The Biology of Canadian Weeds. 142. Camelina alyssum (Mill.) Thell.; C. microcarpa Andrz. ex DC.; C. sativa (L.) Crantz.

Abstract

Can. J. Plant Sci. 89: 791-810. This paper summarizes biological information on three cruciferous weed species: Camelina alyssum, C. microcarpa and C. sativa. Camelina sativa, which had been gathered or cultivated as an oil seed for many centuries in Europe, was the first to reach North America as a weed, towards the mid-19th century, gradually spreading across the prairies, mainly in crops, to British Columbia and the Northwest Territory. The most common of the three species in North America, C. microcarpa, arrived in the late 19th century, and subsequently appeared at numerous crop and uncultivated sites across the country, probably in cargo as the railways expanded. Camelina alyssum appeared in the early 20th century at restricted sites on the prairies, mostly in Saskatchewan. All three species have diminished in importance as crop weeds in western Canada over the past few decades. This reduction could be related to increased weed control by herbicides. Herbicide-resistant biotypes have recently been reported in C. microcarpa. Camelina sativa has attracted renewed interest as an oil crop, because of an adaptation to various climatic conditions, low nutrient requirements and resistance to disease and pests. In Europe, where it is now widely grown, it has shown considerable potential in the food, animal feed, nutraceutical, paint, dye, cosmetic and biofuel industries. In North America, it is being grown on a trial basis mainly for its potential as a biofuel in Alberta, Saskatchewan, the Maritime Provinces, and the northern United States of America.

Full text

The Biology of Canadian Weeds. 142.
Camelina
alyssum
(Mill.) Thell.;
C. microcarpa
Andrz. ex DC.;
C. sativa
(L.) Crantz.
A. Francis and S. I. Warwick
Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, K.W. Neatby Bldg., C.E.F.,
Ottawa, Ontario, Canada K1A 0C6 (e-mail: Suzanne.Warwick@.agr.gc.ca)
.
Received 22 October 2008,
accepted 10 March 2009.
Francis, A. and Warwick, S. I. 2009. The Biology of Canadian Weeds. 142. Camelina alyssum (Mill.) Thell.; C. microcarpa
Andrz. ex DC.; C. sativa (L.) Crantz. Can. J. Plant Sci. 89: 791810. This paper summarizes biological information on three
cruciferous weed species: Camelina alyssum, C. microcarpa and C. sativa. Camelina sativa, which had been gathered or
cultivated as an oil seed for many centuries in Europe, was the first to reach North America as a weed, towards the mid-
19th century, gradually spreading across the prairies, mainly in crops, to British Columbia and the Northwest Territory.
The most common of the three species in North America, C. microcarpa, arrived in the late 19th century, and subsequently
appeared at numerous crop and uncultivated sites across the country, probably in cargo as the railways expanded.
Camelina alyssum appeared in the early 20th century at restricted sites on the prairies, mostly in Saskatchewan. All three
species have diminished in importance as crop weeds in western Canada over the past few decades. This reduction could be
related to increased weed control by herbicides. Herbicide-resistant biotypes have recently been reported in C. microcarpa.
Camelina sativa has attracted renewed interest as an oil crop, because of an adaptation to various climatic conditions, low
nutrient requirements and resistance to disease and pests. In Europe, where it is now widely grown, it has shown
considerable potential in the food, animal feed, nutraceutical, paint, dye, cosmetic and biofuel industries. In North
America, it is being grown on a trial basis mainly for its potential as a biofuel in Alberta, Saskatchewan, the Maritime
Provinces, and the northern United States of America.
Key words: Flat-seeded false flax, small-seeded false flax, large-seeded false flax, came
´
line alysson, came
´
line a
`
petits fruits,
came
´
line cultive
´
e, weed biology
Francis, A. et Warwick, S. I. 2009. La biologie des mauvaises herbes au Canada. 142. Camelina alyssum (Mill.) Thell.; C.
microcarpa Andrz. ex DC.; C. sativa (L.) Crantz. Can. J. Plant Sci. 89: 791810. Cet article dresse le bilan de l’information
connue sur la biologie de trois adventices du groupe des crucife
`
res : Camelina alyssum, C. microcarpa et C. sativa. Camelina
sativa qu’on utilise ou cultive pour son huile depuis plusieurs sie
`
cles en Europe, a e
´
te
´
la premie
`
re a
`
atteindre l’Ame
´
rique du
Nord comme mauvaise herbe, vers le milieu du XIX
e
sie
`
cle. Elle s’est ensuite e
´
tendue peu a
`
peu aux Prairies, surtout dans les
cultures, pour parvenir en Colombie-Britannique et jusque dans les Territoires du Nord-Ouest. C. microcarpa, la plus
commune des trois espe
`
ces en sol nord-ame
´
ricain, a e
´
te
´
introduite vers la fin du XIX
e
sie
`
cle avant d’apparaı
ˆ
tre dans de
multiples cultures et terrains incultes du continent, sans doute avec l’expansion du transport des marchandises et du trafic
ferroviaire. Camelina alyssum est survenue au de
´
but du XX
e
sie
`
cle a
`
quelques endroits des Prairies, principalement en
Saskatchewan. Les trois espe
`
ces ont perdu de l’importance en tant que mauvaises herbes dans l’ouest du Canada ces dernie
`
res
de
´
cennies. On le doit peut-e
ˆ
tre a
`
l’usage accru des herbicides pour lutter contre les adventices. Des biotypes re
´
sistants aux
herbicides ont re
´
cemment e
´
te
´
signale
´
s pour C. microcarpa. Camelina sativa a suscite
´
un regain d’inte
´
re
ˆ
t comme ole
´
agineux, en
raison de sa facilite
´
d’adaptation aux conditions climatiques, de ses faibles exigences nutritives et de sa re
´
sistance a
`
la maladie
et aux ravageurs. En Europe, ou
`
on la cultive couramment, cette plante pre
´
sente un potentiel conside
´
rable comme culture
vivrie
`
re ou fourrage
`
re, comme nutraceutique, ainsi que pour les industries de la peinture, des colorants, des cosme
´
tiques et des
biocarburants. En Ame
´
rique du Nord, on la cultive expe
´
rimentalement en Alberta, en Saskatchewan, dans les Maritimes et
dans le nord des E
´
.-U., principalement en raison de son usage e
´
ventuel comme biocarburant.
Mots cle
´
s: Came
´
line alysson, came
´
line a
`
petits fruits, came
´
line faux-lin, flat-seeded false flax, small-seeded small flax, large-seeded
small flax, biologie des mauvaises herbes
1. Name
I. Camelina alyssum (Mill.) Thell. * Synonyms: Came-
lina parodii Ibarra & Laporte; C. dentata Pers., C.
linicola K. F. Schimp. & Spenn.; flat-seeded false flax
(Darbyshire et al. 2000), came´ line alysson (Darbyshire
et al. 2000), came
´
line a
`
graines plates (Darbyshire 2003).
European and Mediterranean (EPPO) code (Bayer
code): CMAAL.
II. Camelina microcarpa Andrz. ex DC. * Synonyms: C.
bornmuelleriana Hub.-Mor. & Reese; C. pilosa (DC.) N.
W. Zinger; C. sylvestris Wallr.; small-seeded false flax
791
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

(Darbyshire et al. 2000), little-podded false flax, small-
seed falseflax, western flax, hairy gold-of-pleasure (UK)
(Darbyshire 2003); came´ line a` petits fruits (Darbyshire
et al. 2000), came
´
line a
`
petites graines (Darbyshire
2003). EPPO code: CMAMI.
III. Camelina sativa (L.) Crantz * Synonyms: Myagrum
sativum L.; Camelina caucasica (Sinskaya) Vassilcz.; C.
glabrata (DC.) Fritsch ex N.W. Zinger; large-seeded
false flax (Darbyshire et al. 2000), Dutch flax, false flax,
gold-of-pleasure, largeweed falseflax, western false flax,
wild flax (UK) (Darbyshire 2003); came´ line cultive´ e
(Darbyshire et al. 2000), came
´
line a
`
graines mucilagi-
neuses, came
´
line de l’Ouest, came
´
line faux-lin, faux lin,
faux lin de l’Ouest, lin ba
ˆ
tard, moutarde, petit lin,
se
´
same ba
ˆ
tard, se
´
same d’Allemagne (Darbyshire 2003).
EPPO code: CMASA.
The three species are members of the mustard family,
Brassicaceae (Cruciferae), Brassicace
´
es (Crucife
`
res).
The genus Camelina consists of 11 species (Warwick et
al. 2006), of which four have been introduced to North
America: C. alyssum, C. microcarpa, C. rumelica Velen.
(United States only; and will not be discussed further in
this Canadian account) and C. sativa (McGregor 1984;
Frankton and Mulligan 1987; Rollins 1993). Camelina
means ‘‘little flax’’ and sativus means ‘‘planted, culti-
vated, sown not wild’’ (Gledhill 1990). The common
name ‘‘false flax’’ apparently arose in early Europe
where these species were common weeds of flax (Linum
usitatissimum L.), grown for centuries as an oil and fibre
crop, and were harvested along with the flax for some of
the same domestic uses (see Section 6). Although no
longer the case in North America, Camelina spp. are still
frequently present as weeds or intercropped with flax in
Europe (e.g., see Sections 3b, 10).
2. Description and Account of Variation
(a) Species Description * The following descriptions are
based on information taken from the taxonomic litera-
ture (Grubert 1980; Schultze-Motel 1986; Frankton and
Mulligan 1987; Blamey and Grey-Wilson 1989; Rich
1991; Alex 1992; Akeroyd 1993; Davis 1993; Rollins
1993; Douglas and Meidinger 1998; Callihan et al. 2000;
Cheo et al. 2001) and from herbarium specimens. Unless
indicated, the description applies to all three species.
Unusual extremes are in parentheses.
Annual or winter annual herbs from taproot. Stems
simple to branched above,9erect. Basal leaves not
lobed, withered at flowering; cauline leaves alternate,
sessile, lanceolate, 28 cm long210 mm wide, aur-
iculate or sagittate at base, usually clasping, with simple
to star-shaped hairs. Inflorescences ebracteate racemes;
flowers small, in terminal clusters. Petals 4, light yellow,
spatulate, 45 mm long; sepals erect, median nectaries
absent. Stamens 6, in three pairs of unequal length;
ovules 825 per ovary. Fruit pods glabrous, more or less
rounded inflated siliques with a membranous partition
as wide as the pod and a persistent style, valves convex,
narrowly winged, on slender spreading pedicels, tardily
dehiscent. Seeds mucilaginous when wetted. Cotyledons
not notched, incumbent. Camelina alyssum stems slen-
der, sparingly branched, 1050 (70) cm tall,9glab-
rous; C. microcarpa stems occasionally branched, 30
100 cm tall, hairy with simple, rarely forked hairs above,
smaller star-shaped hairs below, sometimes glabrous
above; C. sativa stems single, usually branched above,
3060 cm tall,9glabrous, sometimes with a few simple
and branched hairs. Camelina alyssum leaves9glabrous,
deeply toothed or lobed, rarely subentire, apex broadly
to narrowly tapered; C. microcarpa leaves entire, rarely
subentire, hairy with simple, forked or star-shaped hairs,
apex tapered, rosette leaves of winter annual morph
tightly bunched; C. sativa leaves glabrous or sparsely
hairy with primarily forked hairs, entire or shallowly
toothed, apex acute. Camelina alyssum fruiting racemes
rather short and lax; C. microcarpa and C. sativa fruiting
racemes elongated on ascending pedicels. Camelina
alyssum pods depressed globose, with a flattened apex,
not hard or woody, 610 mm long; C. microcarpa pods
woody, pear-shaped, obtuse at apex, 57(8) mm long;
C. sativa pods leathery, pear-shaped to oblong, dis-
tinctly longer than wide, apex rounded, 79 mm long.
Camelina alyssum seeds reddish, rounded, flattened,
1.52.5 mm long; C. microcarpa seeds oblong, com-
pressed, warty, reddish- to orange-brown, slightly longer
than wide, 0.81.4 (1.5) mm long; C. sativa seeds
oblong, brown, deeply grooved, 23 mm long.
Chromosome counts from Canada include 2n40 for
C. alyssum from Alberta (Moss 1983); n20 for C.
microcarpa from Saskatchewan; 2n40 for C. micro-
carpa from Alberta and British Columbia; and 2n40
for C. sativa from Alberta and British Columbia
(Mulligan 1957, 1984, 2002a, b). Reports from the
United States include 2n40 for C. alyssum from the
northern Great Plains Region and n8 for C. micro-
carpa from Ohio (Easterly 1963). Other counts for C.
alyssum were 2n40 from Sweden and Hungary; for C.
microcarpa were n 13 from France and Morocco,
n19 from the Czech Republic, and 2n16, 1820,
32 and 40 from Russia, 2n 26 from France, Spain and
Morocco, and 2n40 from the Czech Republic, Ice-
land, Poland and Sweden and Iran; and for C. sativa
chromosome counts include 2n12, n14, n20, and
2n26 from France, Spain, China, and Bulgaria,
respectively. Other counts, from Argentina, Scotland,
Sweden, Iceland, and Poland indicate 2n40 (reviewed
in Warwick and Al-Shehbaz 2006). Although 2n 40 is
the most common count, the above deviations in
chromosome number would appear to be due to natural
variability among populations (diploid, tetraploid and
aneuploid numbers; Jalas et al. 1996).
(b) Distinguishing Features * The three Camelina species
can usually be distinguished from other members of the
mustard family with small rounded pods by their light
yellow flowers, their glabrous more or less tear-shaped
792 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

pods with a broad septum as wide as the pod, and their
somewhat arrow-shaped clasping stem leaves, usually
with distinctive branching or star-shaped hairs (Alex
1992). Camelina species are most likely to be confused
with hoary alyssum [Berteroa incana (L.) DC.] because of
their similar size, hairy leaves and stems, small flowers,
small ascending beaked seedpods and small seeds.
However, the seedpods of the latter are elliptic in shape,
hairy rather than glabrous, and are adpressed close to the
stem; the basal leaves are also long-stalked, the stem
leaves are elliptic, greyish from the density of hairs, and
non-clasping; and, the petals are white and deeply
notched (Frankton and Mulligan 1987; Warwick and
Francis 2006).
(c) Intra-specific Variation * Two subspecies are
currently recognized for Camelina alyssum: subsp.
alyssum (Mill.) Thell. and subsp. integerrima (C
ˇ
elak.)
Smejkal, of which only subsp. alyssum has been
recognized in Canada (Warwick et al. 1999). No
subspecies are currently recognized for C. microcarpa
and C. sativa (Warwick et al. 2006).
(d) Illustrations * The three Camelina species are
illustrated in Fig. 1, based on Schultze-Motel (1986);
Frankton and Mulligan (1987); Alex (1992); Cheo et al.
2003; and Douglas and Meidinger (1998).
3. Economic Importance
(a) Detrimental * Camelina microcarpa is currently the
most common and widespread of the three species in
North America (see Figs. 2 and 3; USDA, NRCS 2008),
and is the only one to have evolved herbicide resistant
biotypes (see Section 11). It is a common weed of both
cultivated and abandoned fields, and is often a con-
taminant of livestock feed (Royer and Dickinson 1999).
All three species were considered weedy in the past,
particularly in the prairies where they were described as
weeds of disturbed soils in grainfields, along roadsides
and in waste places, with C. microcarpa more common
than C. sativa, and C. alyssum relatively rare (Scoggan
1957; Boivin 1968; Moss 1983; Looman and Best 1987).
They appear to have first become naturalized as weeds
in Manitoba (Scoggan 1957), eventually spreading into
crop areas in all the Canadian prairies and beyond the
United States border; but were never listed among the
top ranking weeds based on weed surveys conducted
over a 30-yr period from the 1970s (Leeson et al. 2005).
These surveys showed that the species (not treated
individually) have continued to be present in some
crops, mainly in Manitoba and Saskatchewan, but their
overall ranking in order of abundance declined from an
average of 100 out of 182 to 136 out of 148. By the
2000s, field peas are the only crop listed as affected by
Camelina spp. The species were reported in spring wheat
up to the 1990s, but were not reported from barley, oats
and flax after the 1970s. Their presence in the Aspen
Parklands region appears to have been largely confined
to Manitoba and Saskatchewan, but their presence in
spring wheat in the Moist Mixed Grassland region may
have included Alberta. The reduced occurrence of
Camelina species may reflect the increased efficacy of
herbicides used to control broadleaf weeds in grain and
other crops.
In Great Britain, C. sativa was formerly a weed of
spring-sown cereal, flax and lucerne fields (Clapham et
al. 1987). Although now less common with improved
cleaning of seed in those crops, it is frequent in gardens
and waste places; C. microcarpa, formerly a weed of
autumn-sown crops, is now rare; and C. alyssum,
formerly reported as rare in waste places, has not been
reported for several years (Clapham et al. 1987).
In German field trials, dry weight of flax plants grown
with C. alyssum was 46.267.4% of that of flax grown
alone; stem length of flax was reduced more by C.
alyssum than by the other six weed species, and growth
of cotyledonary shoots was strongly inhibited (Balschun
and Jacob 1975).
The presence of glucosinolates, sinapine, and phytic
acid in C. sativa seed can limit its use in animal feed,
causing poor bioavailability of minerals and proteins in
the case of phytic acid, and sometimes tainting the
flavour of the meat in the case of sinapine (Mattha
¨
us
1997; see also Section 7c). In German pig feeding trials,
depression of growth rate, deterioration of consistency
of carcass fat, and a decrease in meat palatability
occurred when the percentage of C. sativa expeller
(pressed oilseed residues) exceeded 5% of the pigs’ diet
(Bo
¨
hme et al. 1997), while Flachowsky et al. (1997)
found that feeding C. sativa expellers to pigs resulted in
a decrease in the induction time of fat development and
destruction of the antioxidant vitamin E. Ryha
¨
nen et al.
(2007) found that while C. sativa seed cake added to the
diet of chickens could enhance omega-3 fatty acids in
the meat without affecting the flavour, the overall effects
were negative, because of weight loss, depressed feed
intake and a lower feed conversion level.
In Australian studies, Lovett and Duffield (1981)
reported that allelochemicals found in aqueous washings
of the leaves of Camelina species were the result of
interactions between various acids normally present in
the leaves and the activities of a free-living bacterium
(Enterobacter cloacae) in the phyllosphere. Bacteria
attracted to these host acids led to the production of
secondary chemical compounds, among them benzyla-
mine, identified as the allelochemical influencing the
chemical interaction at the root level of C. sativa with flax.
(b) Beneficial * In medieval Europe the oil was
apparently used for culinary purposes or lighting, while
stems were used in broom construction (Hedrick 1919;
Plessers et al. 1962; Kno
¨
rzer 1978; Hjelmqvist 1979;
Facciola 1990). Camelina seed was also used to feed
livestock in Germany and France (Marie-Victorin 1964).
In recent years, this species has received renewed
attention as an oil crop. Proposed uses of the oil include
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 793
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

food and nutraceutical uses, and in paints, cosmetics and
biofuels (Putnam et al. 1993; Vollmann et al. 1996;
Angelini et al. 1997; Zubr 1997; Bonjean and Le Goffic
1999; Anonymous 2002; Jankowski and Budzynski 2003;
Mlakar et al. 2003; Gugel and Falk 2006; Dubois et al.
2007; Lu 2008).
Early tests in Ottawa, ON, and Fort Vermilion, AB,
indicated its potential as an oil crop in northern areas
because of its early ripening, high yield, oil content, and
resistance to insect damage (Plessers et al. 1962).
Downey (197l) later suggested that although the meal
was suitable for animal feed, the linolenic acid content in
the oil was too high for the edible oil market and too low
to compete with industrial linseed oil from flax. Recent
trials in Alberta and Saskatchewan (Gugel and Falk
2006) have shown that prospects for developing im-
proved C. sativa germplasm suitable for western Cana-
dian environments are good. Increased seed size and oil
content are characteristics of primary importance, but
more research is needed on growing conditions, and in
Fig. 1. Camelina alyssum. A. stem, leaves and fruiting stems; B. seedpod (Thellung 1919); Camelina microcarpa. C. flowering and
fruiting stem; D. lower stem and root; E. seedpod; and F. seeds (Frankton and Mulligan 1987); Camelina sativa. G. Flowering and
fruiting stem; H. seed pod (Douglas and Meidinger 1998); and I. seeds (Frankton and Mulligan 1987). Seeds of C. alyssum are not
illustrated  see text Section 2 for description. Note species variability in plant size in the description.
794 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

developing markets for oil and meal. Urbaniak et al.
(2008a, b) have recently evaluated the potential of
C. sativa as a high-value oilseed crop under the moist
cool conditions of the Maritime Provinces, and con-
cluded that it can be successfully grown there due to its
adaptability to seeder type (seed drill and forage seeder),
low seeding rate requirements and a wide seeding date
window. In Montana, C. sativa was considered the most
economical crop to produce due to minimal input
requirements (Anonymous 2007; Pilgeram et al. 2007),
with production area reaching over 4000 ha in 2006 and
estimated at 20 250 ha in 2007. It has been found to be
similar to other oilseeds in oil content in tests in Sweden,
Denmark, and the former Soviet Union (Zubr 1997;
Zubr and Mattha
¨
us 2002; see Section 7c).
Current interest in North America is mainly in its
potential as a biofuel additive or for biodiesel produc-
tion in western Canada (Reaney et al. 2006), in Montana
and the Northern Great Plains (Anonymous 2007;
Johnson 2007; Pilgeram et al. 2007), and also in Color-
ado (El Paso County Conservation District 2008). In
Ireland, C. sativa oil evaluated as an undiluted vehicle
Fig. 2. Distribution in Canada of A. Camelina alyssum, based on 14 herbarium specimens from ALTA, DAO, SASK, and WIN; B.
C. microcarpa, based on 241 specimens from ALTA, DAO, NFLD, PMAE, SASK, UAC, UBC, V, WIN; C. C. sativa, based on 92
specimens from ALTA, DAO, PMAE, SASK, UBC, V, and WIN. Information from distribution maps along the St. Lawrence
River (Environment Canada 2002) has been added to the maps for C. microcarpa and C. sativa. Herbarium abbreviations in
Holmgren et al. (1990).
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 795
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

fuel was comparable to other biodiesel fuels in fuel
economy; but the camelina ester had higher than
standard iodine levels and potential problems under
low temperature conditions (Fro
¨
hlich and Rice 2005). In
European fuel tests, cold-pressed and filtered camelina
seed oil increased power output and reduced smoke
Fig. 3. Distribution of A. Camelina alyssum B. C. microcarpa and C. C. sativa in the United States, based on USDA, NRCS (2008).
796 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

emissions in diesel transport engines as compared with
mineral fuel (Bernardo et al. 2003). The C. sativa oil also
resulted in slightly higher fuel consumption and O
2
and
NO emissions (Bernardo et al. 2003).
In Minnesota, it was among vegetable oils showing
promise as a replacement for petroleum oil adjuvants in
herbicide sprays (Robinson and Nelson 1975). Camelina
oil can be transesterified with oleyl alcohol to prepare
long-chain wax esters suitable for applications in
cosmetics and lubricants (Steinke et al. 2000, 2001).
The oil also has potential in the health industry, based
on traditional folk remedy uses in treatment of stomach
ulcers, burns, wounds, eye inflammations and as a tonic
(Rode 2002); and some protective effects of phytic acid
and its products, such as decreasing colon cancer risk,
have been reported (Mattha
¨
us 1997).
The high percentage of unsaturated fatty acids in the
oil, particularly of linolenic acid (omega-3) (see Section
7c), suggests potential use as a food additive (Pilgeram
et al. 2007). In Finland, camelina oil was found to
significantly increase the proportion of a-linolenic acid
in the serum lipids of individuals with high cholesterol,
its cholesterol-lowering effect being comparable to that
of canola (rapeseed) (Brassica napus L.) and olive (Olea
europea L.) oils (Karvonen et al. 2002). Camelina oil is
particularly well-suited for use in cold dishes, as its
storage stability and resistance to oxidation at higher
temperatures are lower compared with other edible oils
(Mattha
¨
us 2004). Camelina sativa oil was found to be
more stable than fish and linseed oils in a study in
Ireland (Nı
´
Eidhin et al. 2003a). Recent experiments in
Slovenia to increase the oxidative stability of the oil
have shown promise in retarding the process of auto-
oxidation by as much as 60% with use of rosemary
extract (Abramovic
ˇ
and Abram 2006). The addition of
phenolic compounds extracted from camelina oil
was found to enhance the oxidative stability of commer-
cially produced safflower oil, Carthamus tinctorius L.
(Abramovic
ˇ
et al. 2007).
Beneficial effects of the oil have also been reported for
animals and their products. In Germany, camelina oil
lowered serum cholesterol and triglyceride levels in
experimental animals (Mattha
¨
us 1997). In Finland, oil
added to chicken feed increased the omega-3 content in
the eggs without the unpleasant flavour often found
with the use of flax oil (Rokka et al. 2002). In a
controlled pig-feeding trial in Ireland, camelina oil
increased plasma omega-3 fatty acids and reduced
plasma omega-6 fatty acids and serum triglyceride levels
(Nı
´
Eidhin et al. 2003b).
Camelina sativa meal has a high protein and energy
content suitable for feeding pigs and ruminants (Schus-
ter and Friedt 1998; Mattha
¨
us and Zubr 2000). Came-
lina meal at 0.7 g 100 g
1
cooked pork meat was an
effective antioxidant of protein and lipids (Salminen
et al. 2006). In France, camelina seed and meal in the
diets of dairy cows led to a decrease in milk fat yield and
softer, more spreadable butter, due to the unsaturated
fatty acids in the camelina feed (Hurtaud and Peyraud
2007). Similarly, in Italian studies, C. sativa seed
supplements at high levels, i.e., 1015% of diet,
significantly increased the percentage of polyunsatu-
rated fatty acids in rabbit meat (Peiretti et al. 2007). This
improved the quality of the meat for human nutritional
needs, reducing saturation, atherogenic and thrombo-
genic indexes (Peiretti et al. 2007). In the United States,
camelina meal was found to be a potential feed
ingredient in turkey poultry starter diets if kept below
5% of the feed, as amounts above that figure led to
decreases in weight and poorer feed conversion (Frame
et al. 2007).
Camelina sativa can also be used for forage if rations
are monitored to limit quantities (Schuster and Friedt
1998; Mattha
¨
us and Zubr 2000). Peiretti and Meineri
(2007) concluded that C. sativa provided good forage
when harvested at a stage before the flowering period,
but that there was a substantial decrease in nutritional
quality afterwards. In Russia, Camelina spp. were
among plants considered for use in fodder mixtures
for domestic animals (Artemov and Velibekova 2003).
In a laboratory study, the presence of free-living
nitrogen-fixing bacteria in the phyllospere of C. sativa
was reported to stimulate the growth of radicles of
germinating flax seed, but only at low concentrations
(Lovett and Sagar 1978; Lovett and Jackson 1980). In
Romania, Toncea et al. (2006) suggested intercropping
C. sativa with flax, as flax growth and development is
not affected by C. sativa, and they might interact to
control weeds. When grown in mixed cropping with peas
in Germany, C. sativa had a significant suppressive
effect as a smother crop and weed antagonist on weed
coverage compared with monocropped peas, particu-
larly in the establishment phase (Saucke and Ackermann
2006). The C. sativa crop can be used for green manure
(Anonymous 2002).
Camelina sativa is among arable weeds used in seed
mixtures for wildflower strips introduced to increase
biodiversity in intensively used arable land in Central
Europe (Kollmann and Bassin 2001).
(c) Legislation * Camelina species are classed as second-
ary noxious weed seeds in Class 3 of Schedule 1 of the
Weed Seeds Order, 2005 (Canadian Government 2005)
and C. microcarpa is on a list of noxious weeds in
Saskatchewan under the Noxious Weeds Act, the Nox-
ious Weeds Designation Regulations, (Chapter N-9.1
Reg. 2) as amended by Saskatchewan Regulations 14/99
(Saskatchewan Government 1999). Camelina species do
not appear on any other provincial noxious or invasive
weed lists in Canada. In the United States, Camelina
species do not appear in any federal or state legislation on
invasive or noxious weeds (USDA, NRCS 2008).
4. Geographical Distribution
Native to Eurasia, the distribution of the three intro-
duced species in Canada and the United States is
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 797
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

mapped in Fig. 2 and 3, respectively. Camelina alyssum
has a limited distribution in the three Prairie Provinces
(Fig. 2A). Camelina microcarpa is found in all provinces
and Yukon Territory (Fig. 2B). Camelina sativa is
reported from all provinces except Newfoundland (see
Meades et al. 2000), but occurs mainly in the four
western provinces, with a limited presence in Ontario,
Quebec, the Maritimes and the Northwest Territory
(Fig. 2C), with a recent report for the Yukon Territory
(not shown, see Cody et al. 2004). In the United States,
C. alyssum has a limited distribution in the states of
North Dakota and Minnesota (Fig. 3A); C. microcarpa
is listed as weedy in 48 mainland states, absent only
from Alabama and Florida (Fig. 3B); and C. sativa is
found in 38 mostly northern states, including Alaska
(not shown), but is absent from California, Nevada,
Colorado, Texas and southeastern states from Arkansas
to Florida (Fig. 3C).
The three species have a distribution extending from
temperate Europe to central Asia and China, north into
Britain and Scandinavia and south into the Himalayas.
They have also been introduced in New Zealand,
southern Australia, and Chile (Hulte
´
n and Fries 1986).
Camelina alyssum is now rare in much of western
Europe but remains well established in parts of central
and eastern Europe. Camelina microcarpa is the most
widely established species in Europe, extending into
Spain, and is particularly common from eastern France
through central Europe to Russia. Camelina sativa is
common in the northern and eastern part of its range,
extending into northern Italy, but rarer in Spain and
around the Mediterranean (Jalas et al. 1996). All three
were reported as established in the Caucasus and
Central Asia (Dorofeev 1996). Camelina microcarpa
has recently been spreading in China (Li et al. 2006).
5. Habitat
(a) Climatic Requirements * Camelina spp. are best
adapted to cool temperate semi-arid climate zones on
dry prairies or steppes, but C. microcarpa in particular
may be found at higher altitudes in warmer zones as, for
example, in the United States and Spain. The three
species are rarely present in Mediterranean or coastal
climates, with the exception of the North Sea periphery
in Europe (Jalas et al. 1996). The three species can
tolerate dry soils and low rainfall, and, because of early
ripening, can grow to maturity before frost in regions
with a short growing season, maturing 21 d earlier than
flaxseed, for example (Shukla et al. 2002). Camelina
sativa can survive early-season water deficits and minor
frosts in spring (Seedtec  Terramax 2008). In northern
France, Bonjean and Le Goffic (1999) found that winter
annual plants of C. sativa could tolerate temperatures of
10 to 148C over a period of several days without
snow cover.
(b) Substratum * Camelina microcarpa is found on
sand, loam, gravel and alkaline soils, and C. sativa on
dry sandy soils (Warwick et al. 1999). Camelina sativa
grows best on sandy, loamy and heavy clay soils that are
dry or moist, but well-drained. It is also adapted to
nutritionally poor soils and tolerates a wide pH range
(Plants For A Future 2004). In the Czech Republic, C.
sativa was among potential industrial and energy crops
that could be successfully grown on soils reclaimed from
mine spoil and fly ash (Petrikova et al. 1996).
(c) Communities in Which the Species Occur * In
Canada, C. alyssum has been found in prairie habitats
and as an urban weed along roadsides. Camelina
microcarpa is more widespread, in grain, flax, corn
and hayfields, pastures, grasslands, rangelands, gardens,
on low slopes, flats and sloughs, in ballast, and along
roadsides, railways, wharves and waste places generally.
Camelina sativa also occurs in many of the same
habitats, but is also reported in alfalfa fields,
open woods, lakeshores, and around grain elevators
(Warwick et al. 1999). Migration of C. microcarpa from
disturbed areas to xerothermic grassland habitats over a
period of several decades was detected in a study in
Slovakia (Elia
´
s 2003).
6. History
The species of Camelina are believed to have originated
in the steppes of southeastern Europe and southwestern
Asia, and are associated with the spread of agriculture
as weeds in flax and other crops (Kno
¨
rzer 1978). There
is archaeological evidence, mostly from carbonized
seeds, that C. sativa has been grown or gathered since
the late Bronze and Iron Ages and later under the
Roman Empire in regions around the North Sea, along
the Rhine, in eastern Europe, and at various Roman
sites, including England (Plessers et al. 1962; Willcox
1977; Kno
¨
rzer 1978; Hjelmqvist 1979; Hanelt et al. 1982;
Henriksen and Robinson 1996). Camelina sativa con-
tinued to be cultivated sporadically in Europe in
mediaeval times, and some believe that C. alyssum and
C. microcarpa evolved from it (Kno
¨
rzer 1978). Prior to
the Second World War, it was cultivated mainly on poor
soils to replace winter-killed oilseed rape (Brassica
napus) (Plessers et al. 1962), but in the past few decades
is being planted experimentally as a potential oilseed
and industrial crop in Europe and North America (see
Section 3b).
It is not known how the species were introduced to
North America, but it seems likely that they came as
contaminants in flax or other crop seeds, and perhaps
less likely in bird seed as suggested for their spread in
Great Britain (Stace 1997). Described as introduced and
occurring in fields and cultivated grounds, C. sativa was
the first of the three species to be mentioned in North
American botanical literature (Torrey and Gray 1838
1840). In Canada, the first report of C. sativa is from
Manitoba as a deliberate introduction at the Red River
settlement in 1863, with subsequent reports in 1873 and
1896 (Scoggan 1957). It subsequently spread in the
798 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

western provinces, reaching the Peace River District by
the 1930s and the Northwest Territory in the 1950s.
Camelina microcarpa was first collected in Manitoba in
1896, and subsequently appeared at numerous sites
across the country, often near railways and along
roadsides, suggesting transport in cargo. It reached
British Columbia in the 1920s, but did not begin to
spread widely there until the 1950s. The earliest report
of C. alyssum was in 1910 in Saskatchewan in fields near
Swift Current, and most herbarium specimens have been
collected around nurseries, greenhouses and fields near
research stations. In eastern Canada, both C. micro-
carpa and C. sativa have been found at scattered sites
in southern Ontario and along the St. Lawrence River
(Sabourin 1992; Environment Canada 2002); but Rous-
seau (1968) listed both species as adventive introduc-
tions in Quebec, with no information as to when they
first appeared.
7. Growth and Development
(a) Morphology * The three species do not possess
unusually distinctive morphological features.
(b) Perennation * The species of Camelina are annuals,
or occasionally winter annuals, reproducing only from
seed (Frankton and Mulligan 1987).
(c) Physiological Data * Seed oil content and percent
fatty acid (FA) data for Camelina spp. (Table 1) are
mostly for C. sativa crop accessions, whether in North
America or in Europe, with the exception of a European
report for wild C. microcarpa and C. sativa accessions
[Miller et al. (1965), as cited in Kumar and Tsunoda
(1980)]. Oil content values in Table 1 are similar to the
3442% obtained in Canadian evaluations in Ottawa,
ON, and Vermilion, AB (Plessers et al. 1962). Figures
from recent French and Danish analyses of fatty acids
(Zubr 2003a; Dubois et al. 2007) were similar to those of
the cited European studies. Generally linoleic and a-
linolenic acid percentages were very high; the latter is
high in OMEGA-3, which could reach over 50%.
Selection has also led to increased linolenic acid content
(Seehuber 1984; Zubr and Mattha
¨
us 2002; Zubr 2003b,
Vollmann et al. 2005, 2007), as has seed irradiation-
based mutagenesis (Vollmann et al. 1996, 1997; Bu
¨
ch-
senschu
¨
tz-Nothdurft et al. 1998), where erucic acid was
also reduced. The average oil content of 3344% is
comparable with that of 4045% in canola species
(Brassica napus and B. rapa L.) from western Canada
(Gulden et al. 2008) and 2545% in flax (linseed oil)
from Europe (Nykter et al. 2006).
In Italy, fatty acid composition and organic matter
digestibility of seeds and vegetative parts of C. sativa
were studied at five morphological stages from vegeta-
tive to ripe seed-pod (Peiretti and Meineri 2007).
Linolenic and palmitic acids increased with later growth
stages, a-linolenic content decreased from 560 g kg
1
at
the vegetative to 484 g kg
1
at the ripe pod stage. The
polyunsaturated fatty acid ratio (linoleic/a-linolenic)
increased from 0.18 at the vegetative stage to 0.38 at
the ripe seed-pod stage, and was 0.59 in the seed. There
were also different proportions of fatty acids in the seeds
than in the plants. Dry matter content, organic matter,
neutral and acid detergent fibre, lignin and gross energy
increased with advancing growth stage, whereas crude
protein, ash and in vitro organic matter digestibility
decreased with each stage of maturity, attributed to the
progressive translocation of the soluble cell contents
from the leaves and stems to the grains.
Several reports provided additional profiles of the
seeds. In western Canada, Gugel and Falk (2006) noted
protein content of 2732%. In Germany, Marquard and
Kuhlmann (1986) reported seed protein content between
23 and 30%, mineral content of 0.8% P, 1.0% K, 0.3%
Ca, 0.24% Mg and 0.01% Na, and tocopherol content
from 539 to 779 ppm. In a study of C. sativa cultivars
grown in 11 European and Scandinavian locations,
average oil content ranged from 39.6 to 44.1%, crude
protein averaged 43.6%, and crude fibre 12.416.8%
(Zubr 2003b). In an analysis of Camelina germplasm
collections from around the world, selected genotypes
showed seed oil contents of up to 480 g kg
1
, with
linolenic acid in the 25 to 42% range and low
concentrations of erucic acid (Vollmann et al. 2005,
2007). Camelina sativa contains the antioxidant vitamin
E (tocopherols, tocotrienols and plastochromanol), the
constituents of which are detailed in Budin et al. (1995),
Rode (2002), and Zubr and Mattha
¨
us (2002). In Polish
studies, storage of camelina products, with or without
the addition of antioxidants, led to a substantial
decrease in the amount of a-tocopherol, the most
bioactive form of vitamin E (Jaskiewicz and Sagan
2003). Heat treatment at 60708C also reduced a-
tocopherol levels by ca. 20%, but had little effect on
protein, fat, fibre, sugars and lysine; while cold seed
pressing reduced the seed fat content by ca. 50%
(Jaskiewicz et al. 2006). The antioxidant activity of C.
sativa extracts was also reported in Mattha
¨
us (2002) and
Mattha
¨
us and Angelini (2005).
The oil has an unusually high cholesterol content, ca.
45 mg 100 g
1
compared with other edible oils at 10 mg
100 g
1
(Mattha
¨
us 2004). In the United States, analysis
of the sterol composition of camelina oil showed ca. 0.54
wt% unsaponifiables, of which over 80% were des-
methylsterols; and the major sterols identified included
1884 ppm sitosterol, 893 ppm campestoerol, 393 ppm
^
5
-avenasterol, 188 ppm cholesterol, 133 ppm brassi-
casterol, and 103 ppm stigmasterol (Shukla et al. 2002).
A German study of the antinutritive compounds
tannins, glucosinolates, phytic acid and sinapine in
different oilseeds showed the following amounts in C.
sativa: 2.2 mg g
1
condensed tannins, 19 mg g
1
phytic
acid, 24 mmol g
1
glucosinolates, of which the main one
was 10-methylsulfinyldecyl, and 2.0 mg g
1
sinapine
(Mattha
¨
us 1997). In Saskatchewan, an analysis of
material from both local and other North American
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 799
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

and eastern European sources revealed that C. sativa
leaves accumulated only derivatives of the flavanol
quercetin (Onyilagha et al. 2003). In Great Britain,
Cole (1976) found no glucosinolate autolysis products in
C. sativa; but Daxenbichler et al. (1991) reported
hydrolysis products of camelinin in both C. microcarpa
and C. sativa, and of erysolin in C. sativa.Inan
Australian analysis of seed meal of wild crucifer species,
the indole glucosinolate 4-hydrooxyglucobrassicin was
not detected in the seed meal of C. sativa, a feature
which could be utilized by plant breeders to lower the
contents of glucosinolates in Brassica oilseed crops
(Sang and Salisbury 1987). Schuster and Friedt (1998)
demonstrated considerable variation (ca. 228 mmol g
1
dry seed) in the glucosinolate content of seeds of C.
sativa at a number of localities in Germany. In Austria,
Krist et al. (2006) analysed volatile compounds and
triacylglycerol (TAG) components in a comparative
study of linseed and camelina oils. Trans-2-butenal,
acetic acid, trans, trans-2,4-heptadienal and trans, trans-
3,5-octadien-2-one were the dominant volatile com-
pounds in camelina oil, and whereas linseed oils showed
a high content of TAG containing three linolenic acid
groups comprising more than 50% of the oils, whereas
camelina oil exhibited a higher content (up to 40%) of
TAG containing one eicosenoic acid moiety.
Camelina plants have been genetically modified with a
castor fatty acid hydroxylase gene in order to increase
hydroxyl fatty acid accumulation in the seeds and
provide an economic alternative to castor oil (Lu and
Kang 2008). In Scotland, C. sativa was successfully
established in vitro, and regeneration of roots, callus
and shoots was obtained using leaf explants and growth
regulators at rates over 10 shoots per explant (Tattersall
and Millam 1999). Regenerated shoots were successfully
transplanted to soil, and flowered and set seed normally.
In Poland, shoot regeneration from hypocotyl explants
taken from C. sativa seedlings germinated in vitro was
successful using different cultures; in one experiment,
concentrations of benzyladenine, 1-naphaleneacetic acid
and kinetin in the medium provided the best results
(Zandecka-Dziubak and Uczkiewicz 1999) and, in
another, abscisic acid added to the culture media led
to the largest increase in the number of shoots and
somatic embryos (Mielcarek et al. 2000). In Denmark,
experiments to introduce fungal disease resistance to
Brassica oleracea L. cultivars through protoplast fusion
between the latter and C. sativa were only partially
successful, as regenerated shoots grew vigorously in
vitro, but the hybrid shoots did not establish in soil
(Hansen 1998).
In a study in Poland of annual crucifers grown under
different light conditions, C. sativa was one of two
species that completed organogenesis before flowering
(under extended light) or before the start of seed
formation on a short day, after which the inflorescences
withered (Rostovtseva 1975).
Table 1. Oil content and fatty acid composition of seeds in Camelina spp. The figures at the top of the columns refer to the fatty acid carbon chain and the number of double bonds in the chain:
palmitic (16:0), stearic (18:0), oleic (18:1), linoleic (18:2), linolenic (18:3), eicosenoic (20:1), behenic (22:0) and erucic (22:1). Fatty acids not given specific values or not included in the table are
included in the total under ‘‘Other’’
Location % oil 16:00 18:00 18:01 18:02 18:03 20:01 22:00 22:01 Other Citation
Camelina microcarpa
Europe (not specified) 34 6 3 14 18 33 16 2 6.5 Kumar and Tsunoda (1980)
Camelina sativa
Canada (AB, SK) 3843 12.516.5 14.719.2 34.140.0 12.616.0 2.43.6 13.615.0 Gugel and Falk (2006)
Canada (NB, NS, PE) 3540 5.37.7 1.42.8 12.615.9 15.420.7 31.037.2 13.915.8 2.84.4 Urbaniak et al. (2008a, b)
USA (MI) 3038 14.119.5 18.824.0 27.034.7 12.014.9 0.04.0 11.817.4 Budin et al. (1995)
Europe (Germany) 3741 6.5 2.3 16.2 18.1 39 13.2 2.6 Marquard and Kuhlman (1986)
Europe (north, central) 15 15.2 36.8 15.5 2.8 Zubr and Mattha
¨
us (2002)
Europe (Slovenia) 33 5.7 2.6 16.4 18.7 32.9 15.8 1.5 3 0.5 Rode (2002)
Europe (not specified) 33 6 3 9 19 38 12 3 6 Kumar and Tsunoda (1980)
800 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

(d) Phenology * In Canada, C. microcarpa flowers from
May to August (Frankton and Mulligan 1987) and into
early autumn in southern Ontario, frequently dropping
seed while still flowering (Alex 1992). In North America,
C. sativa and C. alyssum generally flower from May to
June, and C. microcarpa from May to July (Rollins
1993). In Saskatchewan, C. sativa plants come to
maturity in 85100 d (Seedtec  Terramax 2008).
Camelina sativa flowers from May to July in Europe
(Polunin 1969). In Italy, plants reached the flowering
and fruiting stages 12 mo after emergence, respectively,
and late sowing led to a progressive decline in plant
height and to a shorter growth cycle, which varied from
59 to 73 d (Angelini et al. 1997). In Ireland, C. sativa
sown in late September began flowering on May 10 and
matured on July 20; sown on October 15, it began
flowering on May 20 and matured on August 02; and
sown on December 10, it began flowering on June 01
and matured on August 18; all late-sown plants
established within 2 wk and reached the 46 leaf stage
before halting growth until the following April (Crowley
1999). These small over-wintering first leaves can be
contrasted with the large, bunched rosettes described for
C. microcarpa in the United States (Callihan et al. 2000).
(e) Mycorrhiza * Mycorrhization is rare in the Brassi-
caceae (Medve 1983) and no reports were found for a
mycorrhizal association for the Camelina spp.
8. Reproduction
(a) Floral Biology * Both Camelina sativa and C.
microcarpa are considered self-compatible, autogamous
(i.e., predominantly self-fertilizing) species, whereas the
breeding system of C. alyssum is unknown (Mulligan
2002b). Tedin (1922) and Schultze-Motel (1986) pro-
vided details of the selfing process in Camelina sativa:as
the flower begins to close towards evening, the stamens
are directed towards the stigma, where they deposit the
pollen overnight. In 23 d the flower withers and falls
off. The process is repeated along the stem, which
elongates as each flower appears. Tedin also noticed
that out of 10 000 plants he studied, less than 3% were
pollinated by bees or butterflies. However, Goulson
(2003) presumably erroneously included C. sativa
among insect-pollinated crops that would benefit from
enhanced bee and other pollinator populations.
(b) Seed Production and Dispersal * Early trials in
Canada showed seed yields of 1200 to 1550 kg ha
1
(Plessers et al. 1962). Seed production in recent oilcrop
trials in the Maritime Provinces of Canada in 20052006
(Urbaniak et al. 2008b) was not significantly affected by
seeding date (two dates in May to first week of June, 2
wks apart), but was somewhat affected by seeding rate,
yielding 1338 kg ha
1
at 200 seeds m
1
, 1496 kg ha
1
at 400 seeds m
1
and 1599 kg ha
1
at 600 seeds m
1
.
Thousand-seed weights did not differ significantly at the
different rates, as more branches and capsules per plant
were produced at the lower seeding rates.
In German field trials, seed production of C. sativa
was found to differ according to seeding date and soil
enrichment. Yields of early-seeded plants averaged 1600
kg ha
1
as compared with 1150 kg ha
1
with later
seeded plants. Thousand-seed weights similarly varied
between 0.8 and 1.3 g (Marquard and Kuhlmann 1986).
Schuster and Friedt (1995) reported an average 1000-
seed weight of 1 g, and noted a seed yield of 2.53tha
1
could be reached with spring cultivars of C. sativa.In
field trials in Germany, yields averaged 1340, 1160 and
1800 kg ha
1
over a 3-yr period; maximum yield was
2280 kg ha
1
at a sowing rate of 400 seeds m
2
and N
application of 120 kg ha
1
; branch and pod number,
seeds per pod and seed weight per plant were reduced at
the highest sowing rate of 800 seeds m
2
, whereas yield
and yield components increased with increasing N rate
(Honermeier and Agegnehu 1996; Agegnehu and Hon-
ermeier 1997). In other German field trials, C. sativa
yielded on average 1.9 t ha
1
under less than optimal
growing conditions, and several C. sativa breeding lines
showed a significantly higher yield potential than
control cultivars (Mu
¨
ller et al. 1999). Gehringer et al.
(2006) similarly achieved better yields through breeding
for C. sativa crops grown on marginal, poor soil with
only moderate levels of fertilization (80 kg N ha
1
), a
maximum seed yield of 3.0 t ha
1
having been achieved.
In France, trials of C. sativa cultivars produced max-
imum yields of up to 2.3 t ha
1
, with sowing in late
March and application of 100 kg N ha
1
(Merrien and
Chatenet 1996). In Poland, average seed yield for winter
crop C. sativa was 1.75 t ha
1
and oil yield averaged 605
kg ha
1
(Musnicki et al. 1997). In Italian trials over 3
yr, 1000-seed weight varied from 1.6 to 2.2 g, and seed
yield per plant from 1.9 to 4.3 g (Angelini et al. 1997). In
Austria, yields of up to 2800 kg ha
1
were found in
some selected cultivars, but large-seeded genotypes with
1000-seed weight of up to 1.81 g were inferior to small-
seeded genotypes in terms of yield performance and oil
content (Vollmann et al. 2007).
(c) Seed Banks, Seed Viability and Seed Germination *
There is little information on seed dormancy of the three
weed species in the literature. There is also little
information available on crop volunteers of C. sativa.
Looman and Best (1987) described the seeds of Came-
lina plants on the prairies as having "a very short
dormant period" In a 3-yr study of C. sativa crops in
Ireland, Crowley (1999), mentions "establishment" i.e.,
germination and emergence occurred within 2 wk
irrespective of planting date. In a study of germination
of various cruciferous species in Great Britain, Ellis
et al. (1989) found that C. sativa germination was
proportional to white light photon dose, was inhibited
by high radiance, which normally inhibits germination,
and was substantially promoted by gibberelic acid
(GA
3
). In Maritime Canada, the germination rate of
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 801
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

C. sativa cultivars was 95% or more, although emer-
gence rates averaged between 38 and 45%, depending
upon environmental conditions (Urbaniak et al. 2008b).
(d) Vegetative Reproduction * The three Camelina
species do not reproduce vegetatively.
9. Hybrids
Camelina sativa and C. alyssum (as C. dentata) are
completely interfertile (Tedin 1922). Al-Shehbaz (1987)
suggested that the three Camelina species are likely able
to intercross and that such hybridization is responsible
for the many intermediate types found in nature and the
blurred species boundaries among the three taxa.
Intergeneric hybrids between C. sativa and Brassica
carinata A. Braun (Narasimhulu et al. 1994), B. oleracea
(Hansen 1998; Sigareva and Earle 1999) and B. napus
(Mu
¨
ller and Sonntag 2000) were obtained by protoplast
fusion. Somatic hybrids of Brassica napusCamelina
sativa showed callus regeneration but no shoot regen-
eration (Mu
¨
ller and Sonntag 2000).
10. Population Dynamics
In a German competition study (Kranz and Jacob 1977)
between Camelina sativa and flax, C. sativa absorbed
more phosphate ions and rubidium than flax when
grown alone, and at an even higher rate in mixed
cultures. Results suggested that reduced yields of flax
grown in association with C. sativa are largely the result
of competition for nutrients. Seedlings from fall or
winter-sown C. sativa compete with many annual weeds
if planted at high densities (Seedtec  Terramax 2008).
Although the small seedlings of C. sativa can be
outcompeted in the early stages of growth, a dense
established crop is highly competitive and will smother
most competing species (Dimmock and Edward-Jones
2006).
11. Response to Herbicides and Other Chemicals
In the United States, use of 2,4-D was recommended for
control of C. microcarpa in small grain crops in early
spring, or in rotation with other crops where other
herbicides could also provide control (Lorenzi and
Jeffery 1987). In non-crop areas such as pastures, fence
rows and waste areas, application of 2,4-D, paraquat,
glyphosate or a soil sterilant were recommended, pre-
ferably prior to seed formation, and in the case of
pastures, followed by periodic mowing close to the
ground (Lorenzi and Jeffery 1987). Sulfonylaminocar-
bonyl triazolinone applied to wheat showed good
control of broadleaf weeds in the mustard family,
including Camelina spp., when applied post-emergence
at the 12 leaf stage at 3045 g a.i. ha
1
(Scoggan et al.
1999).
Camelina microcarpa is the only one of the three
species to have evolved biotypes resistant to acetolactate
synthase (ALS) inhibiting herbicides. Plants were
detected in 1999 in a wheat crop at a site in Oregon
when a mixture of chlorsulfuron plus metsulfuron
applied at up to 56 g a.i. ha
1
did not control C.
microcarpa, whereas a combination of bromoxynil,
MCPA and dicamba provided complete control (Han-
son et al. 2004).
In a German study of the metabolism of
14
C-
glufosinate (GA) (Jansen et al. 2000), excised shoots
and leaves of C. alyssum absorbed a high percentage
(87%) of the radioactivity applied, had one of the lowest
portions of non-extractable residues (0.3%) and a high
rate of recovered radioactivity (ca. 88%). Camelina
alyssum was also among plants with a relatively low
metabolic rate (34.3 ng GA h
1
mg
1
protein). Inter-
mediate symptoms of herbicide damage were detected in
this species, but although there appeared to be little
correlation between either the amount of absorbed
14
C
or the metabolic rate and the degree of damage, a
tendency was observed that plant species with metabolic
rates below 70 ng GA h
1
mg
1
protein were affected
comparatively little by GA. In Azerbaijan, photochemi-
cal activity in the leaves of C. sativa growing as a weed
in wheat fields was completely and irreversibly sup-
pressed following application of bromoxynil (Azizov
et al. 1991).
No herbicides are registered for use in Camelina
oilseed crops in the United States or Canada (Anon-
ymous 2007; Seedtec  Terramax 2008). In Ireland,
where autumn sown Camelina sativa remains at the 4- to
6-leaf stage until spring, and is vulnerable to weed
establishment, the use of trifluralin reduced percent
weed cover at harvest from 29 to 3% on average over a
3-yr period (Crowley 1999).
12. Response to Other Human Manipulation
No information was found on weedy populations of the
three species. In western Canada, Camelina sativa can be
sown conventionally in the spring or surface seeded in
late fall on stubble without seedbed preparation and will
emerge before other annual plants in spring (Seedtec 
Terramax 2008). In oilseed cultivation trials in Maritime
Canada, Urbaniak et al. (2008a), determined that
cultivar selection was the most important determinant
of success at sites in all three provinces; and that plant
height, seed yield, oil content, total plant nitrogen (N)
and seed protein all responded positively to N applica-
tion at rates of 60 kg N ha
1
in Nova Scotia or 80 kg N
ha
1
in Prince Edward Island. With increased N
application rates, oil content of the seeds decreased, as
did contents of oleic and eicosenic acids, while response
of linolenic acid was not consistent at each year-site. In
seeding trials in Nova Scotia and Prince Edward Island
(Urbaniak et al. 2008b), seeding date (two dates from
May 06 to June 02, 2 wk apart at each location) did not
affect emergence, plant height, seed yield, or oil content.
Later-seeded crops had lower stearic acid concentration
in the oil. In the trials, a forage-type seeder pro-
duced more uniform and easily-harvested crops than
seed drills.
802 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

In Montana, early seeding before other spring-planted
crops, early to mid-March, produced maximum yields;
whereas delays after March 20 resulted in yield reduc-
tions averaging 45 kg per week of delay (Anonymous
2007). Weed control in Montana was achieved with
high-density planting, and there was little response to
nitrogen, phosphorus, potassium or sulphur fertilizers.
In Minnesota trials (Putnam et al. 1993), performance of
C. sativa sown on frozen stubble without seedbed
preparation or herbicides was equal or superior to
conventionally sown flax sprayed with herbicides. Sur-
face seeding seemed to work better under no-till condi-
tions; machine planting was no better than broadcast
seeding in spring; and the winter-sown stands emerged
mid-April, before most other spring-sown crops and
before significant weed flushes. However, winter sowing
without the use of herbicides, fungicides or pesticides
does not work well in areas with mild wet winters, as in
Ireland, and can result in high levels of weed competi-
tion, disease infection, and excessive lodging early in the
season, leading to difficult harvesting conditions and
low yields (Crowley 1999; Anonymous 2002).
Among crops investigated in Poland for the effects of
sowing date, precipitation and temperature on yield, C.
sativa showed the highest variability, late sowing
moderately decreasing yields, and drought conditions
markedly reducing yields (Tobola and Musnicki 1999).
Also in Poland, pre-sowing treatments of nitrogen,
phosphorus and potassium (NPK) fertilizer at 125 kg
ha
1
sulphur (S) at 25 kg ha
1
or magnesium (Mg) at
5kgha
1
and a top dressing with N at up to 40 kg ha
1
did not affect the yield of C. sativa, the highest yield of
2.14 t ha
1
being achieved following treatment with
split nitrogen at a rate of 80 kg ha
1
, the method of N
application, in solid urea or in solution, having no
bearing on crop yield (Szczebiot 2002). In Rumania, C.
sativa crop yield was increased by 14% with application
of 40 kg P ha
1
and by 27% with 60 kg P ha
1
;
increases of 37 and 58% followed application of 50 and
100 kg N ha
1
, while P increased oil content from 39.2
to 41.9%, and N decreased oil content from 40.9 to
40.1%, respectively (Bugnarug and Borcean 2000). In
Russia, during the last week in August, optimal densities
of C. sativa allowing rosettes to gain enough biomass to
resist winter frost varied from 6 to 7 million germinating
seeds ha
1
(Semenova and Presnyakova 2007).
13. Response to Herbivory, Disease and Higher
Plant Parasites
Herbivory
(a) Mammals * No information was found on grazing
by domestic or wild animals. Camelina spp. were being
considered for use in fodder mixtures for domestic
animals in Russia (Artemov and Velibekova 2003).
(b) Birds and/or Other Vertebrates * No information
was found on seed predation by birds; but Fogelfors
(1984) reported from Sweden that C. sativa plants were
eaten by geese and other birds.
(c) Insects * Camelina microcarpa, which has been
included in the host range of the swede midge Contarinia
nasturtii Kieffer (Diptera: Cecidomyiidae) in Europe,
showed no evidence of infestation in field experiments in
Ontario (Hallett 2007). Few insects have been reported
as pests of C. sativa in areas where it has been previously
grown. In a Saskatchewan study of the flea beetle
Phyllotreta cruciferae Goeze (Coleoptera: Chrysolmeli-
dae) on the host plant Brassica napus and three non-host
plants, including C. sativa (Henderson et al. 2004), the
latter was found to have the highest resistance, with
010% damage to cotyledons compared with 95100%
on B. napus. It was suggested that resistance in C. sativa
could have resulted from either the presence of repellent
or the absence of stimulatory volatile phytochemicals,
perhaps due to a low level of glucosinolates. In host
plant resistance trials in Lethbridge, AB, C. sativa was
also a poor host for the cabbage seed pod weevil
Ceutorhyncus obstrictus (Marsh.) (Carcamo et al. 2007).
In US studies, flea beetles were sometimes observed
on C. sativa, but in extensive multi-year, small plot
trials, insect damage was not sufficient to warrant
control measures (Robinson 1987). Camelina sativa
was the least acceptable of 10 cruciferous taxa as food
for the flea beetles Phyllotreta nemorum L. and P.
cruciferae in a greenhouse experiment in Switzerland
(Nielsen et al. 2001). Contrary to the reports of C. sativa
resistance to insect predation, Dimmock and Edward-
Jones (2006) reported flea beetle (P. cruciferae) preda-
tion on C. sativa grown in crop trials in Wales, UK, with
leaves of young plants typically damaged by scattered
holes. In Romania, the European tarnished plant bug
Lygus rugulipennis Popp. (Heteroptera: Miridae) was the
characteristic species associated with C. sativa grown as
a prospective oil crop (Palagesiu 2000).
(d) Nematodes and Other Invertebrates * In a Swiss
study of predation in mixed wildflower plantings, which
included Camelina sativa, the slugs Deroceras reticula-
tum Mu
¨
ller and Arion lusitanicus Mabille caused the
heaviest damage to seeds of C. sativa, destroying
4351% of the seeds after 1 wk; but predation of C.
sativa was significantly reduced in harrowed plots
(Kollmann and Bassin 2001).
Disease
(a) Fungi * In Canada, white rust, Albugo cruciferarum
(DC.) Gray, was reported on C. microcarpa in Manitoba;
downy mildew, Peronospora parasitica (Pers. ex Fr.) Fr.,
on both C. microcarpa and C. sativa in Manitoba; and
the fungus-like protist club root, Plasmodiophora brassi-
cae Wor., on C. microcarpa in Prince Edward Island
(Conners 1967). Five fungi were reported as secondary
invaders of stagheads caused by A. cruciferarum on
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 803
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

C. microcarpa in Saskatchewan: black mould, Alternaria
alternate (Fr.:Fr.) Keissler; black leaf spot, A. brassicae
(Berk.) Sacc.; moulds, Cladosporium sp. and Epicoccum
sp.; and, foot-rot fungus, Fusarium roseum (Link) Fr.
(Petrie and Vanterpool 1974; Ginns 1986).
Camelina sativa is very resistant to blackspot, Alter-
naria brassicae (Conn et al. 1988; Sharma et al. 2002).
This resistance was attributed to the production of
phytoalexins, camalexin and methoxycamalexin
(Browne et al. 1991), whose concentrations increased
with increasing A. brassicae inoculum (Jejelowo et al.
1991). The latter are the first reported naturally
occurring antifungal compounds which contain a 2-
substituted thiazone and have antifungal properties
similar to those found in the synthetic fungicide,
thiabendazole, a 4-substituted thiazole. Further studies
determined that a combination of phytoalexin produc-
tion and the destruxin B detoxification pathway provide
blackleg resistance in C. sativa (Pedras et al. 2003). In
Saskatchewan, C. sativa was among cruciferous crop
and wild species evaluated in a growth chamber for
resistance to pre-emergence damping off and post-
emergence seedling root rot caused by Rhizoctonia solani
Ku
¨
hn (Yang and Verma 1992). None of the species
proved to be immune to the pathogen. However, C.
sativa was among a few that had significantly higher
percent emergence than the check Brassica napus and
was among species with a very low percentage of
symptomless days post-emergence.
In the United States, the following fungal pathogens
were reported on C. microcarpa: white rust, Albugo
candida (Pers.: Fr.) Kuntze, from Ohio, Oklahoma,
Pennsylvania, Utah and Virginia; clubroot, Plasmodio-
phora brassicae, from New Jersey; and the rust Puccinia
aristidae Tracy from Colorado and Utah. Pathogens on
C. sativa include: Albugo candida from Montana, Ne-
braska, Ohio, Oklahoma, Oregon, Texas, Utah and
Virgina; and Plasmodiophora brassicae from New Jersey
(Farr et al. 2006). In US studies, C. microcarpa was rated
generally resistant to leaf spot, Alternaria brassicicola
(Schwein.) Wiltshire and black rot Xanthomonas campes-
tris (Pammel) Dowson (only 3% showed susceptibility to
either pathogen), whereas C. sativa was more variable,
with 34% on average showing some degree of suscept-
ibility to leaf spot and 10% to black rot (Westman and
Dickson 1998; Westman et al. 1999). In C. sativa trials in
Montana, almost no disease was noted except in 2006
under high rainfall conditions, when downy mildew,
Peronospora camelinae, was found (Anonymous 2007).
European fungal reports for C. alyssum include
powdery mildew, Erysiphe cruciferarum Opiz ex L.
Junell, from Estonia, Poland, and former USSR.
European and Asian fungal reports for C. microcarpa
include: powdery mildew, E. cichoracearum Jacz., from
Germany, E. cruciferarum from China, France, Poland,
Romania, Switzerland and former USSR, and Peronos-
pora camelinae Ga
¨
um. from Algeria, Central Asia,
former Czechoslovakia, Germany and Poland. Other
reports from Europe and Asia on C. sativa include: E.
cruciferarum from Bulgaria, Finland, France, Germany,
Italy, Switzerland and former USSR; the mildew
Hyaloperonospora camelinae (Ga
¨
um.) Go
¨
ker & al.
from Spain; Peronospora camelinae from Austria, Bul-
garia, Central Asia, Germany and Switzerland; white
leaf spot, Pseudocercosporella capsellae (Ell. & Ev.)
Deighton, from Armenia; and leaf and glume blotch,
Septoria camelinae Lobik, from Russia (Farr et al.
2006). Camelina sativa was found to be the most
resistant to the blackleg fungus, Leptosphaeria maculans
(Desm.) Ces. & de Not., among crucifer species tested in
Poland (Karolewski 1999) and Australia (Salisbury
1987), and no recently tested isolates of this species
were virulent on C. sativa (Li et al. 2005). In Poland,
70% of C. sativa plants were infected with clubroot by
the protist Plasmodiophora brassicae, while low inci-
dence of Peronospora parasitica, Albugo candida and
grey mould, Botrytis cinerea (De Bary) Whetzel, were
reported in a 3-yr study (Nowicki 1975). Significant
variation was found in resistance of C. sativa oilseed
crop genotypes to downy mildew, Peronospora cameli-
nae, in Austria (Vollmann et al. 2001), while Dimmock
and Edward-Jones (2006) reported widespread suscept-
ibility in the United Kingdom. Winter-sown C. sativa
crops in Ireland were highly susceptible to the moulds
Botrytis spp. and Sclerotinia spp., the former causing the
most serious losses in yield (Crowley 1999).
(b) Bacteria * In Germany, C. sativa plants collected
from different localities were found to be infected by
bacterial blight caused by Pseudomonas syringae pv.
camelinae (Mavridis et al. 2002).
(c) Viruses * In North America, C. microcarpa is
reported as a host for turnip mosaic potyvirus
(TuMV), cauliflower mosaic (CaMV), turnip crinkle
(TCV) and turnip rosette (RosV) viruses (Royer and
Dickinson 1999). Camelina sativa has been reported as
susceptible to Erysimum latent tymovirus (ELV) and to
turnip yellow mosaic tymovirus (TYMV) (Brunt et al.
1996 onwards). Transmission of TYMV by C. sativa
seed has been reported (Heim 1984). Camelina sativa
was among a number of plants shown to be hosts of the
beet western yellows virus (BWYV) in Germany (Grai-
chen and Rabenstein 1996). In the Czech Republic, C.
sativa was reported as a host of the radish mosaic virus
(RaMV) (S
ˇ
pak and Kubelkova
´
2000).
(d) Phytoplasmas * Camelina sativa plants in experi-
mental plots in Alberta, Canada, were found to be
infected by the aster yellows phytoplasma disease
(Khadhair et al. 2001).
Higher Plant Parasites * No information was found on
higher plant parasites.
804 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

ACKNOWLEDGEMENTS
We wish to thank Susan Flood, Agriculture and Agri-
Food Canada, for preparation of the figures and to
thank the British Columbia government for permission
to use copyright material for the preparation of Fig. 1.
Abramovic
ˇ
, H. and Abram, V. 2006. Effect of added rosemary
extract on oxidative stability of Camelina sativa oil. Acta
Agric. Slov. 87: 255261.
Abramovic
ˇ
, H., Butinar, B. and Nicolic
ˇ
, V. 2007. Changes
occurring in phenolic content, tocopherol composition and
oxidative stability of Camelina sativa oil during storage. Food
Chem. 104: 903909.
Agegnehu, M. and Honermeier, B. 1997. Effects of seeding rates
and nitrogen fertilization on seed yield, seed quality and yield
components of false flax (Camelina sativa Crtz.). Bodenkultur
48:1521.
Akeroyd, J. R. 1993. Camelina Crantz. Pages 380381 in T. G.
Tutin, N. A. Burges, A. O. Chater, J. R. Edmondson, V. H.
Heywood, D. M. Moore, D. H. Valentine, S. M. Walters, and
D. A. Webb, eds. Flora Europaea. 2nd ed. Vol. I. Psilotaceae
to Platanaceae. Cambridge University Press, Cambridge, UK.
Al-Shehbaz, I. A. 1987. The genera of Alysseae (Cruciferae:
Brassicaceae) in the southeastern United States. J. Arnold
Arbor. 68: 185240.
Alex, J. F. 1992. Ontario weeds. Ontario Ministry of
Agriculture, Food and Rural Affairs. Publ. 505. Toronto,
ON. 304 pp.
Angelini, L. G., Moscheni, E., Colonna, G., Belloni, P. and
Bonari, E. 1997. Variation in agronomic characteristics and
seed oil composition of new oilseed crops in central Italy. Ind.
Crops Prod. 6: 313323.
Anonymous. 2002. Gold of pleasure (false flax). European web
publication by Interactive European Network for Industrial
Crops and their Applications (IENICA). [Online] Available:
http://www.ienica.net/crops/goldofpleasure.pdf [2009 Jan. 07].
Anonymous. 2007. Camelina releases 2007. Montana State
University-Northwestern Agricultural Research Center. [On-
line] Available: http://plantsciences.montana.edu/Foundation-
Seed/2007/Camelina.pdf [2009 Jan. 07].
Artemov, I. V. and Velibekova, E. B. 2003. Intensive produc-
tion technologies, making fodder supplies and use of high
protein content rape fodders in animal husbandry. Kormo-
proizvodstvo 9:1519. [In Russian, English abstract].
Azizov, I. V., Aliev, D. A. and Khalilova, N. M. 1991.
Photochemical activity of chloroplasts of wheat and weeds
under the influence of the herbicide bromoxynil. Sel’sk. Khoz.
Biol. 3: 159164. [In Russian, English summary].
Balschun, H. and Jacob, F. 1975. Studies on competition
between fibre flax and various weeds. Wiss. Beitr. Martin
Luther Univ. Halle Wittenb. 1973/11 (P2): 124129. [In
German, English abstract].
Bernardo, A., Howard-Hildige, R., O’Connell, A., Nichol, R.,
Ryan, J., Rice, B., Roche, E. and Leahy, J. J. 2003. Camelina
oil as a fuel for diesel transport engines. Ind. Crops Prod. 17:
191197.
Blamey, M. and Grey-Wilson, C. 1989. The illustrated flora of
Britain and northern Europe. Hodder & Stoughton, London,
UK. 544 pp.
Bo¨ hme, H., Aulrich, K., Schumann, W. and Fischer, K. 1997.
Studies on the suitability of false flax expeller as feedstuff. I.
Feeding value and incorporation limits for pigs. Fett/Lipid 7:
254259. [In German, English summary].
Boivin, B. 1968. Flora of the Prairie Provinces, Part II.
Phytologia 16: 286287.
Bonjean, A. and Le Goffic, F. 1999. Camelina  Camelina sativa
(L.) Crantz: an opportunity for European agriculture and
industry. Oleag. Corps Gras Lipides 6:2834. [In French,
English summary].
Browne, L. M., Conn, K. L., Ayer, W. A. and Tewari, J. P.
1991. The camalexins: new phytoalexins produced in the leaves
of Camelina sativa (Cruciferae). Tetrahedron 47: 39093914.
Brunt, A. A., Crabtree, K., Dallwitz, M. J., Gibbs, A. J.,
Watson, L. and Zurcher, E. J. (eds.). 1996 onwards. Plant
viruses online: Descriptions and lists from the VIDE database.
Version: 20 August 1996. [Online] Available: http://image.
fs.uidaho.edu/vide/refs.htm [2009 Jan. 07].
Bu
¨
chsenschu
¨
tz-Nothdurft, A., Schuster, A. and Friedt, W. 1998.
Breeding for modified fatty acid composition via experimental
mutagenesis in Camelina sativa (L.) Crtz. Ind. Crops Prod. 7:
291295.
Budin, J. T., Breene, W. M. and Putnam, D. H. 1995. Some
compositional properties of camelina (Camelina sativa (L.)
Crantz) seeds and oils. J. Am. Oil Chem. Soc. 72: 309315.
Bugnarug, C. and Borcean, I. 2000. A study on the effect of
fertilizers on the crop and oil content of Camelina sativa L.
Lucr. Stiint. Agric. Univ. Stiint. Agron. Med. Vet. Banat.
Timisoara 32: 541544.
Callihan, B., Brennan, J., Miller, T., Brown, J. and Moore, M.
2000. Mustards in mustards: guide to identification of canola,
mustard, rapeseed and related weeds. Agricultural Publica-
tions, University of Idaho, Moscow, ID. 26 pp.
Canadian Government. 2005. Seeds Act, Weed Seeds Order,
2005. Canada Gazette Vol. 139, no. 14. 2005 Jul. 13.
Carcamo, H., Olfert, O., Dosdall, L., Herle, C., Beres, B. and
Soroka, J. 2007. Resistance to cabbage seedpod weevil among
selected Brassicaceae germplasm. Can. Entomol. 139: 658669.
Cheo, T.-Y., Lu, L. L., Yang, G. and Al-Shehbaz, I. A. 2001.
Brassicaceae. Pages 1193 in Z. Y. Wu and P. H. Raven, eds.
Flora of China. Vol. 8  Brassicaceae through Saxifragaceae.
Science Press (Beijing) and Missouri Botanical Garden Press,
St. Louis, MO.
Cheo, T.-Y., Lu, L. L., Yang, G. and Al-Shehbaz, I. A. 2003.
Brassicaceae. Pages 1134 in Z. Y. Wu, P. H. Raven, and
D.-Y. Hong, eds. Flora of China illustrations. Vol. 8 
Brassicaceae through Saxifragaceae. Science Press (Beijing)
and Missouri Botanical Garden Press, St. Louis, MO.
Clapham, A. R., Tutin, T. G. and Moore, D. M. 1987. Flora of
the British Isles. ed. 3. Cambridge University Press, Cam-
bridge, UK. 688 pp.
Cody, W. J., Kennedy, C. E., Bennett, B. and Caswell, P. 2004.
New records of vascular plants in the Yukon Territory VI.
Can. Field-Nat. 118: 558578.
Cole, R. A. 1976. Isothiocyanates, nitriles and thiocyanates as
products of autolysis of glucosinolates in Cruciferae. Phyto-
chemistry 15: 759762.
Conn, K. L., Tewari, J. P. and Dahiya, J. S. 1988. Resistance to
Alternaria brassicae and phytoalexin-elicitation in rapeseed
and other crucifers. Plant Sci. (Shannon) 56:2125.
Conners, I. L. 1967. An annotated index of plant diseases in
Canada and fungi recorded on plants in Alaska, Canada and
Greenland. Can. Dep. Agric. Res. Branch, Ottawa, ON. Publ.
1251. 381 pp.
Crowley, J. G. 1999. Evaluation of Camelina sativa as an
alternative oilseed crop. Teagasc, Dublin, Ireland. 9 pp.
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 805
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

Darbyshire, S. J. 2003. Inventory of Canadian agricultural
weeds. Agriculture and Agri-Food Canada, Ottawa, ON.
[Online] Available: http://dsp-psd.pwgsc.gc.ca/Collection/
A42-100-2003E.pdf [2009 Jan. 07].
Darbyshire, S. J., Favreau, M. and Murray, M. 2000. Common
and scientific names of weeds in Canada. Agriculture and
Agri-Food Canada, Ottawa, ON. Publ. 1397/B. 132 pp.
Davis, L. W. 1993. Weed seeds of the Great Plains. University
Press of Kansas, Lawrence, KA. 145 pp.
Daxenbichler, M. E., Spencer, G. F., Carlson, D. G., Rose, G.
R., Brinker, A. M. and Powell, R. G. 1991. Glucosinolate
composition of seeds from 297 species of wild plants.
Phytochemistry 30: 26232638.
Dimmock, J. and Edward-Jones, G. 2006. Crop protection in
alternative crops. Outlooks Pest Manage. 17:2427.
Dorofeev, V. I. 1996. Genus Camelina (Brassicaceae) of the
Caucasian flora. Bot. Zh. (St. Petersbg.) 81:9599. [In
Russian].
Douglas, G. W. and Meidinger, D. 1998. Brassicaceae. Pages
6690 in G. W. Douglas, G. B. Straley, D. Meidinger, and J.
Pojar, eds. Illustrated flora of British Columbia. Vol. 2.
Dicotyledons: Balsamaceae through Cuscutaceae. Govern-
ment of British Columbia, Victoria, BC.
Downey, R. K. 1971. Agricultural and genetic potentials of
cruciferous oilseed crops. J. Am. Oil Chem. Soc. 48: 718722.
Dubois, V., Breton, S., Linder, M., Fanni, J. and Parmentier,
M. 2007. Fatty acid profiles of 80 vegetable oils with regard to
their nutritional potential. Eur. J. Lipid Sci. Technol. 109:
710732.
Easterly, N. W. 1963. Chromosome numbers of some north-
western Ohio Cruciferae. Castanea 28:3942.
El Paso County Conservation District. 2008. New crop
Camelina has biodiesel potential. Colorado online publication.
[Online] Available: http://elpasocd.org/Camelina.htm [2009
Jan. 07].
Elia´ s, P. 2003. Camelina microcarpa L. in Slovakia. Acta
Fytotech. Zootech. 3:5761.
Ellis, R. H., Hong, T. D. and Roberts, E. H. 1989. Quantal
response of seed germination in seven genera of Cruciferae to
white light of varying photon flux density and photoperiod.
Ann. Bot. (Lond.) 63: 145158.
Environment Canada. 2002. Biodiversity portrait of the St.
Lawrence  Distribution species list: Camelina microcarpa,
Camelina sativa. [Online] Available: http://www.qc.ec.gc.ca/
faune/biodiv/en/recherche/especes/recherche_species.html
[2009 Jan. 07].
Facciola, S. 1990. Cornucopia  a source book of edible plants.
Kampong Publications, Vista, CA. 677 pp.
Farr, D. F., Rossman, A. Y., Palm, M. E. and McCray, E. B.
2006. Fungal databases, Systematic Botany and Mycology
Laboratory, ARS, USDA. [Online] Available: http://nt.ars
grin.gov/fungaldatabases/ [2009 Jan. 07].
Flachowsky, G., Schaarmann, G., Jahreis, G., Scho¨ ne, F.,
Richter, G. H., Bo¨ hme, H. and Schneider, A. 1997. Influence
of feeding of oilseeds and byproducts from oilseeds on vitamin
E concentration of animal products. Fett/Lipid 99:5560. [In
German, English summary].
Fogelfors, H. 1984. Useful weeds? Part 5. Lantmannen 105: 28.
[In Swedish, English abstract].
Frame, D. D., Palmer, M. and Peterson, B. 2007. Use of
Camelina sativa in the diets of young turkeys. J. Appl. Poult.
Res. 16: 381386.
Frankton, C. and Mulligan, G. A. 1987. Weeds of Canada.
Revision of 1970 edition. Agriculture Canada, Ottawa, ON.
Publ. 948. 217 pp.
Fro¨ hlich, A. and Rice, B. 2005. Evaluation of Camelina sativa
oil as a feedstock for biodiesel production. Ind. Crops Prod.
21:2531.
Gehringer, A., Friedt, W., Lu
¨
hs, W. and Snowdon, R. J. 2006.
Genetic mapping of agronomic traits in false flax (Camelina
sativa subsp. sativa). Genome 49: 15551563.
Ginns, J. H. 1986. Compendium of plant disease and decay
fungi in Canada 19601980. Agric. Can. Res. Branch Publ.
1813. 416 pp.
Gledhill, D. 1990. The names of plants. Cambridge University
Press, Cambridge, UK. 202 pp.
Goulson, D. 2003. Conserving wild bees for crop pollination.
Food Agric. Environ. 1: 142144.
Graichen, K. and Rabenstein, F. 1996. European isolates of beet
western yellows virus (BWYV) from oilseed rape (Brassica
napus L. ssp. napus) are non-pathogenic on sugar beet (Beta
vulgaris L. var. altissima) but represent isolates of turnip
yellows virus (TuYV). Z. Pflanzenkr. Pflanzschutz 103:
233245.
Grubert, M. 1980. SEM-studies on mucilage producing seeds
and fruits. Plant Syst. Evol. 135: 137149. [In German, English
abstract].
Gugel, R. K. and Falk, K. C. 2006. Agronomic and seed quality
evaluation of Camelina sativa in western Canada. Can. J. Plant
Sci. 86: 10471058.
Gulden, R. H., Warwick, S. I. and Thomas, A. G. 2008. The
Biology of Canadian weeds. 137. Brassica napus L. and B. rapa
L. Can. J. Plant Sci. 88: 951996.
Hallett, R. H. 2007. Host plant susceptibility to the swede
midge (Diptera: Cecidomyiidae). J. Econ. Entomol. 100:
13351343.
Hanelt, P., Hammer, K., Kulpa, W. and Schultze-Motel, J.
1982. Catalog of indigenous taxa of cultivated plants collected
1978 in Poland. Kulturpflanze 30: 215244. [In German,
English abstract].
Hansen, L. N. 1998. Intertribal somatic hybridization between
rapid cycling Brassica oleracea and Camelina sativa (L.)
Crantz. Euphytica 104: 173179.
Hanson, B. D., Park, K. W., Mallory-Smith, C. A. and Thill, D.
C. 2004. Resistance of Camelina microcarpa to acetolactate
synthase inhibiting herbicides. Weed Res. 44: 187194.
Hedrick, U. P. (ed.) 1919. Sturtevant’s notes on edible plants.
J. B. Lyon, Albany, NY. 686 pp.
Heim, A. 1984. Transmission of turnip yellow mosaic virus
through seed of Camelina sativa (gold of pleasure). Z.
Pflanzenkr. Pflanschutz. 91: 549551.
Henderson, A. E., Hallett, R. H. and Soroka, J. J. 2004.
Prefeeding behaviour of the crucifer flea beetle, Phyllotreta
cruciferae, on host and nonhost crucifers. J. Insect Behav. 17:
1739.
Henriksen, P. S. and Robinson, D. 1996. Early Iron Age
agriculture: archaeobotanical evidence from an underground
granary at Overbygard in northern Jutland, Denmark. Vege-
tat. Hist. Archaeobot. 5:111.
Hjelmqvist, H. 1979. Contributions to the knowledge of
prehistoric economic plants of Sweden. Op. Bot. 46:1150.
[In German, English abstract].
Holmgren, P. K., Holmgren, N. H. and Barnett, L. C. (eds.)
1990. Index herbariorum, Part I: The herbaria of the world,
8th ed. New York Botanical Garden. New York, NY. 693 pp.
806 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

Honermeier, B. and Agegnehu, M. 1996. Camelina has a future
as a non-food crop. Neue Landwirtsch. 12:4446. [In German,
English abstract].
Hulte´ n, E. and Fries, M. 1986. Atlas of North European
vascular plants, Part I  III, maps and commentaries. Koeltz
Scientific Books, Ko
¨
nigstein, Germany. 1172 pp.
Hurtaud, C. and Peyraud, J. L. 2007. Effects of feeding
camelina (seeds or meal) on milk fatty acid composition and
butter spreadability. J. Dairy Sci. 90: 51345145.
Jalas, J., Suominen, J. and Lampinen, R. (eds.) 1996. Atlas
Florae Europaeae Distribution of vascular plants in Europe.
Vol. 11 (Cruciferae: Ricotia to Raphanus). Helsinki University
Printing House, Helsinki, Finland. 310 pp.
Jankowski, K. J. and Budzynski, W. S. 2003. Energy potential
of oilseed crops. Electronic J. Polish Agric. Univ., ser.
Agronomy 6:110.
Jansen, C., Schuphan, I. and Schmidt, B. 2000. Glufosinate
metabolism in excised shoots and leaves of twenty plant
species. Weed Sci. 48: 319326.
Jaskiewicz, T. and Sagan, A. 2003. The effect of storage and
antioxidant agent on the stability of lipid fraction in false flax
materials. Rosl. Oleiste 24: 587594. [In Polish, English
summary].
Jaskiewicz, T., Matyka, S. and Sagan, A. 2006. Influence of
extrusion on the nutrient content of false flax materials. Pol. J.
Nat. Sci. 3 (Suppl.): 6772.
Jejelowo, O. A., Conn, K. L. and Tewari, J. P. 1991.
Relationship between conidial concentration, germling
growth, and phytoalexin production by Camelina sativa leaves
inoculated with Alternaria brassicae. Mycol. Res. 95: 928934.
Johnson, D. 2007. Camelina: an emerging crop for bioenergy.
In Vitro Cell Dev. Biol. Animal 43: S1223.
Karolewski, Z. 1999. Characteristics of Leptosphaeria maculans
isolates occurring in Wielkopolska region in 19911996.
Phytopathol. Polon. 18: 103111.
Karvonen, H. M., Aro, A., Tapola, N. S., Salminen, I.,
Uusitupa, M. I. J. and Sarkkinen, E. S. 2002. Effect of a-
linolenic acid-rich Camelina sativa oil on serum fatty acid
composition and serum lipids in hypercholesterolemic subjects.
Metab. Clin. Exp. 51: 12531260.
Khadhair, A.-H., Tewari, J. P., Howard, R. J. and Paul, V. H.
2001. Detection of aster yellows phytoplasma in false flax
based on PCR and RFLP. Microbiol. Res. 156: 179184.
Kno¨ rzer, K.-H. 1978. Entwicklung und ausbreitung des lein-
dotters (Camelina sativa s.l.). Ber. Deutsch. Bot. Ges. 91, S.:
187195. [English abstract].
Kollmann, J. and Bassin, S. 2001. Effects of management on
seed predation in wildflower strips in northern Switzerland.
Agric. Ecosyst. Environ. 83: 285296.
Kranz, E. and Jacob, F. 1977. The competition of Linum
usitatissimum with Camelina sativa for minerals. 2: The uptake
of 32P-phosphate and 86rubidium. Flora (Jena) 166: 505516.
Krist, S., Stuebiger, G., Bail, S. and Unterweger, H. 2006.
Analysis of volatile compounds and triacylglycerol composi-
tion of fatty acid seed oil gained from flax and false flax. Eur.
J. Lip Sci. Technol. 108:4860.
Kumar, P. R. and Tsunoda, S. 1980. Variation in oil content
and fatty acid composition among seeds from the Cruciferae.
Pages 234252 in S. Tsunoda, K. Hinata and C. Go
´
mez-
Campo, eds. Brassica crops and wild allies: biology and
breeding. Japan Scientific Societies Press, Tokyo, Japan.
Leeson, J. Y., Thomas, A. G., Hall, L. M., Brenzil, C. A.,
Andrews, T., Brown, K. R. and Van Acker, R. C. 2005. Prairie
weed surveys of cereal, oilseed and pulse crops from the 1970s
to the 2000s. Weed Survey Series Publ. 05-1, Agriculture and
Agri-Food Canada, Saskatoon Research Centre, Saskatoon,
SK. 395 pp.
Li, H., Barbetti, M. J. and Sivasithamparam, K. 2005. Hazard
from reliance on cruciferous hosts as sources of major gene-
based resistance for managing blackleg (Leptosphaeria macu-
lans) disease. Field Crops Res. 91: 185198.
Li, Z.-X., Guo, X.-S, Chen, Y.-S. and Wu, Z.-H. 2006. Some
newly recorded angiosperm plants from Shaanxi Province.
Xibei Zhiwu-Xuebao 26: 19381939. [In Chinese, English
abstract].
Looman, J. and Best, K. F. 1987. Budd’s flora of the Canadian
prairie provinces. Rev. ed. Agriculture Canada, Research
Branch, Ottawa, ON. Publ. 1662. 863 pp.
Lorenzi, H. J. and Jeffery, L. S. 1987. Weeds of the United
States and their control. Van Nostrand Reinhold Co., New
York, NY. 355 pp.
Lovett, J. V. and Duffield, A. M. 1981. Allelochemicals of
Camelina sativa. J. Appl. Ecol. 18: 283290.
Lovett, J. V. and Jackson, H. F. 1980. Allelopathic activity of
Camelina sativa (L.) Crantz in relation to its phyllosphere
bacteria. New Phytol. 86: 273277.
Lovett, J. V. and Sagar, G. R. 1978. Influence of bacteria in the
phyllosphere of Camelina sativa (L.) Crantz on germination of
Linum usitatissimum L. New Phytol. 81: 617625.
Lu, C. 2008. Camelina sativa: a potential oilseed crop for
biofuels and genetically engineered products. Information
Systems for Biotechnology. Montana State University online
publication. 3 pp. [Online] Available: http://www.isb.vt.edu
[2009 Jan. 07].
Lu, C. and Kang, J. 2008. Generation of transgenic plants of a
potential oilseed crop Camelina sativa by Agrobacterium-
mediated transformation. Plant Cell Rep. 27: 273278.
Marie-Victorin, F. E
´
. C. 1964. Flore laurentienne, ed. 2.
University of Montreal Press, Montreal, QC. 925 pp.
Marquard, R., von and Kuhlmann, H. 1986. Investigations of
productive capacity and seed quality of linseed dodder
(Camelina sativa Crtz.). Fette Seifen Anstrichm. 88: 245249.
[In German, English abstract].
Mattha¨ us, B. 1997. Antinutritive compounds in different
oilseeds. Fett/Lipid 99: 170174.
Mattha¨ us, B. 2002. Antioxidant activity of extracts obtained
from residues of different oilseeds. J. Agric. Food Chem. 50:
34443452.
Mattha¨ us, B. 2004. Camelina sativa  revival of an old
vegetable oil? Ernahr. Umschau 51:1216. [In German,
English summary].
Mattha¨ us, B. and Angelini, L. G. 2005. Anti-nutritive consti-
tuents in oilseed crops from Italy. Ind. Crops Prod. 21:8999.
Mattha¨ us, B. and Zubr, J. 2000. Variability of specific
components in Camelina sativa oilseed cakes. Ind. Crops
Prod. 12:918.
Mavridis, A., Paul, V. and Rudolph, K. 2002. Bacterial blight of
Camelina sativa caused by Pseudomonas syringae pv. cameli-
nae. Beitr. Zuchtungsf. Bundesanst. Zuchtungsf. Kulturpfl. 8:
1012.
McGregor, R. L. 1984. Camelina rumelica, another weedy
mustard established in North America. Phytologia 55:
227228.
Meades, S. J., Hay, S. G. and Brouillet, L. 2000. Anno-
tated checklist of the vascular plants of Newfoundland and
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 807
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

Labrador. [Online] Available: http://www.digitalnaturalhistory.
com/meades.htm [2009 Jan. 07].
Medve, R. J. 1983. The mycorrhizal status of the Cruciferae.
Am. Midl. Nat. 109: 406408.
Merrien, A. and Chatenet, F. 1996. Came
´
line: comment
s’e
´
labore le rendement? Oleoscope 35:2427. [English
abstract].
Mielcarek, A., Zandecka Dziubak, J. and Uczkiewicz, T. 2000.
Influence of abscisic acid (ABA) on plant regeneration
of Camelina sativa L. in in vitro conditions. Rosl. Oleiste 21:
309314. [In Polish, English summary].
Mlakar, S. G., Jakop, M. and Bavec, F. 2003. False flax
(Camelina sativa (L.) Crantz). Sodobno Kmetijstvo 36:2830.
[In Slovenian, English abstract].
Moss, E. H. 1983. Flora of Alberta. ed. 2. J. G. Packer, ed.
University of Toronto Press, Toronto, ON. 687 pp.
Mu
¨
ller, J. and Sonntag, K. 2000. Electrofusion of protoplasts
in Brassicaceae. Eucarpia Cruciferae Newsl. 22:2526.
Mu
¨
ller, M., Ordon, F. and Friedt, W. 1999. Perspectives for
efficient cereal and oilseed production in marginal areas. Z.
Kulturtechnik Landentw. 40: 275281. [In German, English
abstract].
Mulligan, G. A. 1957. Chromosome numbers of Canadian
weeds. I. Can. J. Bot. 35: 779789.
Mulligan, G. A. 1984. Chromosome numbers of some plants
native and naturalized in Canada. Nat. Can. (Que.) 111:
447449.
Mulligan, G. A. 2002a. Chromosome numbers determined
from Canadian and Alaskan material of native and naturalized
mustards, Brassicaceae (Cruciferae). Can. Field-Nat. 116:
611622.
Mulligan, G. A. 2002b. Weedy introduced mustards (Brassica-
ceae) of Canada. Can. Field-Nat. 116: 623631.
Musnicki, C., Toboa, P. and Musnicka, B. 1997. Productivity
of oilseed crops under the conditions of the Wielkopolska
region and the variability of their yields. Rosl. Oleiste 18:
269278. [In Polish, English summary].
Narasimhulu, S. B., Kirti, P. B., Bhatt, S. R., Prakash, S. and
Chopra, V. L. 1994. Intergeneric protoplast fusion between
Brassica carinata and Camelina sativa. Plant Cell Rep. 13:
657660.
Nı
´
Eidhin, D., Burke, J. and O’Beirne, D. 2003a. Oxidative
stability of v3-rich Camelina oil and Camelina oil-based spread
compared with plant and fish oils and sunflower spread. J.
Food Sci. 68: 345353.
Nı
´
Eidhin, D., Burke, J., Lynch, B. and O’Beirne, D. 2003b.
Effects of dietary supplementation with camelina oil on
porcine blood lipids. J. Food Sci. 68: 671679.
Nielsen, J. K., Hansen, M. L., Agerbirk, N., Petersen, B. L. and
Halkier, B. A. 2001. Responses of the flea beetles Phyllotreta
nemorum and P. cruciferae to metabolically engineered Arabi-
dopsis thaliana with an altered glucosinolate profile. Chemo-
ecology 11:7583.
Nowicki, B. 1975. The susceptibility of cultivars of cruciferous
plants grown in Poland to club root, Plasmodiophora brassicae
Wor. Rocz. Nauk Rolniczych 4: 205212.
Nykter, M., Kyma¨ la¨ inen, H.-R., Gates, F. and Sjo¨ berg, A.-M.
2006. Quality characteristics of edible linseed oil. Agric. Food
Sci. 15: 402413.
Onyilagha, J., Bala, A., Hallett, R., Gruber, M., Soroka, J. and
Westcott, N. 2003. Leaf flavonoids of the cruciferous species,
Camelina sativa, Crambe spp., Thlaspi arvense and several
other genera of the family Brassicaceae. Biochem. Syst. Ecol.
31: 13091322.
Palagesiu, I. 2000. Contributions to the knowledge of the
entomofauna of some prospective oil plants in Romania. Lucr.
Stiint. Agric. Univ. Stiinte Agricol. Med. Vet. Banatul.
Timisoara 32: 983992.
Pedras, M. S. C., Montaut, S., Zaharia, I. L., Gai, Y. and
Ward, D. E. 2003. Transformation of the host-selective toxin
destruxin B by wild crucifers: probing a detoxification path-
way. Phytochemistry 64: 957963.
Peiretti, P. G. and Meineri, G. 2007. Fatty acids, chemical
composition and organic matter digestibility of seeds and
vegetative parts of false flax (Camelina sativa L.) after different
lengths of growth. Anim. Feed Sci. Technol. 133: 341350.
Peiretti, P. G., Mussa, P. P., Prola, L. and Meineri, G. 2007.
Use of different levels of false flax (Camelina sativa L.) seed in
diets for fattening rabbits. Livestock Sci. 107: 192198.
Petrie, G. A. and Vanterpool, T. C. 1974. Fungi associated with
hypertrophies caused by infection of cruciferae by Albugo
cruciferarum. Can. Plant Dis. Surv. 54:3741.
Petrikova, V., Vana, J. and Ustjak, S. 1996. Production and
utilization of industrial and energy crops on reclaimed soils.
Metod. Zemedelsk. Praxi 17:124. [In Czech, English sum-
mary].
Pilgeram, A. L., Sands, D. C., Boss, D., Dale, N., Wichman, D.,
Lamb, P., Lu, C., Barrows, R., Kirkpatrick, M., Thompson, B.
and Johnson, D. L. 2007. Camelina sativa, a Montana omega-3
and fuel crop. Pages 129131 in J. Janick and A. Whipkey, eds.
Issues in new crops and new uses. ASHS Press, Alexandria,
VA.
Plants For A Future. 2004. Camelina sativa- (L.) Crantz: Gold
of Pleasure. [Online] Available: http://www.pfaf.org/database/
plants.php?Camelinasativa [2009 Jan. 07].
Plessers, A. G., McGregor, W. G., Carson, R. B. and
Nakoneshny, W. 1962. Species trials with oilseed plants. II.
Camelina. Can. J. Plant Sci. 42: 452459.
Polunin, O. 1969. Flowers of Europe  A field guide. Oxford
University Press, London, UK. 662 pp.
Putnam, D. H., Budin, J. T., Field, L. A. and Breene, W. M.
1993. Camelina: a promising low-input oilseed. Pages 314322
in J. Janick and J. E. Simon, eds. New crops. Wiley, New
York, NY.
Reaney, M. J. T., Furtan, W. H. and Loutas, P. 2006. A critical
cost benefit analysis of oilseed biodiesel in Canada. A BIOCAP
Research Integration Program Synthesis Paper. [Online]
Available: http://www.biocap.ca [2009 Jan. 07].
Rich, T. C. G. 1991. Crucifers of Great Britain and Ireland.
Botanical Society of the British Isles, London, UK. 329 pp.
Robinson, R. 1987. Camelina: a useful research crop and a
potential oilseed crop. Minn. Agric. Exp. Stn. Bull. 579. 12 pp.
Robinson, R. G. and Nelson, W. W. 1975. Vegetable oil
replacements for petroleum oil adjuvants in herbicide sprays.
Econ. Bot. 29: 146151.
Rode, J. 2002. Study of autochthon Camelina sativa (L.)
Crantz in Slovenia. J. Herbs Spices Med. Plants 9: 313318.
Rokka, T., Ale´ n, K., Valaja, J. and Ryha¨ nen, E.-L. 2002. The
effect of a Camelina sativa enriched diet on the composition
and sensory quality of hen eggs. Food Res. Int. 35: 253256.
Rollins, R. C. 1993. The Cruciferae of continental North
America. Stanford University Press, Stanford, CA. 976 pp.
Rostovtseva, Z. P. 1975. Primary processes of organogenesis
of annual crucifers in connection with formation of the
808 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

morphogenetic type of metamere. Biol. Nauki (Mosc.) 18:60
65. [In Russian, English abstract].
Rousseau, C. 1968. Histoire, habitat et distribution de 220
plantes introduites au Que
´
bec. Nat. Can. (Que.) 95:49169.
Royer, F. and Dickinson, R. 1999. Weeds of Canada and the
northern United States. Lone Pine Publishing, University of
Alberta Press, Edmonton, AB. 434 pp.
Ryha¨ nen, E.-L., Perttila¨ , S., Tupasela, T., Valaja, J., Eriksson,
C. and Larkka, K. 2007. Effect of Camelina sativa expeller cake
on performance and meat quality of broilers. J. Sci. Food
Agric. 87: 14891494.
Sabourin, A. 1992. Guide des crucife
`
res sauvages de l’est du
Canada (Que
´
bec, Ontario et Maritimes), nouvelle e
´
dition. Les
Amis du Jardin Botanique, Montre
´
al, QC. 249 pp.
Salisbury, P. A. 1987. Blackleg resistance in weedy crucifers.
Eucarpia Cruciferae Newsl. 12: 90.
Salminen, H., Este´ vez, M., Kivikari, R. and Heinonen, M. 2006.
Inhibition of protein and lipid oxidation by rapeseed, camelina
and soy meal in cooked pork meat patties. Eur. Food Res.
Technol. 223: 461468.
Sang, J. P. and Salisbury, P. A. 1987. Wild crucifer species and
4-hydroxyclucobrassicin. Eucarpia Cruciferae Newsl. 12: 113.
Saskatchewan Government 1999. The noxious weeds designa-
tion regulations. Chapter N-9.1 as amended by Saskatchewan
Regulations 14/1999. The Queen’s Printer, Regina, SK. [On-
line] Available: http://www.qp.gov.sk.ca/ [2009 Jan. 07].
Saucke, H. and Ackermann, K. 2006. Weed suppression in
mixed cropped grain peas and false flax (Camelina sativa).
Weed Res. 46: 453461.
Schultze-Motel, W. 1986. Camelina. Pages 340345 in G. Hegi,
ed. Illustrierte flora von Mittel-europa, 3rd ed. Vol. 4. Verlag
Paul Parey, Berlin, Hamburg, Germany.
Schuster, A. and Friedt, W. 1995. Camelina sativa: old face 
new prospects. Eucarpia Cruciferae Newsl. 17:67.
Schuster, A. and Friedt, W. 1998. Glucosinolate content and
composition as parameters of quality of Camelina seed. Ind.
Crops Prod. 7: 297302.
Scoggan, A. C., Santel, H. J., Wollam, J. W. and Rudolph, R.
D. 1999. BAY MKH 6561: A new herbicide for grass and
broadleaf control in cereals. Proc. Brighton Crop Prot. Conf.
1:9398.
Scoggan, H. J. 1957. Flora of Manitoba. Natl. Mus. Can. Bull.
140, Biol. Ser. No. 47. 619 pp.
Seedtec

Terramax 2008. Products: Camelina. Qu’Appelle,
SK. [Online] Available: http://www.terramax.sk.ca/specialty/
camelina.html [2009 Jan. 07].
Seehuber, R. 1984. Genotypic variation for yield and quality
traits in poppy and false flax. Fette Seifen Anstrichm. 5:
177180. [In German, English abstract].
Semenova, E. F. and Presnyakova, E.V. 2007. Biochemical
monitoring of the frost-resistance in winter plants of Camelina
sativa. S-Kh. Biol. 3: 106109. [In Russian, English abstract].
Sharma, G., Dinesh Kumar, V., Haque, A., Bhat, S. R.,
Prakash, S. and Chopra, V. L. 2002. Brassica coenospecies: a
rich reservoir for genetic resistance to leaf spot caused by
Alternaria brassicae. Euphytica 125: 411417.
Shukla, V. K. S., Dutta, P. C. and Artz, W. E. 2002. Camelina
oil and its unusual cholesterol content. J. Am. Oil Chem. Soc.
79: 965969.
Sigareva, M. A. and Earle, E. D. 1999. Camalexin induction in
intertribal somatic hybrids between Camelina sativa and rapid-
cycling Brassica oleracea. Theor. Appl. Genet. 98: 164170.
S
ˇ
pak, J. and Kubelkova´ , D. 2000. Serological variability among
European isolates of radish mosaic virus. Plant Pathol. 49:
295301.
Stace, C. 1997. New flora of the British Isles. ed. 2. Cambridge
University Press, Cambridge, UK. 1130 pp.
Steinke, G., Kirchhoff, R. and Mukherjee, K. D. 2000. Lipase-
catalyzed alcoholysis of Crambe oil and Camelina oil for the
preparation of long-chain esters. J. Am. Oil Chem. Soc. 77:
361366.
Steinke, G., Weitkamp, P., Klein, E. and Mukherjee, K. D.
2001. High-yield preparation of wax esters via lipase-catalyzed
esterification using fatty acids and alcohols from Crambe and
Camelina oils. J. Agric. Food Chem. 49: 647651.
Szczebiot, M. 2002. Effect of mineral fertilization on yielding
of spring false flax and crambe. Rosl. Oleiste 23: 141150. [In
Polish, English abstract].
Tattersall, A. and Millam, S. 1999. Establishment and in vitro
regeneration studies of the potential oil crop species Camelina
sativa. Plant Cell Tissue Organ Cult. 55: 147149.
Tedin, O. 1922. Zur Blu
¨
ten- und Befruchtungsbiologie der
Leindotter (Camelina sativa). Bot. Not. 1922: 177189.
Thellung, A. 1919. Camelina. Pages 368371 in G. Hegi,
Illustrierte Flora von Mittel-Europa, Vol. 1, Part 4. J. F.
Lehmanns Verlag, Munich, Germany. [In German].
Tobola, P. and Musnicki, C. 1999. Yield variability of spring-
sown cruciferous oilseed crops. Rosl. Oleiste 20:93100. [In
Polish, English abstract].
Toncea, I., Hebean, V., Ionita, G. N. and Codreanu, M. 2006.
Experimental results regarding the importance and biological
and technological peculiarities of Camelina (false flax). Probl.
Agrofit. Teoret. Aplic. 28:3747. [In Romanian, English
summary].
Torrey, J. and Gray, A. 1838

1840. A flora of North America.
Vol. 1. Wiley & Putnam, New York, NY. 711 pp.
Urbaniak, S. D., Caldwell, C. D., Zheljazkov, V. D., Lada, R.
and Luan, L. 2008a. The effect of cultivar and applied nitrogen
on the performance of Camelina sativa L. in the Maritime
Provinces of Canada. Can. J. Plant Sci. 88: 111119.
Urbaniak S. D., Caldwell, C. D., Zheljazkov, V. D., Lada, R.
and Luan, L. 2008b. The effect of seeding rate, seeding date
and seeder type on the performance of Camelina sativa L. in
the Maritime Provinces of Canada. Can. J. Plant Sci. 88:
501508.
USDA, NRCS. 2008. Plants database. Natural Resources
Conservation Service, U.S. Dep. Agric. [Online] Available:
http://plants.usda.gov [2009 Jan. 07].
Vollmann, J., Damboeck, A., Baumgartner, S. and Rucken-
bauer, P. 1997. Selection of induced mutants with improved
linolenic acid content in camelina. Fett/Lipid 99: 357361.
Vollmann, J., Damboeck, A., Eckl, A., Schrems, H. and
Ruckenbauer, P. 1996. Improvement of Camelina sativa,an
underexploited oilseed. Pages 357362 in J. Janick, ed. Pro-
gress in new crops. American Society of Horticultural Science
Press, Alexandria, VA.
Vollmann, J., Grausgruber, H., Stift, G., Dryzhyruk, V. and
Lelley, T. 2005. Genetic diversity in camelina germplasm as
revealed by seed quality characteristics and RAPD poly-
morphism. Plant Breed. 124: 446453.
Vollmann, J., Moritz, T., Kargl, C., Baumgartner, S. and
Wagentristl, H. 2007. Agronomic evaluation of camelina
genotypes selected for seed quality characteristics. Ind. Crops
Prod. 26: 270277.
FRANCIS AND WARWICK  CAMELINA ALYSSUM, C. MICROCARPA, C. SATIVA 809
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.

Vollmann, J., Steinkellner, S. and Glauninger, J. 2001. Varia-
tion in resistance of Camelina (Camelina sativa [L.] Crtz.) to
downy mildew (Peronospora camelinae Ga
¨
um). J. Phytopathol.
(Berl.) 149: 129133.
Warwick, S. I. and Al-Shehbaz, I. A. 2006. Brassicaceae:
chromosome number index and database on CD-Rom. Plant
Syst. Evol. 259: 237248.
Warwick, S. I. and Francis, A. 2006. The biology of invasive
alien plants in Canada. 6. Berteroa incana (L.) DC. Can.
J. Plant Sci. 86: 12971309.
Warwick, S. I., Francis, A. and Al-Shehbaz, I. A. 2006.
Brassicaceae: Species checklist and database on CD-Rom.
Plant Syst. Evol. 259: 249258.
Warwick, S. I., Francis, A. and Mulligan, G. A. 1999.
Brassicaceae of Canada. Agriculture and Agri-Food Canada.
1999 Updated version. [Online] Available: http://www.scib.gc.
ca/spp_pages/brass/index_e.php [2009 Jan. 07].
Westman, A. L. and Dickson, M. H. 1998. Disease reaction to
Alternaria brassicicola and Xanthomonas campestris pv. cam-
pestris in Brassica nigra and other weedy crucifers. Eucarpia
Cruciferae Newsl. 20:8788.
Westman, A. L., Kresovich, S. and Dickson, M. H. 1999.
Regional variation in Brassica nigra and other weedy crucifers
for disease reaction to Alternaria brassicicola and Xanthomo-
nas campestris pv. campestris. Euphytica 106: 253259.
Willcox, G. H. 1977. Exotic plants from Roman waterlogged
sites in London. J. Archaeol. Res. 4: 269282.
Yang, J. and Verma, P. R. 1992. Screening genotypes for
resistance to pre-emergence damping-off and postemergence
seedling root rot of oilseed rape and canola caused by
Rhizoctonia solani AG 2-1. Crop Prot. 11: 443448.
Zandecka-Dziubak, J. and Uczkiewicz, T. 1999. Shoot regen-
eration from hypocotyls explants of Camelina sativa L. in in
vitro culture. Rosl. Oleiste 20: 631635. [In Polish, English
summary].
Zubr, J. 1997. Oil-seed crop: Camelina sativa. Ind. Crops Prod.
6: 113119.
Zubr, J. 2003a. Dietary fatty acids and amino acids of
Camelina sativa seed. J. Food Qual. 26: 451462.
Zubr, J. 2003b. Qualitative variation of Camelina sativa seed
from different locations. Ind. Crops Prod. 17: 161169.
Zubr, J. and Mattha¨ us, B. 2002. Effects of growth conditions
on fatty acids and tocopherols in Camelina sativa oil. Ind.
Crops Prod. 15: 155162.
810 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 201.20.182.28 on 10/19/15
For personal use only.