Camelina Sativa)

17 Camelina (Camelina sativa) C. Eynck1 and K.C. Falk2 1Linnaeus Plant Sciences, Inc., Saskatchewan, Canada; 2Agriculture and Agri-Food Canada, Saskatchewan, Canada 17.1 History of Camelina established in the south-eastern part of Europe Cultivation and it became a commonly grown crop in several parts of the European mainland and Camelina (Camelina sativa L. Crantz) is an Scandinavia during the Iron Age (400 BCE –CE ancient oilseed that belongs to the Brassicaceae 500) (Plessers et al., 1962; Knörzer, 1978; (mustard) family. It is also known as gold-of- Bouby, 1998). Seeds of C. alyssum were a sub- pleasure, false flax, large-seeded false flax, stantial part of the human diet, together with wild flax (UK), linseed dodder, Dutch flax, flax and other cereals (Hjelmquist, 1979). The German sesame, or Siberian oilseed (Putnam species that is now known as Camelina sativa et al., 1993; Zubr, 1997). While some of these L. Crantz probably emerged during prehistoric names hint at the plant’s resemblance to flax times from plantings of C. microcarpa and/or (Linum usitatissimum), the latter three denota- C. alyssum (Knörzer, 1978). It is also not clear tions provide an indication of the geographical how or when the event of speciation of C. sativa origin of this species. Thus, archaeological took place. records indicate that the south-eastern Europe– Genetic analyses provide an alternative south-western Asian steppe regions are most approach to address the question of the origin likely the centre of origin of camelina (Knörzer, of a species. A recent study (Ghamkhar et al., 1978; Zohary and Hopf, 2000). 2010) used AFLP fingerprinting to investigate Observations from mostly carbonized the genetic diversity in a previously not acces- seeds and capsules suggest that the evolution sible set of C. sativa accessions from the of camelina as a crop was initiated when the Russian-Ukrainian area, together with acces- wild species Camelina microcarpa and C. alys- sions from central and western Europe, China sum (synonym: linicola, macrocarpa, dentata) and Australia. In accordance with archaeologi- were domesticated in the late Neolithic in cal records, it identified the Russian-Ukrainian south-eastern Europe (Zinger, 1909; Knörzer, region as a genetic diversity hotspot and poten- 1978; Putnam et al., 1993). With the expansion tial centre of origin of the species. of agriculture, camelina very early invaded The importance of camelina as a food crop cultivated land as a weed in flax (Schwanitz, declined during the Middle Ages and it was 1967). But soon the nutritional value of the grown only sporadically until modern times oleiferous seeds was discovered and camelina (Knörzer, 1978; Hjelmquist, 1979). Aside from was grown as a crop in monoculture. In the its limited utilization as edible oil, camelina oil Bronze Age (1500–400 BCE ), camelina was well was the lamp oil of choice until the discovery of ©CAB International 2013. Biofuel Crops: Production, Physiology and Genetics (ed. B.P. Singh) 369 370 C. Eynck and K.C. Falk gas and electricity; later it was used in the pro- 1999), North America (Putnam et al., 1993) duction of soft soaps and coating material. Prior and Australia (Francis and Campbell, 2003), to the 1930s, the crop was still grown on sandy primarily because of the exceptional level of soils in France (Guillaumin, 1946), Belgium, the ω-3 essential fatty acid α-linolenic acid in Holland, the Balkan region and in Russia from its oil, favourable agronomic attributes, its Caucasus to Siberia (Wacker, 1934; Plessers potential as a low-input source of biofuels and et al., 1962; Zubr, 1997). The cultivation of as a feedstock for the oleochemical industry camelina declined further with the introduction (Fröhlich and Rice, 2005). Currently, it is being of oilseed rape. According to Crowley (1999), grown as a feedstock for biofuels in several the lower cost of hydrogenating rape oil and the northern US states (Fröhlich and Rice, 2005; lack of knowledge of potential large-scale Moser and Vaughn, 2010), such as Montana, applications for camelina oil were the major North Dakota and Washington. reasons for the lack of interest in the crop. With the colonization of the New World by European settlers, camelina was introduced for a second time into a new environment as a weed 17.2 Biology in flax (Putnam et al., 1993). In his voluminous encyclopaedia on the fauna of the central and Camelina is a herbaceous, fast-growing annual southern USA, Porcher (1863) described the with both spring and winter forms (Putnam positive agronomic attributes of camelina, prais- et al., 1993). Plants are taprooted and can attain ing its low fertilizer requirements, high yield, heights of between 30 and 110 cm (Putnam early maturity, frost and drought tolerance and et al., 1993; Francis and Warwick 2009; Berti resistance to certain insect pests. In the middle of et al., 2011). Initially, a rosette is formed; the 20th century, a 30-year research programme winter-type camelina as well as spring-type was started at the University of Minnesota, whose camelina sown in the autumn in climates with results added significantly to the body of knowl- mild winters remain dormant in this stage over edge on cultivation and potential uses of winter (Crowley, 1999). In spring or otherwise camelina in the USA (Robinson, 1987). The 1 week to 10 days after a rosette is established, results of this comprehensive study showed that a single erect, either smooth or sparsely haired, camelina was similar to flax, rape, mustard and stem is initiated (Putnam et al., 1993; Francis canola in maturity, yield and requirements for and Warwick, 2009). Depending on the geno- harvesting machinery and that its cultivation was type, stems are more or less heavily branched accompanied with lower production costs than and become woody as they mature. Leaves any of the other oilseed crops. Trials conducted are arrow- to lance-shaped with smooth or in Canada (Plessers et al., 1962) obtained results lobed edges. They are typically 2–8 cm long similar to those in the USA and led to the sugges- and 2–10 mm wide and can be either smooth tion that camelina should be given serious con- or feature a few, primarily dichotomic, hairs sideration as a potential oilseed crop for northern (Putnam et al., 1993; Francis and Warwick, latitudes. 2009). Lacking a petiole, the leaves are usually However, despite numerous favourable clasping and arranged in an alternate manner. accounts of camelina as a low-input oilseed From 4 to 6 weeks after germination, crop, until recently camelina has not been cul- depending on the prevailing weather condi- tivated except for small isolated plantings tions, camelina plants start to flower, with a (Gugel and Falk, 2006). Similar to the situation flowering period of about 2 weeks (Akk and in Europe, the lack of a market for the oil has Ilumäe, 2005). Inflorescences are arranged as hindered the establishment of camelina as a racemes with the flowers in terminal clusters crop in North America. However, with an (Francis and Warwick, 2009). Flowers are small increasing demand for vegetable oils and rising and pale yellow to greenish yellow in colour with petroleum prices this may change. Camelina a diameter of 5–7 mm (Fig.17.1). The organiza- has recently gained renewed interest as an tion of the flower is similar to that of other oilseed crop in Europe (Zubr, 1997; Hebard, cruciferous crop plants but features delicate, 1998; Leonard, 1998; Bonjean and Le Goffic, lance-shaped petals. Camelina is primarily Camelina (Camelina sativa) 371 Fig. 17.1. (a) Raceme, (b) flower and (c) seed pod of C. sativa (photo courtesy Prakash Venglat, Genome Prairie). self-pollinating (Plessers et al., 1962; Zubr, Plessers et al., 1962; Zubr, 1997; Schuster and 1997; Mulligan, 2002). To the best of our Friedt, 1998; Gugel and Falk, 2006; Vollmann knowledge the only study that investigated the et al., 2007). rate of out-crossing in camelina is a field study Seed oil contents of 300–460 g kg−1 seed conducted in Alberta, reporting rates of at max- have been reported (Robinson, 1987; Putnam imum 0.28% at 20 cm distance (Walsh et al., et al., 1993; Budin et al., 1995; Angelini et al., 2012). However, cross-pollination mediated 1997; Zubr and Matthäus, 2002; Zubr, 2003; by insects, i.e. bees, has been suggested by Gugel and Falk, 2006; Vollmann et al., 2007) several authors (Blamey and Grey-Wilson, and a considerable environmental variation in 1989; Corbet et al., 1991). oil content and its composition has repeatedly The fruits of camelina are smooth pear- been demonstrated (Zubr, 2003; Vollmann shaped siliques 7– 9 mm in length. At first sight, et al., 2007). In northerly locations the oil con- they resemble the seed capsules of flax, a char- tent of camelina is usually higher than in more acter that led to denotations such as false flax southerly locations (Plessers et al., 1962; Gugel or wild flax in the European vernacular. Siliques and Falk, 2006), which is probably due to the usually contain 5–15 golden, brown or reddish fact that the predominantly cool and moist brown seeds; however, up to 25 seeds per sil- weather in northern areas is conducive to ique have been reported (Schuster and Friedt, proper seed filling and maturation. 1998). Seeds are oblong-oval and rough, having Camelina oil is largely composed of a deeply ridged surface (Putnam et al., 1993; unsaturated fatty acids with 25–42% α-linolenic Francis and Warwick, 2009). They are small acid (C18:3), 13–21% linoleic acid (C18:2), with 1000-seed weights averaging about 1.0 g 14–20% oleic acid (C18:1), 12–18% gondoic and ranging from 0.3 to 2.3 g, depending on (eicosenoic) acid (C20:1) and 2–4% erucic genotype and growing conditions during seed acid (C22:1) (Budin et al., 1995; Zubr and development (Gugel, personal communica tion; Matthäus, 2002; Vollmann et al., 2007).

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