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a Dep. of Crop and Soil Sciences, Washington State Univ., Pullman, WA 99164-6420
b USDA-ARS Wheat Genetics, Quality, Physiology, and Disease Research Unit, Pullman, WA 99164-6420
c Plant, Soils, and Entomological Sciences, Univ. of Idaho, Moscow, ID 83844-2339. Research was funded in part by the Washington State Grain Alliance
* Corresponding author (kidwell{at}wsu.edu).
ABSTRACT
Hard red spring (HRS) wheat (Triticum aestivum L.) grain must have high grain protein concentration (GPC) to maximize market value. The objective of this research was to develop a HRS wheat cultivar with high GPC and durable resistance to stripe rust (Puccinia striiformis Westend f. sp. tritici), a major foliar fungal disease problem of spring wheat. ‘Kelse’ (Reg. No. CV-1035, PI 653842) HRS wheat was developed and released in July 2008 by the Agricultural Research Center of Washington State University. Kelse was tested under the experimental designations of WA007954, H0100092, and HR98036, which were assigned through progressive generations of advancement. Kelse is a semidwarf cultivar adapted to the intermediate-to-high rainfall (>380 mm average annual precipitation), nonirrigated wheat production regions of Washington State. Kelse was released based on its (i) resistance to the Hessian fly [Mayetiola destructor (Say)], (ii) high-temperature, adult-plant resistance (HTAP) to local races of stripe rust (infection type 0–3), (iii) high GPC, (iv) excellent bread-baking quality, and (v) high grain yield potential in the target production region. Kelse is the first HRS wheat cultivar adapted to the Pacific Northwest region of the United States with HTAP resistance to stripe rust. Based on molecular marker analysis, Kelse has the Lr34/Yr18 and Lr37/Yr17/Sr38 gene clusters for rust resistance. Kelse also has exceptionally high GPC compared with HRS cultivars currently in production.
Abbreviations: DOY, day of year • GPC, grain protein concentration • HRS, hard red spring • HTAP, high-temperature, adult-plant • IT, infection type • PST, Puccinia striiformis Westend f. sp. Tritici • WSU, Washington State University
Grain protein concentration (GPC) has a major impact on the end-use quality of bread products made with flour extracted from hard wheat (Triticum aestivum L.) grain; therefore, this trait is typically a high priority in wheat improvement efforts aimed at improving bread-making quality. Due to improved profit potential compared with soft white spring wheat, the acreage of hard red spring (HRS) wheat has increased dramatically in the Pacific Northwest region of the United States in recent years (Washington Agricultural Statistics Service, 2005–2008). The objective of this research was to develop a HRS wheat cultivar with high GPC, superior agronomic characteristics, and improved end-use quality attributes compared with other HRS cultivars currently in commercial production.
‘Kelse’ (Reg. No. CV-1035, PI 653842) HRS wheat was developed and released in July 2008 by the Agricultural Research Center of Washington State University. Kelse was named in honor of Kelsey L. Kidwell-Yonan, Dr. Kidwell's beloved niece and treasured friend. Kelse was released as a replacement to ‘WestBred 926’ (proprietary cultivar from WestBred LLC, Bozeman, MT), ‘Hank’ (PI 613585; proprietary cultivar from WestBred LLC, Bozeman, MT), ‘Tara 2002’ (PI 617073; Kidwell et al., 2003b), and ‘Scarlet’ (PI 601814; Kidwell et al., 1999) based on its (i) resistance to the Hessian fly [Mayetiola destructor (Say)], (ii) high-temperature, adult-plant resistance (HTAP) to local races of stripe rust (Puccinia striiformis Westend f. sp. tritici), (iii) high GPC, (iv) excellent bread-baking quality, and (v) high grain yield potential in the intermediate-to-high rainfall (>380 mm average annual precipitation), nonirrigated wheat production regions in Washington State.
Methods
Kelse, tested under the experimental designations WA007954, H0100092, and HR98036, which were assigned through progressive generations of advancement, is an F3:4 headrow selection derived from the cross ‘WestBred 906R’ (PI 483455)/SD 2961 (PI 520542)//‘Scholar’ (PI 607557). WestBred 906R is an HRS variety derived from a facilitative male sterile recurrent selection released in 1981 by WestBred LLC, Bozeman, MT. SD 2961 is an unreleased HRS wheat breeding line developed by South Dakota State University in 1987 with the pedigree ‘Butte’ (CItr 17681)//‘Olaf’ (CItr 15930)/MN 6792 (PI number not available). Scholar is an HRS wheat variety released in 1999 by Montana State University (Lanning et al., 2000), with the pedigree MT7746 (PI number not available)/‘Lew’ (CItr 17429)//‘Marberg’ (PI 518816). The final cross for Kelse was completed in the greenhouse in Pullman, WA, in 1997, and the following modified pedigree-bulk method was used to advance early generation progeny. Bulked seed (30 g) from F1 plants was used to establish a 3.6-m2 F2 field plot in 1998. Approximately 100 heads were selected at random from individual F2 plants, and a 40-g subsample of the bulked seed was used to establish a single 7.2-m2 F3 field plot in 1999. Single heads from approximately 150 F3 plants were selected at random and threshed individually to establish F3:4 headrow families in the field in 2000. F1 progeny were advanced at the Washington State University (WSU) Plant Growth/Wheat Research Facility on the WSU campus in Pullman, WA. F2 and F4 progeny were advanced in field nurseries at Pullman, WA, whereas F3 progeny were advanced at the Lind Dryland Experiment Station in Lind, WA.
Following selection among rows for general adaptation, resistance to stripe rust, plant height, and grain appearance, seed from all plants within each selected headrow was bulk harvested to obtain F3:5 seed for early generation quality assessment. A 15-g subsample of each selected head row was cyclone milled (UDY Corporation, Fort Collins, CO) using a 0.5-mm screen. Resulting whole wheat flour was evaluated for protein content and hardness using a Bran+Luebbe InfraAlyzer 450 (Bran+Luebbe GmbH, Norderstedt, Germany). Selections with appropriate grain protein concentration and kernel hardness were evaluated for gluten strength using the sodium dodecyl-sulfate-sedimentation method (American Association of Cereal Chemists, 2000). Twenty-five headrow selections were grown in 7.4-m2 plots in a nonreplicated field trial in Lind, WA, in 2001, and resulting grain was evaluated for grain yield, grain volume weight, grain protein concentration, disease resistances, and milling and baking quality. All subsequent years of field testing involved the same size plot area and data collection strategy using either a randomized complete block design (four replications) (2002–2005) or a general alpha lattice design (three replications) (2006–2008) (Mason et al., 2003). Four of the original 25 selections were advanced to preliminary replicated, nonirrigated field trials in Pullman and Lind, WA, in 2002. Two selections were advanced to state replicated, nonirrigated trials in Pullman, Lind, Connell, and Dusty, WA, in 2003. One line, designated H0100092, was selected to be tested on a regional basis and assigned the new identification number WA007954 in 2004. In 2004 and 2005, WA007954 was evaluated in the Tri-State Regional Nursery, which was established at nine locations each year in Washington, Oregon, and Idaho, in both rainfed and irrigated environments. In 2006, WA007954 was entered in the Washington State University Extension Uniform Cereal Variety Testing Spring Wheat Performance Trials and tested in 2006, 2007, and 2008 throughout eastern Washington at 16, 17, and 15 locations, respectively. Since 2001, WA007954 was evaluated for end-use quality attributes and disease resistance by the USDA–ARS Wheat Genetics, Quality, Physiology, and Disease Research Unit, Pullman, WA. WA007954 was evaluated by the Pacific Northwest Wheat Quality Council in 2007.
Breeder seed of Kelse (WA007954) was produced as a reselection, on the basis of phenotypic uniformity, of 1400 F3:11 headrows grown under irrigation in Othello, WA, in 2007. Approximately 70 headrows were removed from the population, resulting in 1330 selected headrows that were bulked at harvest, resulting in the production of 573 kg of Breeder seed.
Agronomic data generated from 2002 to 2005 were analyzed as a randomized complete block, whereas data from 2006 to 2008 were analyzed using the general lattice (ALS) procedure of Agrobase Generation 2, version 18.3.1 (Agronomix Software, Inc. Winnipeg, Canada). Since four major wheat-producing regions with distinct climatic conditions exist in Washington, data were analyzed across locations within regions instead of over all locations. The 2006 to 2008 data from the high rainfall and irrigated region were combined for analysis. Location means and ranks from 2006 to 2008 were generated using the arithmetic mean of the ALS adjusted mean. The arithmetic means and ALS adjusted means were subjected to analysis of variance, and breeding lines were advanced on the basis of exceptional performance within each location, across locations within a region, and across regions within a year. Once Kelse was selected for release, final data analysis only included entries common to all trials across all years. End-use quality data were analyzed by the Student's paired t test procedure (Cochran and Cox, 1957).
Characteristics
Kelse is an intermediate height, semidwarf spring wheat with lax, tapering, erect curvature inflorescence with tan awns and glumes that are long in length, wide in width and have medium, oblique shoulders and narrow, acuminate beaks. Kelse has oval kernels that are red, hard, and vitreous. Seed of Kelse has a large germ with a medium-depth crease, angular cheeks, and a short, noncollared brush. Kelse lacks anthocyanin pigmentation in the coleoptile and displays a semi-erect juvenile plant growth habit. Kelse's flag leaf is green in color, recurved, twisted, and waxy at Feekes growth stage 10.0 (Large, 1954). The stem of Kelse has three nodes, a waxy bloom, semisolid last internode of the rachis, and pigmented auricles, it lacks anthocyanin pigmentation and pubescence, and its peduncles are erect with an average length of 41 cm. Data generated from 2006 to 2008 demonstrate that Kelse headed (day of year [DOY] 168) 3 d later than Hank (DOY 165) and 5 d later than WestBred 926 (DOY 163) and Tara 2002 (DOY 163). From data collected from 2006 to 2008, the average plant height of Kelse was 78 cm, which was significantly (P < 0.05) taller than WestBred 926 (72 cm) and Hank (70 cm) and equal to Tara 2002 (78 cm). Lodging percentages of Kelse (<1%) when grown with irrigation were comparable to Hank (<1%), and lower than WestBred 926 (3–5%) and Tara 2002 (15–20%) (P < 0.05).
Disease and Insect Resistance
In naturally infected field tests on the Whitlow Farm near Pullman, WA, and at Mt. Vernon, WA, from 2002 to 2005, on the Spillman Agronomy, Plant Pathology, and Whitlow farms near Pullman, WA, in 2006 and 2007, and in breeding nurseries at multiple locations throughout eastern Washington from 2002 to 2007, Kelse displayed high levels of non–race-specific HTAP resistance (infection type [IT] 0–3) at all locations in eastern Washington and moderate resistance (IT 5) at Mt. Vernon, WA. In greenhouse seedling tests conducted in 2006 and 2007 under low temperature cycles (diurnal temperatures gradually changing from 4 to 20°C; Chen and Line, 1992), Kelse was resistant (IT 2) to races PST (Puccinia striiformis Westend f. sp. tritici)-37, PST-45, and PST-100; intermediate (IT 5) to PST-17 and PST-43; but susceptible (IT 7–8) to PST 116 and PST-127. When adult plants were tested in the greenhouse at high temperatures (diurnal temperature cycle gradually changing from 10 to 35°C; Chen and Line, 1995), Kelse was resistant (IT 0–3) to races PST-45, PST-100, and PST 116 and moderately resistant (IT 5) to PST-127. The contrasting reactions of seedlings compared to adult plants with races PST-116 and PST-127 indicate that Kelse has a moderate level of non–race-specific HTAP resistance to stripe rust, which has proven to be durable in other commercial spring wheat cultivars, such as ‘Louise’ (PI 634865; Kidwell et al., 2006). Kelse is the first HRS variety released in the Pacific Northwest region that is confirmed to have HTAP resistance to stripe rust.
Results of pedigree analysis indicate that Kelse may have inherited its stripe rust resistance from SD 2961 and subsequently from ‘Frontana’ (PI 500147; Singh and Rajaram, 1992). DNA marker csLV34, which is linked to the Lr34/Yr18 complex present in Frontana, was tested on Kelse (Lagudah et al., 2006). Results confirmed the presence of Lr34/Yr18 in Kelse. Kelse also was analyzed using the primer pairs VENTRIUP-LN2, linked to the Lr37/Yr17/Sr38 complex from T. ventricosum (Helguera et al., 2003). Results indicate that Kelse contains the Lr37/Yr17/Sr38 genes for rust resistance. Data from the 2008 Western Regional Spring Wheat Nursery indicate that Kelse has a moderate resistance reaction to leaf rust, as well as to race TRTT of stem rust (USDA–ARS, 2009).
On the basis of results from 2 yr of controlled environment insect screening trials conducted at the University of Idaho in 2005 and 2006, Kelse is resistant (100%) to Hessian fly biotypes E, F, and GP. In 2005, Kelse was tested with the susceptible cultivar Alpowa (PI 566596) and the resistant cultivar Macon (PI 617072; Kidwell et al., 2003a) in a five replication trial with four plants per replicate. Kelse and Macon were 100% resistant, whereas Alpowa was 100% susceptible. In 2006, Kelse was tested with the susceptible cultivar Alturas (PI 620631; Souza et al., 2004) and the resistant cultivar UI Cataldo (PI 642361) in a replicated trial with five replicates, and four plants tested per replicate. Kelse and Cataldo were 100% resistant, whereas Alturas was 100% susceptible. On the basis of pedigree and natural field infestation ratings from Pullman, WA, Kelse is susceptible to the Russian wheat aphid [Diuraphis noxia (Mordvilko)].
Agronomic Performance
Kelse was evaluated in 60 replicated field trials under fallow, rainfed, and irrigated conditions in Washington State from 2002 to 2008. In 12 trials conducted from 2002 to 2005 in low (<380 mm average annual precipitation), intermediate (380–460 mm average annual precipitation), and high precipitation zones (>460 mm average annual precipitation) and with irrigation, the average grain yield of Kelse, WestBred 926, and Scarlet were 3225, 3153, and 2755 kg ha–1, respectively. Grain yield averages of Kelse and WestBred 926 did not differ significantly (P < 0.05) and were significantly higher than grain yield averages of Scarlet. Average grain volume weight of Kelse (750 kg m–3) was similar to WestBred 926 (744 kg m–3) and significantly higher than the grain volume weight of Scarlet (736 kg m–3) (P < 0.05).
In 21 rainfed trials conducted from 2006 to 2008 in the semiarid region in Washington State, average grain yields of Kelse, WestBred 926, Tara 2002, and Hank were 2278, 2298, 2305, and 2331 kg ha–1, respectively, and were statistically similar (P < 0.05). In 17 rainfed trials conducted from 2006 to 2008 in the intermediate rainfall zones in eastern Washington, the grain yield averages of Kelse, WestBred 926, Tara 2002, and Hank were 3682, 3615, 3675, and 3695 kg ha–1, respectively, and were not significantly different (P < 0.05). In 10 replicated field trials conducted from 2006 to 2008 in eastern Washington in the high rainfall zones and under irrigation, the grain yield averages of Kelse, WestBred 926, Tara 2002, and Hank were 5160, 5274, 5362, and 5671 kg ha–1, respectively. Grain yield averages of Kelse and WestBred 926 did not differ significantly but were significantly lower than those of Tara 2002 and Hank (P < 0.05).
In these same field trials, the average grain volume weight of Kelse (766 kg m–3) was significantly (P < 0.05) higher than WestBred 926 (755 kg m–3) and Hank (752 kg m–3) and equal to that of Tara 2002 (764 kg m–3). Thousand-kernel weight averages of Kelse, WestBred 926, Tara 2002, and Hank were 38.3, 44.4, 50.0, and 37.7 g, respectively. Thousand-kernel weight estimates were not replicated; therefore, statistical analysis was not conducted on these data.
End-Use Quality
In tests conducted at the USDA–ARS Western Wheat Quality Laboratory in Pullman, WA, using grain produced in 26 breeding and commercial variety testing trials in Washington from 2001 through 2007, the milling and baking qualities of Kelse were compared with WestBred 926, ‘Hollis’ (Kidwell et al., 2004), Tara 2002, and Scarlet. Grain protein concentration averages of Kelse (155 g kg–1) were similar to those of both WestBred 926 and Hollis (149 g kg–1), and were significantly (P < 0.01) higher than those of Tara 2002 (146 g kg–1), and Scarlet (143 g kg–1). Flour yields of Kelse (680 g kg–1) were significantly (P < 0.01) lower than those of WestBred 926 (692 g kg–1), Hollis (701 g kg–1), Tara 2002 (694 g kg–1), and Scarlet (697 g kg–1). Flour ash content for Kelse (4.5 g kg–1) was similar to WestBred 926 (4.3 g kg–1), and significantly (P < 0.01) higher than Tara 2002 (4.0 g kg–1), Hollis (3.8 g kg–1), and Scarlet (3.8 g kg–1). Kelse had a lower average milling score (80.0) than WestBred 926 (82.1), Tara 2002 (84.0), Scarlet (85.3), and Hollis (85.7). Dough mixing time of Kelse (4.2 min) was similar to Scarlet (4.0 min) and WestBred 926 (4.1 min), slightly shorter than Hollis (4.5 min), and significantly (P < 0.01) shorter than Tara 2002 (6.1 min). Mixograph water absorption of Kelse (647 g kg–1) was similar to that of WestBred 926 (642 g kg–1) and Hollis (640 g kg–1) but was significantly (P < 0.01) higher than Scarlet (639 g kg–1) and Tara 2002 (637 g kg–1). Average 100-g pup loaf volume for Kelse (1091 cm3) was comparable to Tara 2002 (1084 cm3) and Hollis (1085 cm3) and significantly (P < 0.01) larger than that of WestBred 926 (1044 cm3) and Scarlet (996 cm3) when compared across production regions. All the above tests were conducted using approved AACC methods (American Association of Cereal Chemists, 2000).
In 2007, Kelse was evaluated by commercial millers and bakers in the Pacific Northwest Wheat Quality Council. These results indicate that Kelse had superior end-use quality attributes when compared with WestBred 926 in nearly every assessment category (data not shown). Out of the 28 wheat varieties tested, Kelse ranked third overall for superior end-use quality attributes, whereas WestBred 926 ranked 18th (Pacific Northwest Wheat Quality Council, 2008). All collaborators preferred the dough handling properties and baking quality of Kelse compared with WestBred 926, which were attributed to the superior protein quality of Kelse compared with WestBred 926. The protein content, protein quality, and baking quality attributes of Kelse overcame any concerns about milling quality and ash content from a commercial users' perspective.
Availability
Foundation seed of Kelse will be maintained by the Washington State Crop Improvement Association under supervision of the Department of Crop and Soil Sciences and the Washington State Agricultural Research Center. Small quantities may be obtained for research purposes by contacting the National Plant Germplasm System. U.S. Plant Variety Protection status for this cultivar is pending.
Acknowledgments
We greatly appreciate the dedicated assistance of the following support personnel who assisted with field testing, quality assessment, and disease screening during the development of this cultivar: Tracy Harris (WSU), Dave Wood (USDA–ARS), Kelley Richardson (USDA–ARS), and Dennis Schotzko (Univ. of Idaho). We also thank Dr. Craig Morris, USDA–ARS Wheat Genetics, Quality, Physiology, and Disease Research Unit, Pullman, WA, and John Burns, WSU Extension Agronomist, for providing access to facilities, equipment, and personnel for conducting end-use quality and agronomic evaluations, respectively. We are grateful to the Washington State Grain Alliance and the Washington State Agriculture Research Center for providing funding for this research. Marker analysis was supported by the National Research Initiative of USDA's Cooperative State Research, Education and Extension Service, CAP Grant No. 2006-55606-16629.
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 January 5, 2009.
References
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