Published online 31 May 2007
Published in Crop Sci 47:905-914 (2007)
© 2007 Crop Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA
REVIEW & INTERPRETATION
Background and Importance of ‘Minnesota 13’ Corn
A. Forrest Troyera,* and
Lois G. Hendricksonb
a Dep. of Crop Sciences, Univ. of Illinois, Urbana, IL 61801
b Univ. of Minnesota Archives
* Corresponding author (atroyer{at}uiuc.edu).
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ABSTRACT
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Background knowledge of germplasm helps corn (Zea mays L.) breeders develop inbreds and predict hybrids. The background of ‘Minnesota 13’ is still not generally understood. We provide an explanation for the mystery and provide conclusive evidence for Minnesota 13's origin from plant breeding records at the University of Minnesota, whose archives contain corn breeding nursery books. We also cite Minnesota Agriculture Experiment Station annual reports, bulletins, and other pertinent sources. A seed lot collected in 1888 designated as University No. 13 and reported as being improved by Prof. Willet Hays and Andrew Boss in an 1890 bulletin was later destroyed by a seed house fire. They collected a new seed lot in 1893 and the same designation, No. 13, was again used. We show that this second seed lot was common corn grown in the St. Paul, MN, area. We also review the improvement of Minnesota 13 and its influence on the history of corn growing in Minnesota, Wisconsin, and France. We diagram and discuss the background of Pioneer Hi-Bred Inbred PH207 and the importance of Minnesota 13 as 13% of the documented background of U.S. hybrid corn.
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INTRODUCTION
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BACKGROUND KNOWLEDGE of germplasm helps corn (Zea mays L.) breeders develop inbreds and predict hybrids. The origin of ‘Minnesota 13’ was a mystery for most of a century and has only been recently understood. As agriculture spread slowly across the North American continent from 1600 to 1890, the number of corn cultivars increased from about 250 to 1000 (Montgomery, 1916). Crossbreeding Northern Flint and Southern Dent races to develop new cultivars showed benefits appreciated by some experts as early as 1800 (Wallace and Brown, 1988).
Hudson (1994) detailed the corn growing–meat packing movement westward. By 1839, Tennessee and Kentucky joined Virginia to form the original U.S. Corn Belt after early settlers and veterans with land grants found the land attractive. Tennessee led the nation in corn and swine production. By 1858, corn production moved 322 km (200 mi) north and 640 km (400 mi) west to center in Illinois, Ohio, and Missouri. Cincinnati earned the nickname "Porkopolis" by leading the nation in registered slaughterhouses. It shipped pork up and down the Ohio River and over the mountains, and lighted much of the Western Hemisphere with 378 000 L (100 000 gallons) of lard oil annually for lamps. In 1878, Iowa surpassed Illinois in corn production, and about 500 counties made up the U.S. Corn Belt. Corn production moved west because livestock agriculture required corn for animal feed to produce meat for an increasing human population in an expanding USA.
According to Darwin (1868), when a new breed (cultivar) has spread widely, it gives rise to new strains and sub-breeds as a result of natural selection and of artificial (human) selection. The 750 newer, open-pollinated corn cultivars flowered earlier and were more tolerant to droughts, being adapted farther north and farther west (Montgomery, 1916). We document Minnesota 13's more recent origin that moved dent corn still farther north and to higher elevations after American western expansion and provide information about the origin, background, breeders, selection, and importance of Minnesota 13, along with some history of corn and corn growing in Minnesota, Wisconsin, and France.
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MINNESOTA AGRICULTURE BEFORE MINNESOTA 13 CULTIVAR
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The site for Fort Snelling, where the Minnesota joins the Mississippi River at St. Paul and Minneapolis, was established in 1805 by Lt. Zebelon Pike, who was seeking the source of the Mississippi River (Blegen, 1963). Cornfields could be seen growing in the valleys near the site. Domesticated flint corn had been grown in Minnesota since about 1500. Nonetheless, nineteenth-century European settlers considered Minnesota unsuitable for corn growing because dent corn brought from longer growing-season areas did not mature (Hays, 1904).
Professor Willet Hays (1890, 1894, 1895a, 1895b, 1904) described Minnesota corn growing during the 1890s. He called grain more valuable than the fodder in most parts of the state and noted that cultivars grown for fodder were much larger and more thickly planted. According to Hays, early maturing flint and sweet varieties gave the best results for fodder and for grain in the northern third of the state. In the central third, both flints and dents were used. He noted that farmers in the northern two-thirds of the state knew little about suitable corn cultivars to plant or how to save the seed. In the southern third, dents were used almost exclusively. Along the northern edge of the U.S. Corn Belt, small patches of several varieties were planted together without regard to type. Hays (1889) conducted studies in Iowa and in Minnesota showing that shallower cultivation of corn increased yield over deeper cultivation.
In 1894, Minnesota had 155 000 farms averaging 69 ha (170 acres) (Blegen, 1963). Major crops included wheat (Triticum aestivum L.), 1.1 million ha (2.7 million acres); oat (Avena sativa L.), 514 000 ha (1.3 million acres); corn, 326 000 ha (805 000 acres); barley (Hordeum vulgare L.), 198 000 ha (488 000 acres); flax (Linum usitatissimum L.) seed, 126 000 ha (312 000 acres); cultivated hay, 105 000 ha (258 000 acres), and potatoes (Solanum tuberosum L.), 39 000 ha (97 000 acres). During the 1890s, the Crookston (northwest) and Grand Rapids (north central) Minnesota Agricultural Experiment Branch Stations were established to provide better testing to recommend crops and crop cultivars for northern Minnesota (Anonymous, 1896; Troyer and Stoehr, 2003). Before Minnesota 13, only 325 000 ha (800 000 acres) of corn were grown annually in Minnesota.
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DEVELOPERS OF MINNESOTA 13 CULTIVAR
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Willet Hays was born (19 Oct. 1859) and raised on a central Iowa farm near Eldora (Fig. 1). He attended Oskaloosa, Drake, and Iowa State Colleges, earning his bachelor's and master's degrees in 1885 and 1886 at Iowa State. He joined the University of Minnesota faculty in 1888. He helped organize North Dakota State College and Experiment Station in 1891 and 1892 and returned to Minnesota as professor in 1893. The father of American scientific plant breeding, he established the American Breeders Association (now the American Genetic Association) in 1903 (Troyer and Stoehr, 2003). Hays invented and practiced the first systematic pure-line selection with progeny tests (landrace breeding) of oats in the USA at the Minnesota Experiment Station in 1888 (Stanton, 1936). He did hands-on plant breeding from 1888 until 1905 when he became U.S. Assistant Secretary of Agriculture (Boss, 1929; Troyer and Stoehr, 2003; Stoehr and Troyer, 2005). Hays did the first decade of selection on the development of Minnesota 13.
Andrew Boss was born (3 June 1867) and raised on a Minnesota farm near Wabasha. In 1891, he graduated from the University of Minnesota School of Agriculture at University Farm, St. Paul, when he was hired as foreman of the experiment station farm. He was active in plant breeding from 1890 through 1894. He was promoted to full professor in 1905 and become long-time vice director of the Minnesota Experiment Station in 1917. A strong advocate of Professor Willet Hays (Boss, 1929), Boss was active in the initial years of Minnesota 13 development and in Minnesota's increase in corn growing.
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ORIGIN OF MINNESOTA 13
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The University Number Book of 1890 lists accessions of cultivars in numerical order and provides descriptive information including: name, from whom received, post office, state, county, date received, and class (e.g., endosperm color, grain type, silage; Fig. 2). For University Number 13, the entries are St. Paul, DeCou & Co., St. Paul, Minn., Ramsey, 4/1-1893, and Yellow Dent, respectively (Fig. 3). The book lists 215 cultivars from 1890 to 1902. The list of accessions apparently was recopied, with the University numbers of discarded accessions reused; original entries began in 1890, and replaced entries began in 1893. Some of the more popular cultivars listed include Pride of the North, from Northrup King in 1897; Gehu and Northwestern Dent, from Oscar Will in 1897; Reid Yellow Dent, from James Reid in 1901; and Funks 90-day, from Funk Bros. in 1902 (1890 Number Book, p. 80, University of Minnesota Archives).
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Figure 2. The 1890 University Number Book and 1894 Corn Nursery Book (University of Minnesota Archives). Leather-bound ledgers of high-rag-content paper with entries in India ink.
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Figure 3. The 1890 Number Book (p. 80; University of Minnesota Archives). Partial descriptions of the first corn accessions.
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On 1 Apr. 1893, at the request of Willet Hays, Andrew Boss purchased a yellow dent corn cultivar from DeCou & Co. (St. Paul Wire Works and North Star Seed Co.) located in St. Paul, MN (Thomas, 1963). In Director W.M. Liggett's audit ledger for the experiment station, two seed corn purchases from DeCou & Co. are recorded for $1.68 and $1.18 in April 1893 (Fig. 4). One of these two purchases became Minnesota 13.
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Figure 4. University Farm Account Book, April 1893 (University of Minnesota Archives). DeCou & Co. corn purchases.
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In the 1894 Minnesota Corn Nursery Book (University of Minnesota Archives), the word "common" has been added above the cultivar name St. Paul, and the firm name DeCou & Co. has been added in the description information (Fig. 5). The writing matches that of Andrew Boss, who considered the variety to be the common corn grown in the St. Paul area. A few small lots of this seed were sent out as North Star Yellow Dent (p. 150–152, 1893 Minnesota Corn Nursery Book; University of Minnesota Archives), which was probably DeCou & Co.'s name for what Andrew Boss called the St. Paul Yellow Dent common corn that became Minnesota 13. The name North Star comes from "L' etoile du Norde" (The star of the North), which has been on the Minnesota state seal since 1851.
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Figure 5. 1894 Minnesota Corn Nursery Book (p. 72), showing the written in portions for No. 13. (University of Minnesota Archives.)
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What does "common" mean? Dictionaries emphasize "everyday," "usual quality," and "frequent occurrence." Grain-trade terminology dating back to the 1846 Grain Dealers Journal includes a dozen adjectives to describe grain to potential buyers, including choice, fair, common, sound, unsound, yellow, white, mixed, clean, dirty, bright, and hot (Lowell Hill, pers. comm.). To a seeds man, "common" might mean that the variety is not stated. In this case, it probably refers to a cultivar frequently found growing in, and adapted to, the St. Paul area. Hence, St. Paul Yellow Dent common corn was the background of Minnesota 13. Breeders used common corn as a shortcut to local adaptation to develop improved, better-adapted varieties. Leaming Corn, for example, was developed by Jake Leaming near Wilmington, OH, from a common corn grown in the Cincinnati area (Troyer, 1999).
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METHODS USED FOR SELECTING MINNESOTA 13
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The University of Minnesota Experimental Farm is the northwestern quarter of Section 21, Roseville Township, Ramsey County. It is presently bordered by Larpenteur Avenue on the north and Como Avenue on the south, and is immediately east of the St. Paul Campus. The farm is virtually on the 45th parallel. The soil type is predominantly Waukegan (fine-silty, over sandy or sandy-skeletal, mixed, superactive, mesic Typic Hapludolls). Surface layer is about 25 cm (10 in) thick; subsoil is about 52 cm (21 in) thick. It has moderate permeability in upper material and rapid permeability in underlying material. Available water capacity is moderate; surface run-off is slow. It is formed on outwash plains, in valley trains, and on stream terraces. Slope ranges from 0 to 2%. Mean annual air temperature is about 7°C (45°F), and mean annual precipitation is about 71 cm (28 in). Maximum daylength is 15.6 h. Grain yields are no longer taken on the research farm because of bird damage. Based on USDA program payment for the same soil type in the same township, the corn yield is 87% of the Minnesota state average (Troyer, 1999). Natural selection of common corn on the farm would favor earlier flowering plants for the shorter season; the soil type would favor plants with more tolerance to moisture stress. Human selection for Minnesota 13 emphasized earlier maturity (mature ears), higher yield, and higher grain N (protein) (Hays, 1895b).
Hays recognized the individual plant as the unit of improvement, remarking, "There are Shakespeares among plants" (DeVries, 1907; Boss, 1929). Previous breeders bulked seed of selected plants. Hays grew progeny tests ear-to-row from each single plant and used each row's heritability to judge the parent, selecting on a row basis. He originated the centgener breeding method and used it to develop improved varieties of oat, wheat, corn, flax, and dry beans (Vicia faba L.) (Bressman, 1936; Troyer and Stoehr, 2003). In the centgener method, one grows 100 plants generation–1 from a single selected plant, selects the best plant, and then repeats the process. Hays (1894) described harvest as follows: "Harvest the entire centgener, placing all poor plants in a bundle, laying 10% or more of the best plants out side by side. Strip the husks back on these best plants to expose the ears, and select about 50% of these good plants with superior ears. Dry the ears from these best plants, record the weight of each, and record the nitrogen percent of the few that weigh best."
Corn centgeners were thinned to 31 000 plants ha–1 (12 400 acre–1), which was a relatively high plant density for the period. In 1894, six two-row plots were grown each 4.2 m (13.8 ft) long (1894 Corn Nursery Book, p. 44; University of Minnesota Archives), which supplied 454 kg (1000 lb) of dry seed. After 3 yr of centgener selection, 9800 kg (300 bu) were distributed as ‘University 13’ in 1896. In following years, a seed corn plot was planted, poor plants were detasseled or entirely removed, and the best ears from the best plants were selected for distribution as Minnesota 13. The variety was kept free from outcrosses, but the effort in selection was for yield rather than for uniformity (Corn Nursery Book, 1896).
According to a corn breeding essay, likely by Hays, attached to the 1900 Corn Nursery Book (University of Minnesota Archives), they now purchased very little seed of new varieties for increase and sale to good farmers in distant parts of the state who did the same. The demand for Minnesota 13 far exceeded the supply. In 1899, thinning was overlooked resulting in a high plant density (perhaps nearly twice normal). After 1900, Hays saved five ears per centgener, resulting in the first grid system or modified mass selection scheme. From 1905 through 1908, 37, 43, 43, and 45 centgeners were grown, respectively, and selected (Minnesota Corn Nursery Books, 1905–1908, University of Minnesota Archives). Selection and distribution continued through 1910. More than 25 424 kg (1000 bu) were distributed as Minnesota 13 at a dollar per 25.4 kg (bu), which covered production costs.
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DESCRIPTION OF MINNESOTA 13
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Minnesota 13 is a mid-early (95 RM) yellow dent corn. It is early maturing but heavy yielding, adapted from southern Minnesota northward. Ears are 15 to 21 cm (6–8.5 in) long, 16.5 cm (6.5 in) in circumference, and slightly tapered, and they have 12 to 16 kernel rows, medium-size red cobs, and medium-size shank diameter and length. Kernels are medium yellow, medium deep, and medium broad with a medium smooth, dimpled dent. Kernels are compactly set on the cob. The butts are well rounded, and the tips well filled. Plants are 2 to 2.1 m (6.5–7 ft) tall, and ear height is 71 to 79 cm (28–31 in) high (Shoesmith, 1910; Wallace and Bressman, 1923).
A number of strains of Minnesota 13 were subsequently selected. The Morris strain from the Minnesota branch station was popular in west–central Minnesota. The Thorpe strain, popular in North Dakota, was developed by M.L. Thorpe, a prominent corn grower near Mayville, ND. He obtained his original seed stock from the Minnesota Experiment Station. The Haney strain of Minnesota 13 was developed by J.G. Haney, an agronomist with International Harvester. He obtained seed from the Thorpe farm in 1913 and selected near Grand Forks, ND (Olson et al., 1927).
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WHY THE MYSTERY?
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In various reports, Andrew Boss (1929) described Minnesota 13 as a new corn variety obtained by selection of individual plants. He probably considered his additions of "common corn" and "DeCou & Co. St. Paul" in the 1894 Corn Nursery Book sufficient. Certainly, they would have been sufficient if more widely known. In reports on Minnesota 13 calling for materials and methods, Hays repeatedly described the method but omitted materials. To him, the plant breeding method of improvement was the important thing. We find no comments about Minnesota 13's background in his research papers or his correspondence.
The 1890 Minnesota AES Bulletin 11 (Hays, 1890) indicates that Minnesota 13 is from a particular variety, but a 1894 Minnesota Annual Report (Hays, 1894) tells that a seed house fire in fall 1890 destroyed records and carefully selected seed lots. The 1890 Minnesota AES Bulletin 11 is the original source of errors in the literature on the background of Minnesota 13. In the same article, Hays (1890) reports a 3 flint/1 sugary endosperm segregation ratio in the F2 generation 10 yr before the rediscovery of Mendel's laws. In addition, a series of name changes over years for progeny of the DeCou & Co. seed lot further complicated matters. In the 1893 Minnesota Annual Report (Hays, 1893, p. 204), it is called North Star Yellow Dent from DeCou & Co. in St. Paul. In the 1894 Minnesota Annual Report (Hays, 1894, p. 237), it is called St. Paul Yellow Dent from DeCou & Co. in St. Paul. The 1-yr results for North Star Yellow Dent in 1893 and for St. Paul Yellow Dent in 1894 give the correct three- and four-year averages for St. Paul Yellow Dent and University 13 in the next two annual reports (Hays, 1896, p, 311–312). This gives us North Star Yellow Dent, St. Paul Yellow Dent, and University 13 as previous names for the DeCou & Co. seed lot that became Minnesota 13.
The mystery continued because Dr. H.K. Hayes, who came to Minnesota in 1915 was primarily interested in corn inbreds and hybrids, and not cultivars. He was probably the strongest advocate for hybrid corn during that period (Crabb, 1948). His interest in Minnesota 13 was as a source of good inbreds, and he shrugged off questions about its background (Raymond Baker, pers. comm., 1995).
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IMPORTANCE OF MINNESOTA 13 CULTIVAR
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About 324 000 ha (800 000 acres) of corn were grown in Minnesota in 1893 when Willet Hays began breeding improvement of Minnesota 13 cultivar. Minnesota 13 moved ear corn (for grain) growing northward 80 km (50 mi) in the USA in a decade (Hays, 1904). Minnesota acreage doubled to 891 000 ha (2.2 million acres) in 1911 and quintupled to 2 million ha (5 million acres) by 1932. Minnesota 13 became the most popular early cultivar in the northern U.S. Corn Belt and at higher elevations. It was recommended by the Arizona, Colorado, Idaho, Minnesota, Montana, Nebraska, Nevada, New Hampshire, North Dakota, Oregon, South Dakota, Utah, Vermont, Wisconsin, and Wyoming Experiment Stations within 40 yr of its development (Jenkins, 1936). About 3.0 million ha (7.5 and 7.3 million acres) of corn were grown in Minnesota in 2004 and 2005, with average yields of 9971 and 10 912 kg ha–1 (159 and 174 bushels acre–1), respectively (USDA-NASS, 2006). Earlier-flowering, higher-yielding dent corn moved farther north and to higher elevations.
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INITIAL BREEDERS OF MINNESOTA 13 INBREDS
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Herbert K. Hayes was born (11 Mar. 1884) on a corn and tobacco (Nicotiana tabacum L.) farm near Granby, CT. He earned a B.S. at Massachusetts Agricultural College in 1908 and M.S. and D.Sc. at Harvard University in 1911 and 1921 under Dr. E.M. East. Hayes came to Minnesota in 1915 and began studies on inbreeding and hybridizing corn that continued throughout his career. He was promoted to professor and to department head in 1928. Drs. Hayes and E.C. Stakman developed popular Thatcher and other rust-resistant cultivars of hard red spring wheat (Johnson, 2000). Hayes initiated the Foundation Seed Project in the early 1930s and the construction of the Minnesota Crop Improvement Association building in the late 1940s and was senior author of the popular Methods of Plant Breeding (Hayes and Immer, 1942; Hayes et al., 1955). Hayes developed the first useful Minnesota 13 corn inbreds with the help of Drs. Harvey Brewbaker and Iver Johnson (Johnson, 2000).
Why did Minnesota inbreds persist? Hayes was the most experienced and most enthusiastic hybrid corn breeder in the Corn Belt during the 1920s. He started inbreeding corn in Connecticut in 1909 and in Minnesota in 1915 (Crabb, 1948). An early corn-breeding anecdote tells of him developing a Northern Flint inbred with many flag leaves on the husks. Self-pollination was facilitated by cutting back the ear shoot husk cover so that the silks would emerge sooner while the plant was still shedding pollen. This was common practice at the time. Unfortunately, the inbred would not self-pollinate by itself; the silks would ball up in the husks and would not emerge. The inbred was incapable of large-scale seed increase. To prevent this from recurring, Hayes decreed that ear shoots would not be cut for inbred line development. Thus, Minnesota developed early, short-season inbreds that were strong silking and more tolerant to moisture stress. Fresh silks receptive to pollen are more than 90% water (Troyer, 2000). Such inbreds better tolerate drought and persisted as the U.S. Corn Belt expanded north after American westward expansion.
Dr. R. Alexander Brink was born (16 Sept. 1897) on a dairy farm near Woodstock in southern ON, Canada. He earned a bachelor's degree at Ontario Agricultural College at Guelph in 1919, and a master's degree at the University of Illinois in 1922 under Dr. C.F. Hottes. Hottes arranged an Emerson Fellowship at Harvard with Dr. E.M. East, where Brink received the D.Sc. in 1923.
Brink became an assistant professor of genetics at Wisconsin in 1922. He replaced Dr. E.W. Lindstrom who became head of Iowa State's Genetics Department. Brink became professor in 1932 and chairman of the Department of Genetics from 1939 through 1951. He was a great department leader. He brought to the department Dr. Joshua Lederberg, the first University of Wisconsin faculty member to become a Nobel laureate. Brink was elected to the National Academy of Sciences in 1947 and was managing editor of the journal Genetics from 1951 to 1956. He made pioneer studies of paramutation and of transposable elements in corn and developed the popular and hardy Vernal alfalfa (Medicago sativa L.), released in 1954. He also developed a nonbitter, coumarin-free sweet clover (Melilotus officicinalis L.) cultivar (Nelson and Owen, 1995; Brink, 1983).
Brink's association with Drs. East and D.F. Jones in Connecticut made him a strong advocate of hybrid corn (Nelson and Owen, 1995). His first priority on arriving at Madison in 1923 was to start a hybrid corn program in the Genetics department. Agronomist E.J. Delwiche also started inbreeding corn in 1923 at the Spooner branch station in northwestern Wisconsin; he was followed in 1939 by A.M. Strommen. In 1925, Brink expanded hybrid corn breeding by arranging its coordination with the Agronomy department, Plant Pathology department, and the USDA (Nelson and Owen, 1995). They obtained 100 S2 (twice self-pollinated) lines that were self-pollinated near Oconomowoc, WI, in 1920 from Dr. M.T. Jenkins; about half of these lines traced back to Minnesota 13 (Crabb, 1948).
The first Wisconsin corn hybrid was released in 1933. Twenty-five years later, the annual average production of hybrid corn in Wisconsin required 500 000, 25.4-kg (bu) bags of seed. Acreage increased from 450 000 to 680 000 ha (1.1–1.7 million acres). Yield increased from 2188 to 3919 kg ha–1 (35–62.5 bu acre–1), and Wisconsin corn production increased from 890 to 2759 million kg (38.5–108.5 million bu) yr–1 (Rohweder et al., 2003). Dr. Glenn Pound, dean of the College of Agriculture, estimated Wisconsin-bred corn hybrids added $20 million annually to the Wisconsin field corn crops of 1950 to 1970. Questions were no longer raised about the agricultural significance of genetics after hybrid corn came into wide use (Brink, 1983). In 2005, Wisconsin harvested 1.17 million ha (2.9 million acres) for grain that averaged 9 281 kg ha–1 (148 bu acre–1) (USDA-NASS, 2006).
Dr. Norman P. Neal was born (13 May 1897) in Blenheim, NZ, earning a bachelor's degree from the University of New Zealand in 1920. Neal saw his first cornfield after arriving in the USA in 1926. He earned master's degree and doctorate from the University of Wisconsin in 1927 and 1935 under Brink. Neal assumed leadership of the Wisconsin hybrid corn program in 1931 and was awarded the Bledsoe Medal for outstanding contributions to agriculture by the University of New Zealand. He was elected a Fellow of the American Society of Agronomy and of the Crop Science Society of America and was a member of the American Association of Advancement of Science and of the Genetics Society of America (Neal, 1989).
Neal visited France in September and October 1951 and toured corn-breeding stations in France in June and July 1952 (Neal, 1989). A report he published on increasing hybrid corn production in France covered different types of corn hybrids, corn for silage, an outline of a selection program for France, suggestions for tolerance to the cold springtime weather, the use of cold germination tests for seed quality, the need for a guarantee of seed quality and a reserve supply of seed, ventilation necessary for ear corn cribs, and corn utilization (Neal, 1952).
French corn grain yields were about 1 Mg ha–1 from 1840 to 1900. They increased to 1.5 Mg ha–1 from 1935 to 1950. Neal provided Wisconsin inbreds W67, W67C, and W79 (all related to Minnesota 13), as well as W73A, W431, and W567 (all related to ‘Wisconsin 25’), which were used in France in the 1950s (Neal, 1989; Gerdes et al., 1993). Hybrid INRA 200 [(WH x WJ) (F2 x F7)], launched in 1957, contained two inbreds out of Wisconsin 25 from the Spooner, WI, program (Troyer, 2000). A decade or two later Wisconsin second-cycle Inbreds W33 (W9 x WH), W117 (643 x Minnesota 13), W182E (WD x W22), and W401 [(W33 x Wisconsin 25) W67C] were important in France. Inbred W401 contributed to higher plant-density tolerance (Gerdes et al., 1993; Daniel Segonds, pers. comm., 2006). The combination between this early U.S. Dent material and that derived from European local Flint (often French inbreds F2 and F7) proved particularly suitable (Wallace and Brown, 1988; Troyer, 2006).
After Neal's visits and advice in the early 1950s (Neal, 1952, 1989), French corn breeders, agronomists, and farmers increased grain yields to 2.8, 5.0, 5.7, 7.0, and 9.0 Mg ha–1 for 1960, 1970, 1980, 1990, and 2000, respectively. They increased hybrid corn for grain area in France from 325 000 to 600 000, 1.483 million, 1.757 million, 1.912 million, and 1.834 million ha for 1950, 1960, 1970, 1980, 1990, and 2000, respectively. Recent hot, dry summer seasons reduced grain yields, thus increasing proportion of corn area harvested for silage. French 2005 corn grain yield averaged 8.5 t ha–1 on 1.413 million ha. Neal helped hybrid corn spread northward to shorter seasons in the USA and in France (Neal, 1989) (French corn data from Yves Barrier, INRA, Lusignan, France, and from Josiane Lorgeou, ARVALIS, Boigneville, France, pers.comm.).
Perry M. Collins was born (29 Mar. 1902) and raised on a diversified grain and livestock farm near Menlo, IA, about 64 km (40 mi) west of Des Moines. He enrolled at Iowa State College in agriculture but dropped out due to illness. He entered into a farming operation before school started again the next fall. He was interested in livestock breeding and studied books on genetics and animal breeding. He visited Henry A. Wallace in Des Moines in 1926, after reading about the latter's hybrid corn experiments. Wallace asked Collins to grow an isolated crossing block to produce six hybrid corn samples, which Collins did. In late August 1931, Collins accepted a salaried position with Hi-Bred Corn Co. (later Pioneer Hi-Bred International), as its first, full-time employee. He worked with Mr. Raymond Baker near Des Moines in 1931 and 1932.
An accomplished, largely self-taught, corn breeder, Collins started a corn breeding nursery in northern Iowa near Rolfe in 1933. He later credited survivors of the 1934 and 1936 severe heat and drought years (Troyer, 1983) for much of his future success. He had considered discarding them because the years were so atypical. He moved still farther north to Algona, Iowa in 1938. Pioneer Hi-Bred purchased a quarter section of land and picked out the best 40.5 ha (100 acres) for a corn breeding nursery. Mr. Collins spent the rest of his 37 year career, through 1967, breeding corn at Algona IA. He was on a first name basis with many of the progressive farmers in northern Iowa because part of his responsibilities in the early years had been to sell seed corn. He became acutely aware of what farmers wanted in a corn hybrid. He became an active member of the First United Methodist Church and a longtime adult Sunday school teacher. He was an honorary member and past president of the Algona Rotary Club and served as president of the Algona School Board from 1948 to 1960. He also served on the board of directors of the Iowa School Association for three years.
Collins annually developed more than 100 new inbreds and tested hundreds of new hybrids. He discarded many rows of inbred development during flowering for poor synchrony (silk delay). He developed an excellent testing program of six locations in each of three maturity zones. He was an early user of mechanization for harvesting yield trials and originated a successful, double-cross-hybrid breeding scheme, where he systematically substituted an improved, closely related inbred for an older inbred in a superior double-cross formula. He developed several, multimillion-ha Pioneer hybrids, including 349, 352, 3658, 368, 368A, 371, 3755, 3775, 377A and 388. Pioneer 349, first commercially grown in 1947, was the reason competitors voted to drop 5- and 10-yr averages in the Iowa Corn Yield Test reports, where it was usually the highest yielding. Pioneer 349 was grown on more than 10 million ha (Troyer, 1996, 2004b). The first Pioneer single-cross hybrid, 3755, was parented by two of Collins' inbred lines.
Considering the infancy of hybrid corn technology, the competition from 500 local Iowa seed corn companies (Johnson, 1957), and the Great Depression's damper on spending during his early tenure, Collins is arguably the most effective hybrid corn breeder ever, developing most of the first- and second-cycle Pioneer inbreds with Minnesota 13 background.
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INBREDS DERIVED FROM MINNESOTA 13
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Jenkins (1936) reported that five states had developed 25 outstanding inbred lines from Minnesota 13, 7 more than the 18 that eight states had developed from Reid Yellow Dent. Minnesota 13 inbreds had been developed at the following experiment stations: Colorado (9045, 9049, 9064, 9059, 9064, 9070, 9074, 9088, 9092, 9094, 9096, and 9099), Iowa (US153—sole parent of Wisconsin inbred W153), Minnesota (C11, C14, C46, and C49—C stands for culture), North Dakota (91, 106, 178, 207, 210, 211, 212, and 267), and Wisconsin (WM13). Many more Minnesota 13 inbreds were developed after 1936 (Henderson, 1984; Gerdes et al., 1993).
According to Jenkins (1936), the following hybrids containing Minnesota 13 inbreds had been distributed. In Minnesota, Minhybrids 301, 401, and 402 contained Minnesota inbreds C11 and C14 in their females, and in Wisconsin, hybrids 350, 404, 406, 450, 455, 520, 525, and 530 contained Wisconsin inbred WM13 in their pedigree. The first (1934) popular, early maturity, pure yellow endosperm hybrid in northern Iowa and southern Minnesota was Minhybrid 301/Pioneer 355 [(C11 x C14) B164]. Henry A. Wallace had traded Inbred B164 and Purdue Yellow Dent (Duddleston's 461) to Hayes for Inbreds C11 and C14 during the former's attendance at a political rally in St. Paul in 1930 (Troyer, 1999, 2004a).
Wisconsin inbreds W117 and W153R were popular in the 1970s. W117 provided early flowering, rapid grain fill, and fast drying of grain. W153R is a source of Northern leaf blight (Exserohilum turcicum) resistance and of higher test weight. It reduces tillering in hybrids. Wisconsin inbred WM13 (also named M13 and W13) was a grandparent of Ohio inbred OH43 and the source of OH43's early flowering. Montana inbred MT42 was the nonrecurrent parent in Minnesota inbreds A632 and A634. A632 was 7.4, 15.2 and 9.7% of total U.S. seed requirement in 1970, 1975, and 1979, respectively. A634 was 4.2, 7.8, and 3.0% of total U.S. seed requirement in 1970, 1975, and 1979, respectively. North Dakota inbred ND203 is the nonrecurrent parent in Minnesota inbred A635. A635 was 0.7, 0.9, and 1.2% of total U.S. seed requirement in 1970, 1975, and 1979 respectively (Sprague, 1971; Zuber, 1975; Zuber and Darrah, 1980).
Mikel and Dudley (2006) used pedigree information from U.S. Patent and Plant Variety Protection Act records to provide understanding of protected corn germplasm, listing Pioneer Hi-Bred Inbred PH207, developed by Dr. L.J. Hoffbeck at Tipton, IN (40° N lat), as one of seven progenitor inbreds from which much of today's corn germplasm is derived. Walter Vandeventer developed one inbred line parent of PH207 at Tipton, IN. Figure 6 shows the parentage of two Pioneer Modern Iodent inbreds and eight Pioneer Early Iodent inbreds, including PH207.
In 1942, Baker grew 16 ha (40 acres) of breeding populations, segregating for longer ears as a detasseled portion of a production field with Idt x I205 as the male to make seed that was 87.5% Iodent Reid. His objective was to increase hybrid yields of Idt. His scale of program was inspired by Edgar Anderson's (1944) views on effective germplasm and population size. Raymond would later joke that perhaps 8 ha of materials would have been sufficient. He selected for ear length at harvest using ears from the male rows as a comparative check. He kept fewer than 200 ears from the B164 x Lan nonrecurrent portion of the field. He grew the selected ears ear-to-row for two generations and selected for longer ears and better germination before test crossing. Inbreds Idt)2B and Idt)2C resulted (Fig. 6).
In 1958 and 1959, Raymond Baker and A.F. Troyer made breeding populations segregating for maturity; the objective was to make Inbreds Idt)2B and Idt)2C earlier. The inspiration for selection method came from working with Drs. E.L. Pinnell and E.H. Rinke at St. Paul, MN (45° N lat), who developed Inbreds A632 and A634 (Rinke and Sentz, 1961). They suggested a single-cross source of earliness to increase number of earliness genes available. Baker and Troyer chose early inbreds that performed well in early commercial hybrids as sources of earliness. In the early 1960s, Troyer grew populations of 1000 to 1200 plants of several versions of BC1 and BC2 generations, segregating for maturity at twice normal plant density and N fertilizer to select the earliest flowering 10 to 12 plants (1%) to self, to backcross, or to sib mate at Mankato, MN (44° N lat). These selections became the sole pathway to several important commercial Early Iodent inbreds, including Inbred PH207 (Fig. 6; Troyer, 1999, 2004a).
Baker developed Pioneer Hi-Bred/Minnesota Inbred B164 from Troyer Reid via Duddleston's 461 (Crabb, 1948; Troyer, 1999, 2004a). Lindstrom (1931) developed Iowa inbred Lan, a source of long ears and sister to Iowa inbreds L289, L304A, and L317 (Troyer, 2004a). He also developed Idt, a sister to Iowa Inbred I205 (Troyer, 1999). Inbred PH207 is an Early Iodent inbred. The sources of earliness include Minnesota inbreds A78 from Northwestern Dent cultivar and A109, A237, and C49 from Minnesota 13. Collins developed Inbred C49)2A. Inbred PH207 became popular in the USA in the mid-1970s and in France in the late 1970s and early 1980s. Selection for earliness probably increased the proportion of Minnesota 13 cultivar.
Minnesota 13 accounts for 13% of the documented U.S. hybrid corn background tracing back to Minnesota inbreds A109, A237, C49, South Dakota inbred SD105, and Wisconsin inbred WM13 (Fig. 7; Smith et al., 1990; Gerdes et al., 1993; Troyer, 1999, 2004a). Minnesota inbred A109 has Minnesota inbred C49 from Minnesota 13 as a parent. Minnesota inbred A237 has Minnesota inbreds C11 and C23 from Minnesota 13 as parents. South Dakota inbred SD105 was developed from Brookings 86 cultivar—also a Minnesota 13 derivative. In addition, Wisconsin inbred WM13 was developed from Minnesota 13. (Fig. 6, 7).
Dedication
We dedicate this paper to the late Penny Krosch, long-time University of Minnesota archivist. She wrote: "We think we have the corn cultivar accession book you are looking for, but we don't recognize it. Please come help us find it."
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ACKNOWLEDGMENTS
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Raymond Baker encouraged me (A.F. Troyer) to study the background of Minnesota 13. Dr. E.H. Rinke suggested the Minnesota Experiment Station Annual Reports as sources of information. Searching for the corn cultivar accession book led to the University of Minnesota Archives. Eric Wellin, Marcia Thompson, and Julie Rautio helped with the figures. We received helpful suggestions from Bill Ambrose, Alain Charcosset, James Coors, James Crow, Bill Crum, Arnel Hallauer, Daniel Segonds, Howie Smith, and Bill Tracy. The review process helped the paper. Thank you.
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NOTES
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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 October 15, 2006.
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ABSTRACT
INTRODUCTION
