2003 to 2006. In the greenhouse experiments, Faller was specifically evaluated for resistance to the predominant stem rust pathotypes Pgt-TMLK, -QTHJ, -QFCQ, -RTQQ, -TPMK, -THTS, and -TCMJ, and to leaf rust pathotypes MCDL and THBJ.
Seed Purification and Increase
Faller was first produced from a bulk of purified F5 head-rows selected in 2001 at the NZ nursery. Faller was further purified by selecting one hundred spikes from the quality drill strips (F7) grown at Prosper, ND, in 2004. These spikes were threshed individually and seeded as head-rows at Casselton in 2005. Nonuniform rows were discarded and the remaining rows were bulked and planted in the 2005–2006 winter nursery grown near Yuma, AZ, as the first seed increase of Faller. Further seed increase of Faller was continued by the NDSU Seed-Stock program in summers 2006 and 2007. The purity of Faller was maintained throughout the increase process based on visual elimination of off-type plants (taller, different spike color, and presence or absence of awns).
Statistical Analysis
Data were analyzed using SAS-JMP version 6.0.3 (SAS Institute, Cary, NC). Grain yield and other agronomic data such as grain volume weight from the AYT and HRSW-VT were subjected to analysis of variance across locations within years and a combined analysis across location-years was performed whenever error variances were homogeneous. The analysis of AYT, HRSW-VT, and URN included only entries common to the trials across years. A mixed model with environments and genotypes as fixed effects and replications within environments as random effects was used for within-year analyses. Similarly, across-year analyses were also done according to a mixed model with genotypes and location-year combinations as fixed effects and replications within location-year combinations as random effects. Tukey's honestly significant difference test (
= 0.05) was used to compare the least squares means for the genotype effects.
Characteristics
Agronomic and Botanical Description
Faller is an awned, medium-early maturing, and semidwarf hard spring wheat. It has a lax head type with plant height (85 cm) significantly taller than Alsen (PI 615543; Frohberg et al., 2006) and Reeder (PI 613586) (80 cm), but shorter than Dapps (PI 633862; Mergoum et al., 2005a) (93 cm). Faller's height was, however, similar to Steele-ND (PI 634981; Mergoum et al., 2005b), Howard (PI 642367; Mergoum et al., 2006b), and Parshall (PI 613587) (Table 1
). Similarly, in the URN trials, the height of Faller was 84 cm, comparable to the height of Verde (PI 592561) (79 cm) and 2375 (82 cm), but significantly shorter than Keene (PI 598224 PVP) (96 cm) and Chris (CItr 13751) (100 cm) (Table 2
). Faller heads the same day as Reeder, Howard, Glenn (PI 639273; Mergoum et al., 2006a), and Steele-ND, and significantly later than Alsen, Parshall, and Dapps (Table 1). Compared with Verde, Keene, and Chris, Faller heads on the same day as Verde and Keene and 1 d earlier, but not significantly, than Chris (Table 2). Plant lodging (straw strength) was measured on a scale of 0 to 9 where 0 = completely erect and 9 = totally flat at harvest, Faller was resistant (lodging score = 0.6), similar to Howard, Glenn, and Dapps, with lodging scores of 0.5, but significantly less susceptible than Steele-ND (lodging score = 2.0) (Table 1). Similarly, the lodging score of Faller grown in the URN was 0.9, similar to Verde but significantly less than Keene and Chris (Table 2).
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Faller was observed for 10 crop cycles (F3–F12 generations) from 1999 to 2006 and was observed to be uniform and stable during the last four generations of seed increase (head-row increases and large drill strip increases in 2004, Breeder seed increase in 2005, and Foundation seed increase in 2006). Faller remains stable in its essential and distinctive characteristics when sexually reproduced. Variants are limited to (i) taller plants (5–30 cm) that occur at a frequency of less than 1 in 1000 plants and (ii) awnless plants at a trace frequency of less than 1 in 10,000 plants. The variants in Faller are within commercially acceptable limits for all described traits.
Disease Reactions
Under severe FHB disease pressure, the average disease severity (Stack and Frohberg, 2000) recorded on Faller from the field scab nursery (27%) was significantly lower (P < 0.05) than the susceptible check Reeder (55%) and the very susceptible check 2398 (72%) (Table 3
). In the same trials, the average FHB severities recorded on Alsen, Glenn, Parshall, Steele-ND, Dapps, and ND 2710 (PI 633976, Frohberg et al., 2004) were 26, 22, 38, 31, 42, and 13%, respectively. Alsen was released in 2000 as the first NDSU HRSW cultivar with resistance to FHB and has been widely grown in North Dakota since 2001. Glenn (Mergoum et al., 2006a) was also an NDSU HRSW released in 2005 with FHB resistance and was the leading grown cultivars in North Dakota in 2007. Similarly, Howard and Steele-ND are both NDSU HRSW releases with FHB resistance derived from T. dicoccoides (Mergoum et al., 2006b, 2005b). Under greenhouse conditions (data not shown), average FHB severity of Faller was 34.2%, similar to the scores of Alsen (29%) and Glenn (30%); and significantly lower (P < 0.05) than 89 and 73% registered for the susceptible checks 2398 and Reeder, respectively. The same tests indicate that visually scabby kernels of Faller (21%) was also very low (P < 0.01) compared with the susceptible checks 2398 (74%) and Reeder (60%), but similar to Alsen (24%), Steele-ND (29%), Howard (31%), and higher than the resistant check ND 2710 (8%).
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Grain Yield Performance and Quality Parameters
Based on 27 location-years of testing in the HRSW-VTs and AYTs, grain yield of Faller (4467 kg ha–1) was significantly (P < 0.05) higher than all previously released NDSU cultivars, including Alsen (3763 kg ha–1), Glenn (3743 kg ha–1), Parshall (3607 kg ha–1), Steele-ND (4052 kg ha–1), Reeder (3625 kg ha–1), and Howard (3943 kg ha–1) (Table 1). In 37 location-years of testing in the URN trials conducted in 2005 and 2006, Faller yielded 4014 kg ha–1 compared with 3213, 3743, and 2738 kg ha–1 for Keene, Verde, and Chris, respectively (Table 2). Faller has average thousand kernel weight (29.8 g) compared with 29, 30.2, 28.4, 29.5, 26.9, 26.5, and 26.7 registered for Howard, Glenn, Steele-ND, Dapps, Alsen, Parshall, and Reeder (Table 1). Mean grain volume weight of Faller (757 kg m–3) over 26 location-years in HRSW-VT was similar to Reeder (753 kg m–3) and Dapps (756 kg m–3), but significantly (P < 0.05) lower than Glenn (797 kg m–3) and Howard (778 kg m–3) (Table 1). In the URN, the mean grain volume weight of Faller was 752 kg m–3 compared with 753, 767, 758, 742, and 737 kg m–3 of cultivars Verde, 2375, Keene, Chris, and Marquis, respectively (Table 2). Marquis is an old spring wheat cultivar released in 1911 (Underdahl et al., 2008). Grain protein of Faller (150 g kg–1) was comparable to Howard (15.3 g kg–1), Steele-ND (15.3 g kg–1), and Reeder (155 g kg–1), but lower (P < 0.05) than Glenn (15.7 g kg–1), Dapps (165 g kg–1), Alsen (157 g kg–1), and Parshall (156 g kg–1) (Table 1). Similarly, in the URN trials, Faller's grain protein was 14.1 g kg–1 compared with 14.4, 14.1, 14.4, 15.1, and 14.3 g kg–1 recorded on Verde, 2375, Keene, Chris, and Marquis, respectively (Table 2). Additional quality parameters including falling number, flour extraction, dough and baking parameters for Faller and major HRSW cultivars included in the HRSW-VT grown in ND from 2003 to 2006 are reported in Table 4
. These data show that falling number of Faller (423 s) was not significantly different from those of the most commonly grown HRSW cultivars, including Howard (427 s), Steele-ND (425 s), Alsen (412 s), Parshall (415 s), and Reeder (431 s), and excepting Glenn (401 s) (Table 4). Flour extraction (Table 4) of Faller (711 g kg–1), although higher than all checks, was significantly superior only to Glenn (676 kg–1), Alsen (686 kg–1), and Reeder (678 kg–1). Mixing time of Faller was 8.1 min, significantly longer than Reeder (7.0 min), but shorter than Glenn (9.3 min) and similar to Alsen (9.0 min), Parshall (8.3 min), Howard (8.2 min), and Steele-ND (8.5 min). The mixing tolerance score (15.8 min) was shorter than Glenn (20.6 min), but longer than Howard (12.2 min), Steele-ND (13.5 min), and Reeder (12 min); and similar to Alsen (16.2 min) and Parshall (14.9 min). Loaf volume of Faller (1042 mL) was comparable to Alsen, Howard, Steele-ND, Parshall, and Reeder, but significantly lower than Glenn (1102 mL). The water absorption of Faller (64.3%) was not significantly different than the checks (Table 4).
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Breeder seed of Faller will be maintained by the Seed Stocks Project, Agricultural Experiment Station, North Dakota State Univ., Fargo ND 58105-5051. Multiplication and distribution rights of other classes of Certified seed have been transferred from NDSU to the NDSU Research Foundation, 1735 NDSU Research Park Drive, Fargo, ND 58105-5002. Faller is protected under the U.S. Plant Variety Protection Act with recognized classes of Foundation, Registered, and Certified seed (PVP 200700066). A seed sample has been deposited in the USDA-ARS National Center for Genetic Resources Preservation where it will become available for distribution after expiration of PVP. Small quantities of seed for research purposes may be obtained from the corresponding author for at least five years from the date of this publication. Seed distribution of Faller for research purposes will be made according to the provisions of the Wheat Worker's Code of Ethics (Annual Wheat Newsletter, 1995).
Acknowledgments
The authors thank T. Olson (Dep. of Plant Sciences, NDSU, Fargo, ND), for quality analysis; J. Faller, J. Halley, and all the colleagues at the NDSU REC for their help in conducting trials and nurseries to evaluate Faller.
Footnotes
All rights reserved. No part of this periodical may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permission for printing and for reprinting the material contained herein has been obtained by the publisher.
Received for publication March 20, 2008.
References
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