Example Pennycress (Thlaspi Arvense L.)
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Plant Science 227 (2014) 122–132 Contents lists available at ScienceDirect Plant Science j ournal homepage: www.elsevier.com/locate/plantsci Review New approaches to facilitate rapid domestication of a wild plant to an oilseed crop: Example pennycress (Thlaspi arvense L.) a,∗ b c John C. Sedbrook , Winthrop B. Phippen , M. David Marks a School of Biological Sciences, Illinois State University, Campus Box 4120, Normal, IL 61790 USA b School of Agriculture, Western Illinois University, 1 University Circle, Macomb, IL 61455, USA c Department of Plant Biology, University of Minnesota, 1445 Gortner Avenue, 250 Biological Sciences Center, Saint Paul, MN 55108, USA a r t i c l e i n f o a b s t r a c t Article history: Oilseed crops are sources of oils and seed meal having a multitude of uses. While the domestication of Received 20 May 2014 soybean and rapeseed took extended periods of time, new genome-based techniques have ushered in Received in revised form 23 July 2014 an era where crop domestication can occur rapidly. One attractive target for rapid domestication is the Accepted 25 July 2014 winter annual plant Field Pennycress (Thlaspi arvense L.; pennycress; Brassicaceae). Pennycress grows Available online 4 August 2014 widespread throughout temperate regions of the world and could serve as a winter oilseed-producing cover crop. If grown throughout the USA Midwest Corn Belt, for example, pennycress could produce as Keywords: much as 840 L/ha oils and 1470 kg/ha press-cake annually on 16 million hectares of farmland currently Oilseed left fallow during the fall through spring months. However, wild pennycress strains have inconsistent Thlaspi arvense Pennycress germination and stand establishment, un-optimized maturity for a given growth zone, suboptimal oils Domestication and meal quality for biofuels and food production, and significant harvest loss due to pod shatter. In this Next generation sequencing review, we describe the virtues and current shortcomings of pennycress and discuss how knowledge TILLING from studying Arabidopsis thaliana and other Brassicas, in combination with the advent of affordable next generation sequencing, can bring about the rapid domestication and improvement of pennycress and other crops. © 2014 Elsevier Ireland Ltd. All rights reserved. Contents 1. Introduction . 123 2. Phylogeny, geographic distribution, and traits of pennycress. 123 2.1. Phylogeny . 123 2.2. History and geographic distribution of collected germplasm . 123 2.3. Growth characteristics and anatomy. 125 2.4. Agronomic traits . 125 3. Overview of domestication . 126 4. New approaches to facilitate rapid domestication . 126 4.1. Next generation sequencing and the pennycress genome . 126 4.2. Targeting Induced Local Lesions IN Genomes (TILLING) . 127 4.3. Targeted genome editing . 128 5. Traits and genes to target for pennycress domestication. 128 5.1. Seed dormancy . 128 5.2. Oil quality . 128 5.3. Seed glucosinolates . 129 5.4. Flowering time and maturation . 129 5.5. Pod shatter . 129 Abbreviation: TILLING, Targeting Induced Local Lesions IN Genomes. ∗ Corresponding author. Tel.: +1 309 438 3374; fax: +1 309 438 3722. E-mail address: [email protected] (J.C. Sedbrook). http://dx.doi.org/10.1016/j.plantsci.2014.07.008 0168-9452/© 2014 Elsevier Ireland Ltd. All rights reserved. J.C. Sedbrook et al. / Plant Science 227 (2014) 122–132 123 5.6. Seed size and oil content . 130 5.7. Orthologs in pennycress . 130 6. Conclusion . 130 Acknowledgements . 130 References . 130 1. Introduction affordable next-generation sequencing tools, attractive gene tar- gets, and demonstrated transformability (all of these topics are described herein), the time is ripe for pennycress to be rapidly Plant oils in the form of triacylglycerols are among the most domesticated. energy-rich abundant biomolecules in nature, accumulating in the embryos and endosperm of a variety of plant species where they are energy stores used by the germinating seedlings [1]. Triacyl- 2. Phylogeny, geographic distribution, and traits of glycerols also accumulate to high levels in fruit mesocarp tissues of pennycress trees such as oil palm (Elaeis guineensis), olive (Olea europaea), and avocado (Persea americana), functioning as edible attractants that 2.1. Phylogeny facilitate seed dispersal [2,3]. Soybean (Glycine max L.) and rapeseed (Brassica napus) are two Pennycress is a member of the mustard family and is closely major seed oil-producing crops worldwide, being cultivated on 105 related to a large number of agronomically and scientifically rel- million and 25 million hectares of land in 2012/13, respectively, evant plants including the oilseed crops rapeseed (B. napus and producing over 300 million metric tons of seed [4]. Soybean seeds Brassica rapa varieties), canola (rapeseed variety double-low in contain about 20% oil and 40% protein (dry basis) whereas rapeseed erucic acid and glucosinolate), and camelina (Camelina sativa), the seeds contain about 40% oil and 23% protein [5,6]. Oils from these vegetables cabbage, cauliflower, and broccoli (all domesticates of and other oilseed crops are valuable commodities having great ver- Brassica oleracea), and the model plant Arabidopsis (Arabidopsis satility, for example being used as food and cosmetics ingredients, thaliana) [18,19]. The mustard family is divided into three lineages cooking oils, industrial lubricants, chemical feedstocks for soaps where Arabidopsis and camelina are members of lineage I, and and detergents, and more recently as feedstocks for biodiesel and species of the Brassica genus along with pennycress are members jet fuels [1,7,8]. Meal, which is the seed material including protein of lineage II [19]. that remains after oil extraction, also serves as an important food While pennycress belongs to extended II lineage of Brassicaceae, and feed commodity. the taxonomic standing of members of the Thlaspi genus had long The demand for products derived from oilseed crops contin- been in dispute. In 1997 through the analysis of nuclear ribosomal ues to rise due to population growth and increasing globalization. internal transcribed spacer and chloroplast sequences, members In addition, a large body of evidence points toward the burning of the Thlaspi genus conspicuously clustered into a polyphylogenic of fossil fuels as causing ecosystem-damaging climate change [9], group [20]. This analysis supported the movement of nine of four- placing an impetus on lowering our carbon footprint by replacing teen so-called Thlaspi species out of the Thlaspideae tribe. Notably, liquid fossil fuels with plant oils-derived biodiesel and jet fuels. Thlaspi caerulescens, a species well studied for its bioremediation These energy-dense biofuels are essential for meeting transporta- potential [21], was moved to the genus Noccaea. While these find- tion needs given that less energy-dense biofuels such as ethanol ings were reported some time ago, some of the broader scientific derived from corn grain or cellulosic biomass cannot feasibly run literature has been slow to reflect these changes [22]. Besides pen- heavy equipment and airplanes. nycress, other members of the Thlaspideae tribe include Thlaspi In order for oilseed crop production to meet burgeoning ceratocarpum and Thlaspi alliaceum (roadside pennycress) along demands in an.