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Multiple Models for Rosaceae Genomics[OA] View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Digital.CSIC Genome Analysis Multiple Models for Rosaceae Genomics[OA] Vladimir Shulaev*, Schuyler S. Korban, Bryon Sosinski, Albert G. Abbott, Herb S. Aldwinckle, Kevin M. Folta, Amy Iezzoni, Dorrie Main, Pere Aru´ s, Abhaya M. Dandekar, Kim Lewers, Susan K. Brown, Thomas M. Davis, Susan E. Gardiner, Daniel Potter, and Richard E. Veilleux Virginia Bioinformatics Institute and Department of Horticulture, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061 (V.S., R.E.V.); Department of Natural Resources and Environmental Sciences, University of Illinois, Urbana, Illinois 61801 (S.S.K.); Department of Horticultural Science, North Carolina State University, Raleigh, North Carolina 27695 (B.S.); Department of Genetics and Biochemistry, Clemson University, Clemson, South Carolina 29634 (A.G.A.); Department of Plant Pathology (H.S.A.), and New York State Agricultural Experiment Station, Department of Horticultural Sciences (S.K.B.), Cornell University, Geneva, New York 14456; Horticultural Sciences Department and Plant Molecular and Cellular Biology Program, Gainesville, Florida 32611 (K.M.F.); Department of Horticulture, Michigan State University, East Lansing, Michigan 48824 (A.I.); Department of Horticulture and Landscape Architecture, Washington State University, Pullman, Washington 99164 (D.M.); Institut de Recerca i Technologia Agroalimenta`ries (IRTA), Centre de Recerca en Agrigeno`mica (CSIC-IRTA-UAB), 08348 Cabrils, Barcelona, Spain (P.A.); Department of Plant Sciences, University of California, Davis, California 95616 (A.M.D., D.P.); United States Department of Agriculture-Agricultural Research Service, Beltsville Agricultural Research Center, Genetic Improvement of Fruits and Vegetables Lab, BARC-West, Beltsville, Maryland 20705 (K.L.); Department of Plant Biology, University of New Hampshire, Durham, New Hampshire 03824 (T.M.D.); and HortResearch, Palmerston North 4442, New Zealand (S.E.G.) The plant family Rosaceae consists of over 100 genera and 3,000 species that include many important fruit, nut, ornamental, and wood crops. Members of this family provide high-value nutritional foods and contribute desirable aesthetic and industrial products. Most rosaceous crops have been enhanced by human intervention through sexual hybridization, asexual propa- gation, and genetic improvement since ancient times, 4,000 to 5,000 B.C. Modern breeding programs have contributed to the selection and release of numerous cultivars having significant economic impact on the U.S. and world markets. In recent years, the Rosaceae community, both in the United States and internationally, has benefited from newfound organization and collaboration that have hastened progress in developing genetic and genomic resources for representative crops such as apple (Malus spp.), peach (Prunus spp.), and strawberry (Fragaria spp.). These resources, including expressed sequence tags, bacterial artificial chromosome libraries, physical and genetic maps, and molecular markers, combined with genetic transformation protocols and bioinformatics tools, have rendered various rosaceous crops highly amenable to comparative and functional ge- nomics studies. This report serves as a synopsis of the resources and initiatives of the Rosaceae community, recent devel- opments in Rosaceae genomics, and plans to apply newly accumulated knowledge and resources toward breeding and crop improvement. ROSACEOUS CROPS FOR HUMAN HEALTH Prunus, Rubus, and Fragaria; see below). Other noned- AND NUTRITION ible species with almost exclusively ornamental value include rose, hawthorn, potentilla, cotoneaster, and Rosaceae, comprised of over 100 genera and 3,000 pyracantha. The products of this family are in high species, is the third most economically important plant demand for their nutritional and aesthetic values, and family in temperate regions (Dirlewanger et al., 2002). their cultivation drives regional economies (Fig. 1). The United States is a leading producer of almond, The Rosaceous tree genera, including Malus, Pyrus, apple, plum, peach, pear, raspberry, sour cherry, sweet Prunus cherry, and strawberry (in the genera Malus, Pyrus, and , are predominantly grown for their fruit and originated in regions extending from Asia west to the Caucasus (Vavilov, 1951). The early Malus and Pyrus lineages gave rise to the cultivated apple * Corresponding author; e-mail [email protected]. (Malus 3domestica), considered to be an ancient allo- The author responsible for distribution of materials integral to the polyploid, with domesticated European and Asian findings presented in this article in accordance with the policy described in the Instructions for Authors (www.plantphysiol.org) is: pears representing multiple species. The fruit-producing Vladimir Shulaev ([email protected]). Prunus also diverged into multiple species. Plums hold [OA] Open Access articles can be viewed online without a sub- the central position within this genus, with numerous scription. species having originated in Europe, Asia, China, and www.plantphysiol.org/cgi/doi/10.1104/pp.107.115618 North America (Smartt and Simmonds, 1995). The Plant Physiology, July 2008, Vol. 147, pp. 985–1003, www.plantphysiol.org Ó 2008 American Society of Plant Biologists 985 Shulaev et al. Figure 1. Representative rosaceous crops exhibiting diversity of fruit types: A, P. persica, peach, fleshy drupe; B, P. armeniaca, apricot, fleshy drupe; C, M. 3domestica, apple, pome; D, F. 3ananassa, strawberry, achenes; E, R. 3hybrida, rose, achene; F, Pyrus communis, pear, pome; G, P. avium, sweet cherry, fleshy drupe; H, P. domes- tica, plum, fleshy drupe; I, Rubus idaeus, raspberry, drupelets. cherry subgenera evolved in Asia Minor, particularly 1999). L-Ascorbic acid, quercetin, kaempferol, myricetin, Iran, Iraq, and Syria (Vavilov, 1951), whereas both p-coumaric acid, gallic acid, and ellagic acid are well- peach and apricot have a Chinese center of origin. Pots known antioxidants and/or cancer-inhibiting com- recovered from cave dwellings in Europe contained pounds that have been identified in these fruits. cherry pits dated from 5,000 to 4,000 B.C. (Marshall, Epidemiological evidence suggests that diets rich in 1954). fruit and vegetables significantly reduce cancer risk During ancient times, rosaceous fruits populated (Caragay, 1992; Ziegler et al., 1996). In vitro and in vivo native human habitats, and fruits from these species studies with animal models provide evidence that fruit were valuable sources of food. Subsequent centuries of and leaf extracts from many Rosaceae species inhibit selection and domestication produced the dramatic some cancers or have strong antioxidant activities (Yau increase in fleshy fruit size that distinguishes today’s et al., 2002). Ellagic acid, present in strawberry, red commercial fruit from their wild relatives. raspberry, arctic bramble, cloudberry, and other rosa- The economic importance of edible rosaceous crops ceous berries (Mandal and Stoner, 1990; Ha¨kkinen et al., derives from their flavorful fruits and nuts that provide 1998; Masuda et al., 1999; Ha¨kkinen and To¨rro¨nen, unique contributions to dietary choices of consumers 2000; Harris et al., 2001; Cordenunsi et al., 2002), has and overall human health. Rosaceous fruits are con- been shown to affect cell proliferation and apoptosis, sumed in multiple forms, including fresh, dried, juice, suggesting a potential anticancer role. Some strawberry and processed products. The variety of flavors, tex- cultivars have the highest concentrations of L-ascorbic tures, and levels of sweetness and acidity offered by acid among all fruits (Haffner and Vestrheim, 1997). these fruits satisfies diverse consumer tastes and Other phenolic compounds belonging to distinct chem- choices. Rosaceae fruits are also a major human dietary ical classes, many of which are also potential antioxi- source of phytochemicals, such as flavonoids and other dants and anticancer agents (Eichholzer et al., 2001; phenolic compounds, cyanogenic glucosides, phytoes- Schieber et al., 2001), have been isolated from rosaceous trogens (Mazur et al., 2000), and phenols that could fruits (Macheix et al., 1991). potentially yield health and disease-fighting advan- Throughout the past century, agricultural research tages (Macheix et al., 1991; Swanson, 1998; Selmar, has focused on increasing crop production to feed the 986 Plant Physiol. Vol. 147, 2008 Rosaceae Genomics growing human population. However, as malnutrition data from six nuclear and four chloroplast loci pro- and infectious and nutrition-related diseases remain vided strong support for the division of Rosaceae into common, the research focus has begun to shift to three subfamilies (Potter et al., 2007; Fig. 2). These include improving the dietary value of different foods include Dryadoideae (including Cercocarpus, Dryas, and identifying novel compounds with pharmacolog- and Purshia; x 5 9), Rosoideae (including Fragaria, ical properties (Kishore and Shewmaker, 1999). Tradi- Potentilla, Rosa, Rubus, and others; x 5 7), and Spi- tionally, new traits for improvement of crop plants raeoideae (including Kerria, Spiraea, and others; x 5 8, have been primarily derived from their related wild 9, 15, or 17). Each of the latter two subfamilies has been species. However, genomics and bioinformatics ad- further divided into supertribes, tribes, and subtribes. vances over the past decade have provided new op- With less emphasis on
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