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Research Gaps and Challenges in the Conservation and Use of North American Wild Lettuce Germplasm

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Research Gaps and Challenges in the Conservation and Use of North American Wild Lettuce Germplasm Published October 24, 2019 REVIEW & INTERPRETATION Research Gaps and Challenges in the Conservation and Use of North American Wild Lettuce Germplasm Aleš Lebeda,* Eva Křístková, Miloslav Kitner, Ľuboš Majeský, Ivana Doležalová, Colin K. Khoury, Mark P. Widrlechner, Jinguo Hu, Daniel Carver, Harold A. Achicanoy, and Chrystian C. Sosa A. Lebeda*, E. Křístková, M. Kitner, and Ľ. Majeský, Dep. of Botany, Faculty ABSTRACT of Science, Palacký Univ. in Olomouc, 78371 Olomouc, Czech Republic; The North American crop wild relatives (CWR) I. Doležalová, Crop Research Institute, Centre of the Region Hana for of lettuce (Lactuca L.) represent an underex- Biotechnological and Agricultural Research, Šlechtitelů 29, 78371 Olomouc, plored pool of genetic diversity of potential Czech Republic; C.K. Khoury and D. Carver, USDA-ARS, National Laboratory for Genetic Resources Preservation, 1111 South Mason St., Fort Collins, CO, value to breeding programs. The 10 species 80521, USA; C.K. Khoury, H.A. Achicanoy, and C.C. Sosa, International Center belong to three different groups: a native clade for Tropical Agriculture (CIAT), Km 17, Recta Cali-Palmira, Apartado Aéreo including at least six allotetraploid species [L. 6713, 763537 Cali, Colombia; M.P. Widrlechner, Departments of Horticulture biennis (Moench) Fernald, L. canadensis L., L. and of Ecology, Evolution and Organismal Biology, Iowa State University, Ames, floridana (L.) Gaertn., L. graminifolia Michx., L. IA 50011, USA; J. Hu, USDA-ARS, Washington State University, Pullman, hirsuta Muhl. ex Nutt., and L. ludoviciana (Nutt.) WA 99164-6402, USA; D. Carver, Natural Resource Ecology Laboratory, Riddell], a diploid clade with one species [L. Colorado State Univ., Fort Collins, CO 80524-1499, USA; C.C. Sosa, Grupo tatarica (L.) C. A. Mey. subsp. pulchella (Pursh) de investigación en Evolución, Ecología y Conservación EECO, Programa de Stebbins], and a clade related to the cultivated Biología, Facultad de Ciencias Básicas y Tecnologías, Universidad del Quindío, taxon (L. sativa L.) with three non-native species Armenia, Colombia. Received 31 May 2019. Accepted 4 Oct. 2019. Assigned (L. saligna L., L. serriola L., and L. virosa L.). In to Associate Editor Jorge da Silva.*Corresponding author ([email protected]). this review, we examine the role of herbarium Abbreviations: AFLP, amplified fragment length polymorphism; CA50, and genebank holdings in taxonomic and other circular area of 50-km radius; CGN, Centre for Genetic Resources, the foundational studies, as well as for germplasm Netherlands; CWR, crop wild relatives; ERSex, ecological representativeness exploration and use. We compile the state of score ex situ; ERSin, ecological representativeness score in situ; FCSc-mean, knowledge on the ranges of lettuce CWR in final combined conservation score (mean); FCSex, final conservation score North America, modeling the potential distri- ex situ; FCSin, final conservation score in situ; GBIF, Global Biodiversity Information Facility; GISH, genomic in situ hybridization; GRIN, Germplasm butions of the species and assessing their ex Resources Information Network; GRSex, geographical representativeness score situ and (for native species) in situ conservation ex situ; GRSin, geographical representativeness score in situ; IPK, Leibniz status. We categorize seven of the species as Institute of Plant Genetics and Crop Plant Research; ITS, internal transcribed high priority for further conservation and three spacer; LGCS, USDA Lactuca Germplasm Collection—Salinas; LGRCD, Lactuca as medium priority, with none currently consid- Genetic Resources Collection, University of California, Davis; NPGS, US ered low priority or sufficiently conserved. National Plant Germplasm System; PGRC, Plant Gene Resources of Canada; Further, we review morphological, phenological, SRS, sampling representativeness score; SSR, simple sequence repeat; WIEWS, genetic diversity, and pest and disease infor- World Information and Early Warning System on Plant Genetic Resources for mation with regard to North American species. Food and Agriculture; WRPIS, Western Regional Plant Introduction Station. We conclude by outlining the critical gaps and describing a way forward for addressing chal- rop wild relatives (CWR) represent a large pool of lenges in the conservation and use of North Cgenetic diversity from which to draw new allelic variation American wild lettuce germplasm. required in breeding programs (Maxted et al., 2006). Crop wild relatives have been extremely valuable in adapting crop varieties Published in Crop Sci. 59:2337–2356 (2019). doi: 10.2135/cropsci2019.05.0350 © 2019 The Author(s). This is an open access article distributed under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). CROP SCIENCE, VOL. 59, NOVEMBER–DECEMBER 2019 to changing pest and disease pressures, farming practices, and seed size (Lebeda et al. 2007; Simko et al., 2014a, market demands, and climatic conditions (Dempewolf et 2014b). All these forms except for two (stem and oilseed) al., 2017). Unfortunately, CWR are a threatened resource are typically consumed raw (Lebeda et al., 2019). (Kell et al., 2012) and measures need to be taken to protect As one of the earliest domesticated vegetables (up to them, both in the wild and in genebanks (Castañeda- 10,800 yr ago), lettuce belongs to a core group of ancient Álvarez et al., 2016; Khoury et al., 2019b; Vincent et al., crops (Hancock, 2012; Zhang et al., 2017). Lettuce is 2019). Limited resources for conservation management considered to be of polyphyletic origin and most likely demand careful planning to prioritize those taxa in most was selected from the genepool of L. serriola (Lebeda et urgent need of conservation (Kell et al., 2017; Magos al., 2007; Kuang et al., 2008) [most probably L. serriola f. Brehm et al., 2017). In this paper, we review how wild integrifolia (Gray) S.D. Prince & R.N. Carter], followed North American Lactuca species, both native and non- by simultaneous introgressions of traits from other closely native, may contribute to the breeding of lettuce varieties related taxa (Lebeda et al., 2019). We suspect that certain and where efforts must be concentrated to harness their other species (L. azerbaijanica Rech. f., L. altaica Fisch. & potential value. C.A. Mey., L. aculeata Boiss. & Kotschy, and L. scario- The genus Lactuca is composed of >100 species, the loides Boiss.) (Zohary, 1991; Koopman et al., 1998; Lebeda majority occur in Asia (51 species) and Africa (43 species), et al., 2007) played roles in the evolution of cultivated some others in Europe (17) (Lebeda et al., 2004b). From lettuce (Lebeda et al., 2019). Based on recent findings, North America, up to 12 species have been described the center of diversity of these related Lactuca species is (Lebeda et al., 2007, 2019; Kilian et al., 2017). Unfortu- in southwestern Asia (i.e., eastern Turkey, Armenia, and nately, from a taxonomic viewpoint, the genus has not northwestern Iran) (Kuang et al., 2008), and the origin been the subject of a modern monograph (Lebeda et al., of lettuce may also spring from this region, especially the 2019). However, useful perspectives on the biosystematics Euphrates–Tigris valley (Zohary and Hopf, 1993). and phylogeny of North American Lactuca species have Although considerable progress has been achieved both recently been published (Jones et al., 2018). in the areas of fundamental research on Lactuca germplasm Within North America, native Lactuca species are and its practical applications during the last 35 yr (Lebeda widely distributed from Canada south to Florida and et al., 2007, 2009, 2014, 2019), the study of Lactuca species Mexico. These species are, in most cases, biennial (Lebeda occurring in North America (Strother, 2006; Lebeda et et al., 2019). Lactuca species native to North America al., 2012, 2019) has been relatively neglected. As part of likely arose relatively recently, in the Pliocene (Kilian a recent volume reviewing the status of CWR in North et al., 2017). Six of them [L. biennis (Moench) Fernald, America (Greene et al., 2019), the first comprehensive L. canadensis L., L. floridana (L.) Gaertn., L. graminifolia overview of wild and weedy North American Lactuca was Michx., L. hirsuta Muhl. ex Nutt., and L. ludoviciana published (Lebeda et al., 2019). Building on the findings (Nutt.) Riddell] are known to be allotetraploids (x = 17) of that review, here we highlight the most important (Strother, 2006; Jones et al., 2018). research gaps and challenges related to the conservation Additionally, three weedy, Old World species (L. and use of wild North American Lactuca. saligna L., L. serriola L., and L. virosa L.) are found in North America (Lebeda et al., 2012), as well as the domesti- PHYLOGENY AND TAXONOMY OF cate, cultivated lettuce (L. sativa L.) (Lebeda et al., 2007). NORTH AMERICAN LACTUCA All these species were introduced to the New World by In the most recent comprehensive treatment of the European settlers (Lebeda et al., 2019), with the introduc- Asteraceae, the genus Lactuca was included in the subtribe tion of L. sativa possibly dating back to Columbus’s second Lactucinae of the tribe Cichorieae (Kilian et al., 2009). The voyage of 1494 (Hedrick, 1972). Two additional taxa, L. Lactucinae host ?200 species, with the greatest diversity terrae-novae Fern. and L. tatarica (L.) C.A. Mey. subsp. located in southwest Asia and the Sino-Himalayan region pulchella (Pursh) Stebbins, are included in the “Synony- (Kilian et al., 2009, 2017; Wang et
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