Safflower in California

Safflower in California The Paulden F. Knowles personal history of plant exploration and research on evolution, genetics, and breeding Safflower in California The Paulden F. Knowles personal history of plant exploration and research on evolution, genetics, and breeding edited by Patrick E. McGuire Ardeshir B. Damania Calvin O. Qualset Citation: McGuire, P.E., A.B. Damania, and C.O. Qualset (eds.) 2012. Safflower in California. The Paulden F. Knowles personal history of plant exploration and research on evolution, genetics, and breeding. Agronomy Progress Report No. 313, Dept. of Plant Sciences. University of California. Davis CA USA. Cover photographs: P.E. McGuire Preface photograph: Photographer unknown, UC Davis Dept. of Plant Sciences collection. Text photographs: P.F. Knowles Contents v Editors’ summary: P.F. Knowles’ genetic resources legacy x i Preface i x Foreword xii Dedication 1 Introduction 1 Early history in California 2 Commercial establishment in California 3 Expanding the germplasm base 7 Distribution of cultivated safflower 10 Preservation of germplasm 10 Germplasm evaluation 12 Species relationships in Carthamus 15 Fatty acid composition of safflower oil 16 Male sterility 17 Miscellaneous traits 19 References Tables 3 Table 1. Itinerary of 1958 collection trip. 4 Table 2. Herbaria visited en route in 1958. 6 Table 3. Itinerary of 1975 collection trip. 7 Table 4. Local names for safflower. 11 Table 5. Selections of safflower with resistance to four races of Fusarium wilt and registered with Crop Science Society of America. 11 Table 6. Summary of number and origin of Verticillium wilt-resistant introductions. 18 Table 7. Association of flower colors with chromatography colors. Figures 5 Figure 1. Route map of 1964–65 trip covering 31,000 miles. 14 Figure 2. Genome evolution in genus Carthamus. iii Photographs x Paul Knowles, UC Davis 4 Two-row planter used for safflower in India in 1965. 8 Minor use of safflower as a vegetable in 1965. 9 Transporting harvested safflower in India in 1965. 13 Threshing and winnowing safflower in India in 1965. 16 Processing safflower for oil in Egypt in 1958. Appendices 23 Appendix 1 Table 1. Count of accessions of cultivated safflowerCarthamus ( tinctorius) collected in 1958. 25 Appendix 1 Table 2. Count of accessions of wild species of safflower (genusCarthamus ) collected in 1958. 26 Appendix 1 Table 3. Count of accessions of cultivated safflowerCarthamus ( tinctorius) collected in 1964–65. 27 Appendix 1 Table 4. Accessions of wild species of safflower (genusCarthamus ) collected in 1964–65. 28 Appendix 1 Table 5. Accessions of Carduncellus species collected in 1964–65. 29 Appendix 1 Table 6. Safflower germplasm selected in China in 1988. 33 Appendix 2. Publications of P.F. Knowles on oilseed crops. 41 Appendix 3. List by species, year, and PI number of 1175 safflower accessions associated with P.F. Knowles in the collection at the USDA-ARS-RPIS, Pullman WA USA. iv Editors’ summary: P.F. Knowles’ genetic resources legacy At the start of Paul Knowles’ career, safflower was only incidentally grown in California. By virtue of his work, the crop was introduced and improved and production and distribution became a California industry. Central to this progress was germplasm acquisition, evaluation, conservation, and utilization carried out with Knowles’ vision and leadership. His achievements were far reaching. Today, when crop genetic resources are paradoxically both increasingly recognized as highly valuable and yet confronted with ever greater risk of being lost, his story has valuable lessons. • Accessible, wide-ranging genetic diversity is essential for the genetic improvement of a crop. Knowles recognized early on that safflower breeding programs in the US had a narrow genetic base and this lack of germplasm was restricting progress. • Accessibility of genetic diversity requires international exchange and collaboration. Knowles was supported, directly and indirectly, in his efforts by his institution, several US agencies (e.g., USDA, USAID, US Dept. of State), the UN Food and Agriculture Organization, and numerous academic and national researchers and institutions and private growers and sellers in the many countries in which he traveled. He initiated and facilitated exchanges of germplasm, information, and scholars. • Collection of germplasm should be made across all environments and geographic regions in which the target species or crop grows. The mutations he reports and worked with for safflower fatty acid composition arose in very small popula- tions in widely dispersed and unpredictable sites (India, Iran, Israel, Portugal, and Russia) and would never have been found without access to wide-ranging germplasm collections. • Useful genetic diversity can be found in landraces of the crop from regions where it has a long history of cultivation and in wild species closely related to the crop species. He reports useful genes and traits for cultivated safflower from landraces (e.g., the fatty acid composition mutations, flower color variants, growth habit, disease resistances, and male sterility) and from wild species (disease resistances). v • Indigenous germplasm is being rapidly replaced with introductions and newly developed varieties in many regions and environments, it is critical that material continues to be collected for ex situ conservation from these areas. He recognized early in his collecting career that safflower landraces that had been grown for centuries in some countries were being replaced by crops and cultivars from developed nations. • Research on species relationships and distribution, ecology, ethnobotany, cytogenetics, and basic biol- ogy are crucial for efficient and directed utilization of landraces and wild species for improvement of related crops. Over his career, his graduate students, postdocs, and research colleagues compiled a broad record of research efforts on safflower that encompassed basic agronomy, pathology, cytogenetics, genetics, biochem- istry, and biogeography. The productivity of this research effort depended in a large way on the safflower genetic resources collection (the World Collection) to which he was a major contributor. At the same time, these research efforts enhanced the value of the World Collection as the resulting data became part of the information associated with the collection. • Evaluation and characterization of genetic resources (landraces and breeding lines) under diverse environments and stress conditions are essential for identifying characters useful for crop improvement. His work in California with the World Collection and screening for disease resistances and adaptive traits and the work with collaborators in other countries and environments exemplify the value of the evaluation and characterization process for ultimately successful germplasm utilization. • Novel genetic diversity can lead directly to successful cultivar development. Safflower cultivar ‘UC-1’, released by Knowles in 1968, was essentially the first commercial variety of a new type of safflower oil, a new crop. Its oil properties (high oleic acid) differed in fatty acid composition from that of traditional safflower oil (high linoleic acid) as a result of a mutation discovered in a safflower accession from India. The oil of UC-1 was chemically similar to olive oil. The identification of the novel oil type was made possible only by screening diverse collections of safflower germplasm for variation in fatty acid properties. The Indian accession was crossed in Knowles’ breeding program to a Nebraska cultivar (N-10) in 1957 to initiate the incorporation of the novel oil type into an agronomically acceptable background. By 1967, after a backcrossing breeding program, certified seed of the new cultivar was available and it was the foundation for California production for several years and for commercial breeding programs thereafter. • Crop introduction is facilitated by agronomic research and outreach and extension to cultivate growers, producers, and markets for production. His work in this area early in his career laid the foundation for all his subsequent work. The rising success of safflower as a California crop gave impetus and support for his subsequent breeding, evolution, and geographic interests in the crop and its wild related species. • Plant ‘breeding’ in the broadest sense of plant selection, use, and improvement is a story of human curiosity, need, innovation, and persistence. On his travels, he didn’t interact only with breeders and seed dealers, he sought out farmers, market sellers, and users of the crops in which he was interested. This report on safflower touches on some of the many anecdotal presentations of the ways of growing, harvesting, processing, and using (in art, medicine, industry, and cuisine). The similarities and contrasts that he observed from one country and tradition to another in all these steps with a common plant are enlightening and a testimony to his own sense of curiosity and the apparent value he placed on each person’s contribution, testimony, and information. • Serendipity plays a significant role in revealing the utility of germplasm collections. Two examples from Knowles’ work illustrate this: 1. Lettuce is known commonly as a leafy vegetable, and more distantly in time, it was cultivated for its stem, but on both his 1958 and 1964–1965 collection trips, he observed the use of a primitive lettuce vi in Egypt as an oil crop in areas where safflower was also cultivated. Its seeds were being

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