The Biology of Canadian Weeds. 137.

Brassica napus L. and B. rapa L.

Robert H. Gulden1, Suzanne I. Warwick2, and A. Gordon Thomas3
1
Department of Plant Science, 222 Agriculture Building, 66 Dafoe Road, University of Manitoba, Winnipeg,
Manitoba, Canada R3T 2N2 (e-mail: gulden@cc.umanitoba.ca); 2Eastern Cereal and Oilseed Research Centre,
Agriculture and Agri-Food Canada, K.W. Neatby Bldg., Central Experimental Farm, Ottawa, Ontario, Canada K1A
0C6; and 3Saskatoon Research Centre, Agriculture and Agri-Food Canada, 107 Science Place, Saskatoon,
Saskatchewan, Canada S7N 0X2. Received 10 December 2007, accepted 21 April 2008.
Gulden, R. H., Warwick, S. I. and Thomas, A. G. 2008. The Biology of Canadian Weeds. 137. Brassica napus L. and
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22

B. rapa. Can. J. Plan Sci. 88: 951
996. Brassica napus and B. rapa are native to Eurasia. In Canada, these species are
commonly referred to as volunteer canola, while feral populations of B. rapa are referred to as birdrape. Brassica napus and
B. rapa have been grown commercially for their seed oil content in western Canada since the middle of the last century and
volunteer populations are common in fields. Escaped populations of both species are also found along roadways, railways
and in waste areas; however, only B. rapa is known to have naturalized, self-sustaining feral populations in these habitats
in eastern Canada. Despite these escaped and feral populations, B. napus and B. rapa are mainly a concern in agricultural
fields where their combined relative abundance has increased over the past few decades. In the mid 1990s, herbicide-
resistant genotypes of B. napus were released for commercial production. Herbicide-resistance and the stacking of genes in
volunteer populations conferring resistance to multiple herbicides have contributed to increased difficulties in controlling
volunteer B. napus in some crops. However, yield loss resulting from volunteer populations is not well documented in
Canada.

Key words: Brassica napus, Brassica rapa, herbicide resistance, transgene escape, volunteer canola, weed biology
For personal use only.

Gulden, R. H., Warwick, S. I. et Thomas, A. G. 2008. La biologie des mauvaises herbes au Canada. 137. Brassica napus L. et
B. rapa L. Can. J. Plant Sci. 88: 951
996. Brassica napus et B. rapa sont des plantes originaires d’Eurasie. Au Canada, on
les désigne souvent sous l’expression « canola spontané », tandis que les peuplements sauvages de B. rapa sont appelés
« navette ». Dans l’Ouest canadien, Brassica napus et B. rapa sont cultivés commercialement pour l’huile que renferment
leurs graines depuis le milieu du siècle dernier et les repousses spontanées sont devenues monnaie courante dans les
champs. On observe des peuplements des deux espèces le long des routes et des voies ferrées ainsi que dans les terrains
vagues, cependant on sait que seule B. rapa s’est acclimatée pour donner des populations sauvages et stables dans les
mêmes habitats, dans l’est du pays. Malgré cela, B. napus et B. rapa demeurent surtout préoccupants dans les cultures, où
leur abondance relative, une fois combinés, s’est accrue au cours des dernières décennies. Au milieu des année 1990, des
génotypes de B. napus résistants aux herbicides ont été homologués pour la culture commerciale. Cette résistance
et l’accumulation des gènes qui la codent dans les peuplements spontanés ont concouru à compliquer la lutte contre les
repousses de B. napus dans certaines cultures. Quoi qu’il en soit, les pertes de rendement qui en résultent sont mal étayées
au Canada.

Mots clés: Brassica napus, Brassica rapa, résistance aux herbicides, fuite transgénique, canola spontané,
biologie des mauvaises herbes

1. Name
Brassica napus L.
Argentine canola, canola, rapeseed,
oilseed rape, canola colza [not listed as a weed in
Crompton et al. (1988) and Darbyshire et al. (2000);
volunteer listed as a weed in Darbyshire 2003]. BAYER
code for B. napus BRSNN.
Brassica rapa L. (syn. B. campestris L.), bird rape
(Darbyshire et al. 2000), bird’s rape, bird-rape, field
mustard, field-mustard, wild turnip, rutabaga (Cromp-
ton et al. 1988; Darbyshire 2003), common turnip,
birdsrape mustard (US) (Darbyshire 2003); moutarde
des oiseaux (Darbyshire et al. 2000), chou champêtre,
chourave, chou-rave, colza, colza-navette, moutarde,
moutarde champêtre, moutarde d’Allemagne, moutarde
des champs, moutarde sauvage, navet sauvage, navette,
navette d’Allemagne, navette des oiseaux, navette d’été,
navette fourragère, navette oléagineuse, navette sau-
vage, rave, sénevé à feuilles lisses (Crompton et al. 1988;
Darbyshire 2003), navet blanc (Darbyshire 2003).
BAYER code for B. rapa BRSRA. Brassicaceae or
Cruciferae, mustard family, Brassicacées or Crucifères.
Brassica rapa (AA genome, 2n 20 chromosomes)
and B. oleracea L. (CC genome, 2n 18 chromosomes)

951
 952 CANADIAN JOURNAL OF PLANT SCIENCE
are the progenitor species of B. napus (AACC genome,
2n 38 chromosomes) (U 1935; Palmer et al. 1983;
Parkin et al. 1995). Both Brassica rapa and B. napus
have been developed as oilseed crop species. Brassica
rapa was first grown in Canada as an industrial oilseed
crop and developed in the late 1970s as the first canola-
quality (low glucosinolate and low erucic acid) crop
(Polish canola, canola navette) cultivated in Canada. It
has also been cultivated as a European and Asian
vegetable (turnip, Chinese cabbage etc.) and Indian
oilseed (reviewed in Specht and Diederichsen 2001) and
grows as a recent crop escape (volunteer) or naturalized
weed (self-sustaining populations) in many parts of the
world. Canola-quality B. napus (Argentine canola,
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22
canola colza) was developed in Canada in the late 1970s.
2. Description and Account of Variation
(a) Species Description * Annual, winter annual or
biennial; taproot slender, although cultivated forms of
B. rapa (turnip) and B. napus (rutabaga) have swollen
roots. Stems erect, 0.5
1.3 m high, simple to freely
branched above, glabrous (B. napus) or with coarse
trichomes (B. rapa, rarely in B. napus). Basal rosette
leaves are not persisting, short-petiolate, 5
20 cm long,
ovate to elongated, lobed to deeply divided; glaucous
(blue-green), hairless (B. napus) or having numerous
For personal use only.
distinct hairs with a slightly bulbous base, and giving the
appearance of dots on the leaves when viewed from
above (B. rapa); upper stem or cauline leaves are
reduced, often entire (B. napus) or simply toothed (B.
rapa) with enlarged bases that clasp the stem, completely
in B. rapa and often partially clasping in B. napus.
Flowers arranged in terminal many-flowered branched
raceme, clustering at the end of the stem and branches,
and conspicuously elongating in fruit. Open flowers of
the raceme overtopping the buds (B. rapa), below the
buds (B. napus), or at same level as buds (both species).
Fruiting pedicels ascending to spreading, 1
2.5 cm long.
Flowers with 4 sepals, spreading; 4 petals arranged in a
cross, yellow (bright yellow to yellow in B. rapa and pale
yellow in B. napus); obovate to broadly obovate;
6
11 (B. rapa) and 10
16 (B. napus) mm across; stamens
6 (4 long and 2 short, tetradynamous arrangement);
ovules 1
12, style narrow and persistent. Fruit is a linear
nearly cylindrical silique, with distinct valve (lower) and
beak (upper) segments; valves dehiscent, many-seeded,
4.5
7.5 (10) cm long, 2
5 mm thick (narrower 2
4 mm in
B. rapa and 3.5
5 mm in B. napus), smooth or slightly
torulose (i.e., cylindrical with slight constrictions at9
regular intervals), with one central vein; beak portion
usually seedless, ca. 1=4
1u3 the length of the fruit or
9
16 mm (B. napus) and 1/3
1/2 fruit length or 8
22 mm
in B. rapa). Seeds are arranged in one row in the fruit,
nearly spherical to round, reddish brown, brown, or
black, 1.1
2 mm (B. rapa) and 1.8
2.7 mm (B. napus),
surface appearing finely net-veined (i.e. reticulate alveo-
late) at high magnification, not mucilagenous when wet.
Cotyledons conduplicate (folded longitudinally around
the radicle in the seed). Cotyledons in the seedlings are
broadly kidney-shaped, 6
12 mm wide, with a charac-
teristic deep, wide rounded notch at the end. (Frankton
and Mulligan 1987; Alex 1992; Sabourin 1992; Rollins
1993; S. I. Warwick and I. Al-Shehbaz unpublished
data).
The chromosome number for Brassica rapa is n10,
2n 20 and for B. napus is n 19, 2n 38 (reviewed in
Warwick et al. 2000; Warwick and Al-Shehbaz 2006).
There are no counts reported for weedy B. napus in
Canada (reviewed in Mulligan 2002a). Canadian counts
for B. rapa include: n 10 from British Columbia
(Taylor and Mulligan 1968), 2n 20 from New Bruns-
wick (Mulligan 1959) and Quebec (Gervais 1979).
(b) Distinguishing Features * Brassica rapa and B.
napus can be easily recognized by a combination of
characters: broad kidney-shaped cotyledons, having a
deep, wide, rounded notch at the end; lower leaves
petiolate and toothed, entire to lobed, but upper ones
sessile, with enlarged bases clasping the stem; yellow
four-petalled flowers, ca. 1.2 cm across, with erect-
spreading sepals; fruit pods are usually hairless, stalked,
valves are strongly one-nerved and split lengthwise at
maturity, pod is terminated by a seedless beak, and
seeds are brown-black. Brassica rapa and B. napus may
be confused with a number of other annual yellow-
flowered crucifers: Brassica nigra (L.) Koch (black
mustard), Brassica juncea (L.) Czern. (Indian mustard),
Sinapis alba L. (white mustard), Sinapis arvensis L. (wild
mustard), and Raphanus raphanistrum L. (wild radish).
Wild mustard differs from all of these mustards by
having a seed in the beak of the fruit in addition to those
within the valves. Both wild mustard and white mustard
have stiff downward directed hairs on the stem, but
distinctly different fruit pods (see key below). Bird rape
and canola are distinguished by the absence of down-
ward-directed hairs on the stems (B. rapa), or the
absence of hairs and a bluish-green or glaucous tinge
(B. napus), clasping upper leaves, smooth pods with a
single vein and a long thin seedless beak. Wild radish is
similar to bird rape in general habit, but differs in
having more deeply divided lower leaves, flower petals
with conspicuous dark veins that are usually yellow, but
may be white or purple; and distinctive fruit pods as
indicated in the key below. Indian mustard is glabrous,
with little or no hair, has paler yellow flowers, upper
leaves are smooth and narrow, fruit pods have a slender
long stalk (1.2 cm), and a slender seedless beak, and
brownish-red seeds. Black mustard resembles wild
mustard when young, but grows much taller (up to
2.5 m); the lower leaves have coarser lobed divisions,
upper leaves are narrow without teeth; flowers are
smaller and the inflorescence is widely branched; seed
pods are shorter (1
2 cm), erect and pressed to the stem,
and the seedless beak is needle-like. Mature plants with
fruit can be identified with the following key in Warwick
et al. (2000) adapted from Mulligan and Bailey (1975),
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
A. Fruit pods with numerous ribs and strongly con-
stricted, breaking off at the constrictions with one
seed in each pod segment ........................Raphanus
raphanistrum.
A. Fruit pods, either 1- or 3
5 veined, not strongly
constricted, valves splitting lengthwise.
B. Fruit pods each with 3 to 5 conspicuous
parallel veins, beak one-seeded or strongly
flattened; stem with short stiff downward-
directed hairs.
C. Fruit pods and at least base of beak with
white bristly hairs; beak flattened, curved,
equalling to or exceeding length of valves,
beak seedless.............................. Sinapis alba.
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C. Fruit pods glabrous or rarely with bristly
hairs; beak glabrous, long conical, straight,
slightly winged, usually one-seeded, beak
considerably shorter than length of the val-
ves........................................ Sinapis arvensis.
B. Fruit pods each 1-veined, i.e. with a conspicuous
midrib and much weaker lateral veins; beak neither
one-seeded nor strongly flattened; stem lacking
short stiff downward-directed hairs.
D. Upper stem leaves stalked or narrowed into a
stalk-like base, not clasping; beak less than
6.0 mm, not gradually tapering.
For personal use only.
E. Leaves green, pods appressed to
stem, 1.0
2.0 cm, with mature pods
more or less four-sided, beak 1.5
3.0
mm, needle-like, fruiting pedicels
short (1.5
3.0 mm), appressed to
stem ..........................Brassica nigra.
E. Leaves glaucous, pods ascending,
3.0
5.0 cm, with mature pods
slightly flattened, perpendicular to
partition, beak 5.0
10.0 mm long,
fruiting pedicels 8.0
14.0 mm, sprea-
ding......................... Brassica juncea.
D. Upper stem leaves half to completely clasping stem,
broadened at base; beak longer than 6.0 mm,
gradually tapering.
F. Open flowers overtopping or equal to the
buds; petals bright yellow, 6.0
11 mm; beak
or terminal fruit segment 8
22 mm, 1/3 to ½
length of the valves .................Brassica rapa.
F. Flower buds overtopping or rarely at the
same level of open flowers; petals pale yellow,
10
16 mm; beak or terminal fruit segment
usually 9
16 mm, 1/4 to 1/3 length of the
valves ..................................... Brassica napus.
Seeds of B. napus and B. rapa may be difficult to
distinguish from those of other Brassicaceae crops
(mustard species) and weeds (S. arvensis). However,
under magnification, all Brassica crop species, B. rapa,
B. oleracea, B. nigra, B. napus, B. juncea, and S. alba and
S. arvensis can be easily distinguished on the basis of
seed coat surfaces (Mulligan and Bailey 1976). The seed
953
coat pattern of B. napus is intermediate between that of
B. oleracea and B. rapa. Andrew et al. (1987) suggested
that B. napus and S. arvensis could be distinguished
based on the primary seed surface wax, with B. napus
featuring nonacosane (C29H60) and S. arvensis featuring
hentriacontane (C31H64).
(c) Intraspecific Variation * Numerous intraspecific
taxa (subspecifc and varietal taxonomic rank) have been
described for both B. rapa and B. napus, corresponding
to various crop types (reviewed in Specht and Dieder-
ichsen 2001). The wild or weedy type of B. rapa has been
referred to as subsp. sylvestris, although Specht and
Diederichsen treat this as a synonym of subsp. oleifera.
Molecular-based phylogenetic studies (Song et al. 1988a;
Warwick and Black 1991) have indicated that B. rapa
and B. oleracea, genomes A and C, respectively con-
stitute a separate lineage from that containing B. nigra
(genome B) and S. arvensis. Genetic maps developed for
both the A and C genomes have indicated a high percent
of similarity (Parkin et al. 1995).
A number of genetic or phylogenetic relationships
have been proposed for taxa within B. rapa, based on
morphology, geographical distribution, isozymes and
nuclear RFLP data (reviewed in Song et al. 1988b;
Specht and Diederichsen 2001). Various data have
indicated a division of B. rapa into two main groups, a
Western and an Eastern group, perhaps corresponding
to two independent centres of origin (Denford and
Vaughan 1977; Prakash and Hinata 1980; Song et al.
1988b; McGrath and Quiros 1992). The western or
European group includes turnip and turnip rape from
which India-Asian oilseed types are putatively derived.
The eastern or Asian group contains the various Asian
vegetables indicated above.
Molecular markers have been extensively utilized as
tools for assessing genetic diversity and genetic relation-
ships in Brassica crop species, restriction fragment
length polymorphisms (RFLPs) (Song et al. 1988b),
random amplified polymorphic DNA (RAPD) markers
were used to discriminate among cultivars within B.
rapa (Ren et al. 1995; Zhang et al. 2000; He et al. 2002,
2003; Wang et al. 2002) and several studies have used
amplified fragment length polymorphisms (AFLP) mar-
kers to assess genetic variation in Brassica crop species,
including B. rapa (Das et al. 1999; Guo et al. 2002; Zhao
et al. 2005) and B. napus (e.g., Xu et al. 2001; Liu et al.
2005).
(d) Illustrations * Brassica napus and B. rapa are
illustrated in Fig. 1. Several web-based illustrations of
seedlings, mature plants, inflorescences, fruit and seed
are also available, e.g., http://plants.usda.gov/ or http://
www.weedscience.com. Photographs of seedlings and
mature plants of B. rapa are available in Bouchard and
Néron (1999).
 954 CANADIAN JOURNAL OF PLANT SCIENCE
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For personal use only.
Fig. 1. Brassica rapa a: habit 1/8, c. flower 3, d. fruit1, and e. seed 10; b.upper part of plant of B. napus1/6 (adapted from
Mohlenbrock 1980). Brassica napus lower stem, root, flower, fruit, and seed morphology similar to B. rapa except for increased size,
see description in Section 2a.
3. Economic Importance
(a) Detrimental * Brassica napus and B. rapa are
grown as oilseed crops in Canada. Despite the ubiqui-
tous presence of volunteer B. napus and B. rapa in
areas where they are grown as a crop, reports of their
ability to interfere with crops are limited in Canada.
Volunteer canola (B. napus and B. rapa) has increased
in abundance during the past three decades from 31st
and 34th in the mid- to late 1970s to a rank of 22th in
the mid-1980s to 12th in the mid-1990s in post-weed
control surveys in western Canada (Thomas and Wise
1983c; Thomas 1991; Thomas and Donaghy 1991;
Thomas et al. 1996). The relative abundance of a
weed species is calculated from the relative frequency,
relative uniformity, and relative density of a species
(Thomas et al. 1996). In volunteer canola, all three
parameters have increased substantially over the past
two decades (Thomas 1991; Thomas and Donaghy
1991; Thomas et al. 1997). However, the relationship
between volunteer canola frequency observed in sur-
veys and the canola acreage grown in the previous year
was tenuous over this period (Table 1), suggesting that
factors such as environment, tillage practices (Gray
et al. 1996) and increased acreages of crops in
which volunteer canola is more difficult to control
(e.g., pulse crops) (Thomas and Wise 1983c; Thomas
et al. 1996) also contributed to the occurrence of
volunteer canola.
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L. 955

Table 1. The infestation index for Brassica napus and/or B. rapa (volunteer canola) in annual field crops in the prairie provinces based on weed surveys
conducted since 1976

Survey Area of canola as percentage of total crop area in Frequency of volunteer canola in Canola infestation index
Province year year previous to surveyz (%) non-canola fieldsy (%) (frequency/area)x

Manitoba 1978 6.1 2.3 0.38

1979 12.3 6.5 0.53
1981 8.6 2.9 0.34
1986 9.9 5.0 0.50
1997 16.3 13.7 0.84
2002 21.0 19.2 0.91
Saskatchewan 1976 7.7 6.4 0.83
1977 3.2 5.8 1.81
1978 6.0 3.5 0.58
1979 11.1 3.9 0.35
1986 9.8 5.1 0.52
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1995 20.5 15.7 0.76

2003 12.1 8.0 0.66
Alberta 1987
1989 17.4 17.9 1.03
1997 16.9 10.7 0.63
2001 20.4 15.2 0.74

z
Crop acreage data for each year were obtained from Statistics Canada, Field Crop Reporting Series, November Estimates of Principal Field Crops,
Catalogue No. 22
002-XIE.
y
Unpublished data derived from the various Weed Survey Series Publications listed in Table 3.
x
The canola infestation index is the ratio of the frequency (%) of volunteer canola in surveyed fields of other annual crops and the relative area of
canola grown the year previous to the survey. The value of the index would be constant if the incidence of volunteer canola in fields did not change.

In the United Kingdom, Lutman et al. (1996) et al. 1986; Moyer and Huang 1997) and elsewhere
For personal use only.

summarized data from a number of field studies that
investigated competition of several crops with oats
(Avena fatua L.). A relative damage coefficient based
on dry weights was about equal in domesticated winter
B. napus (domesticated B. napus that is planted in the
autumn and grown as a winter-annual crop) and oats.
The same index was greater ( less competitive) in
barley (Hordeum vulgare L.), flax (Linum usitatissimum
L.), and field pea (Pisum sativum L.) and was lower
( more competitive) in faba beans (Vicia faba L.),
which were more competitive than oats. In Chile, yield
loss in field pea of 2.3% was observed at a volunteer B.
napus density of 1 plant m 2 and the economic thresh-
old for using herbicides to control volunteer B. napus
was estimated at 2 plants m 2 (Diaz et al. 1994). In
fields in western Canada, densities greater than this are
often observed several years after the last canola crop
has been grown (see Section 5c), although economic
thresholds for Canadian conditions have not been
estimated.
Brassica napus and B. rapa produce allelopathic
compounds with detrimental effects of Brassica crop
herbage or residue on germination of other crops. Vera
et al. (1987) compared the growth of barley, oats, wheat
(Triticum aestivum L.), flax and B. rapa following soil
incorporation of B. rapa, B. napus, and B. juncea
herbage (244 kg ha1) and found consistent reductions
in seedling counts and final yield in all crops, while the
incorporation of 5000 kg ha1 of Brassica crop residue
had no impact on seedling counts and final crop yield.
The allelopathic effect was the same for B. napus and
B. rapa. A number of studies in Canada (Mason-Sedun
(Vaughn and Boydston 1997; Peterson et al. 2001)
have found that water soluble and volatile compounds
were responsible for allelopathic effects caused by fresh
tissue of B. napus and B. rapa. However, leachates from
B. napus and B. rapa residues had the least potent
allelopathic potential among the Brassica species studied
(Mason-Sedun et al. 1986). Leachates from domesti-
cated spring B. napus residue were more potent to
stinkweed (Thlaspi arvense L.), flixweed (Descurainia
sophia L.), and downy brome (Bromus tectorum L.) than
to winter wheat (Triticum aestivum L.), suggesting the
possibility of selective management of these weeds in no-
till, direct-seeded farming systems in Alberta (Moyer
and Huang 1997). Water soluble and volatile isothio-
cyanates have been identified as allelopathic compounds
(Vaughn and Boydston 1997; Petersen et al. 2001).
Potency of isothiosyanates is dependent on form at
each phase. For example, in aqueous extracts, aryl-
isothiocyanate from domesticated winter B. rapa was
the most potent inhibitors of germination in several
species [T. aestivum L., Sonchus asper (L.) Hill, Matri-
caria inodora L., Amaranthus hybridus L., Echinochloa
crus-galli (L.) Beauv., Alopecurus myosuroides Huds.]
(Petersen et al. 2001), while allyl-isothiocyanate and
methyl-isothiocyanate were the most active allelopathic
compounds in the volatile phase and consistently
provided I50 values (the concentration at which 50%
of germination is inhibited) for germination of soybean
[Glycine max (L.) Merr.], corn (Zea mays L.), canola,
alfalfa (Medicago sativa L.), dandelion (Taraxacum
officinalis Weber), cucumber (Cucumis sativus L.), and
wheat (Triticum aestivum L.) at concentrations below
 956 CANADIAN JOURNAL OF PLANT SCIENCE
1 ppm. In soil, the persistence of isothiocyanates from
B. rapa was short (48 h) (Petersen et al. 2001).
Pollen produced by Brassica sp. may elicit allergic
reactions, such as bronchial asthma, in humans (Bucur
and Arner 1978; Welch et al. 2000). The proportion of
the population that is sensitive to airborne Brassica
allergens is relatively small with 9% of people reporting
allergic symptoms during flowering season of B. napus
in Scotland (Soutar et al. 1995); however, allergy skin
tests suggested that 23 to 43% of the population have
sensitivity to Brassica pollen (Bucur and Arner 1978;
Soutar et al. 1995). It has also been suggested that the
condition of some horses with chronic obtrusive lung
may be exacerbated by exposure to B. napus during
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flowering (Soutar et al. 1995). Allergens may not be
contained in pollen only. Butcher et al. (1994) identified
22 volatile organic compounds released from B. napus
during flowering. The bulk of these comprised alpha-
farnesene, beta-myrcene, linalool, and (E)-3-hexen-1-ol
acetate and it is thought that some of these compounds
were suspected of eliciting allergic reactions. In the
United Kingdom, chronic exposure to canola flour,
which is used for animal fodder, has been implicated in
causing occupational asthma in farmers (Alvarez et al.
2001). In Finland, Poikonen et al. (2006) recently
reported increased incidence of allergic reactions in
For personal use only.
small children with atopic dermatitis who were tested
for food allergies using the skin prick test. In a follow up
study (Puumalainen et al. 2006), 9.5 to 14.5 kD napin
seed storage proteins in B. napus and B. rapa were
identified as the causal compounds.
The meal and foliage of domesticated Brassica is
used as animal fodder. Breeding efforts have reduced
erucic acid and glucosinolate levels in the meal to levels
that have eliminated the anti-nutritive effects when fed
to livestock. However, Broadway (1989) examined
trypsin proteinase inhibitors, which are thought to be
antigrazing compounds, in foliage and storage organs
of seven species of Brassicaceae, including B. napus and
B. rapa. In general, greater levels (up to 13 times) of
trypsin inhibitors were found in foliage of cultivated
Brassica spp. than in the foliage of wild Brassica spp.
Trypsin inhibition in B. napus subsp. oleifera was
652 mg mg1 protein and that of B. rapa ranged
from 322 (subsp. rapifera) to 1220 mg mg 1 protein
(subsp. chinensis).
Genes from herbicide-resistant (HR) volunteers of B.
napus and B. rapa may directly be transferred to non-
HR genotypes grown as crops in the same or neighbour-
ing fields (Hall et al. 2000; Rieger et al. 2002; Beckie
et al. 2003). Currently, more than 150 spring B. napus
genotypes with diverse traits are licensed for commercial
production in Canada (Canadian Food Inspection
Agency 2008). Future planned releases include geno-
types with an even more diverse range of properties.
A risk of these traits crossing the species barrier to more
distantly-related wild relatives, S. arvensis, Erucastrum
gallicum Willd., and Raphanus raphanistrum, exists, but
is considered small (Warwick et al. 2003). However,
successful transfer and introgression of such novel genes
to B. rapa may result in populations that may be more
difficult to control (Section 9).
Volunteer B. napus and B. rapa can serve as reservoir
or alternate hosts to insect, nematode, fungal, viral, and
bacterial pests. These pests cause damage to these
species when grown as crops or may cause damage to
other cruciferous vegetable crops (Section 13).
(b) Beneficial * The annual acreage seeded to canola
(B. napus and B. rapa) in Canada has ranged between
3.6 and 5.2 million ha in recent years (2001
2005), with
the vast majority grown in western Canada (Canola
Council of Canada 2006). Although that acreage was at
one point equally divided between the two species,
cultivar improvements in the past two decades have
resulted in B. napus genotypes being planted almost
exclusively. Today, both species are grown principally
for their seed oil content in Canada with total annual
seed production ranging from 4.5 to 9.7 million tonnes
per year (2001
2005) (Canola Council of Canada 2006).
Brassica napus and B. rapa were domesticated initially
for medicinal purposes (Kimber and McGregor 1995).
In some countries, B. napus and B. rapa greens continue
to form part of the human diet. Recently, the feasibility
of forage B. napus greens as an alternative vegetable to
rutagaba or swede turnip greens (B. napus var. rapifera
L.) was investigated in Newfoundland (Spaner and Lee
2001). Forage B. napus was visually more appealing
than the traditional turnip tops, however, a taste panel
ranked B. napus last. The nutritional qualities of
B. napus and B. rapa greens in Virginia compared well
with those obtained from a national data base for turnip
(B. rapa subsp. rapa) and mustard (B. juncea) (Bhardwaj
et al. 2003).
In some countries, the long-time use of B. napus and
B. rapa for grazing or silage continues (Bell 1995).
Domesticated B. napus has high in vitro digestibility
with digestible protein levels comparable to alfalfa
(Wilman et al. 1996) and lambs grazed on forage
B. napus in Australia had higher rates of gain than
lambs grazed on irrigated perennial pasture (Hopkins et
al. 1995). In cattle, however, high levels of B. rapa forage
may influence products derived from milk. Milk thio-
cyanate content may increase to undesirable levels and
cause problems with the manufacture of Cheddar cheese
(Moate et al. 1996).
In the Middle Ages, the use of domesticated B. napus
and B. rapa broadened when oil extracted from the seeds
(approximately 45% of seed dry weight) was used as
lamp oil, for making soap and in rare cases, as a cooking
oil (Kimber and McGregor 1995). Seed oil use was
further expanded upon arrival of the steam engine where
it was used as a lubricating oil on water-washed parts.
Brassica napus and B. rapa were first grown commer-
cially in Canada in the early 1940s for industrial oil
(National Research Council of Canada 1992). To solve
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
the rape oil supply problem caused by the war,
domesticated B. napus and B. rapa seed were sourced
from Argentina and Poland, respectively. Intensive
breeding efforts undertaken in Canada in the 1970s
resulted in modified seed oil quality, which reduced
potential health risks to humans and removed anti-
nutritional compounds in the seed meal allowing its safe
use as a protein supplement in animal rations (see
Section 7c). Although dark seeded genotypes are grown
almost exclusively in Canada at this time, yellow seeded
B. napus is higher in seed oil and lower in chlorophyll
content (Rakow and Raney 1993) than dark seeded
genotypes. Meal from yellow seeded B. napus and
B. rapa also tended to have greater seed protein (0.6
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22
to 3.7%) and seed sucrose (1.4 to 2.8%) (Simbaya et al.
1995). Dietary fibre was correlated negatively to protein
content, while most other nutritional parameters were
not different between seed coat colour classes in the
latter study.
Brassica napus and B. rapa are valuable nectar and
pollen sources for honey production. In Saskatchewan,
the frequency of pollen from Brassicaceae (B. napus and
B. rapa) in honey samples was only second to that of
Melilotus sp. (Feller-Demalsy et al. 1987). Pollen from
Brassicaeae was absent in only 1 of 42 honey samples
and was the major source of pollen in many samples.
For personal use only.
The relative proportion of Brassicaceae pollen in honey
samples in Saskatchewan was lower in a previous study
(Louveaux 1966) which may be reflective of the lower
annual acreage seeded to domesticated B. napus and
B. rapa at the time (Canola Council of Canada 2008). In
a recent study, the steroid fraction of B. rapa bee pollen
was shown to induce apotosis in human prostrate cancer
cells (Wu and Lou 2007).
In the Netherlands, forage B. napus genotypes
have been used in soil reclamation and to prevent
out-of-season nitrogen (N) leaching (Vos and van der
Putten 1997; Vos et al. 1998). Forage B. napus has also
been grown effectively for phytoremediation of selenium
(Se) (Ba_uelos and Mayland 2000) and molybdenum
(Mo) (McBride et al. 2000). Brassica napus grown for Se
phytoremediation may be used in animal rations as long
as the Se content is known and the forage B. napus
content of the ration is adjusted to not exceed recom-
mended threshold levels. Selenium uptake by B. napus is
much greater when Se is available as inorganic selenate
rather than seleniforous organic material (Ajwa et al.
1998). Similarly, B. napus grown for Mo phytoremedia-
tion may be used as animal fodder as long as the Cu:Mo
ratio is not below 2:1 (McDowell 1985).
After oil extraction, B. napus seed meal may be used
as a natural pesticide (Bhardwaj et al. 1996); however,
only if the meal is high in glucosinolates. Glucosinolates
are sulphur-containing organic anions containing b-d-
thioglucose groups that are present in leaf tissue and
seed in several different forms (Mithen 1992). These
compounds are thought to be grazing deterrents, and
herbivore damage has been shown to induce elevated
957
leaf glucosinolate levels in these species (Cipollini et al.
2003). In soil, glucosinolate metabolites from B. napus
meal, such as isothiocyanate and thiocyanate, appear to
have greater biological activity than glucosinolates as
shown by a wireworm (Limonius infuscatus Mots.)
bioassay (Brown et al. 1991). Siemens et al. (2002)
investigated the cost of defence against herbivory (i.e.,
glucosinolate production) in weedy B. rapa during
interference with Lolium perenne L., a generalist compe-
titor, after developing divergent lines from a natural
population through artificial selection. Under competi-
tion with L. perenne, growth rates were the same in high
and low glucosinolate lines, indicating little cost to
defence against herbivory under competition. In the
absence of interference, however, herbivore exposure
decreased growth rates most prominently in low gluco-
sinolate lines (20%), while the growth rate of high
glucosinolate lines decreased by only 8% in the same
treatment. Furthermore, L. perenne root expansion was
inhibited most strongly by glucosinolate metabolites.
The importance of tissue glucosinolate content to
resistance to pests of B. napus and B. rapa is less clear
(Giamoustaris and Mithen 1997; Stowe 1998; Sexton
et al. 1999). The relationship between glucosinolates and
pest and disease resistance was reviewed in Mithen
(1992). A correlation between the type of glucosinolate
compound and maturity class in vegetable B. rapa has
been reported (Kim et al. 2003) where, in the early-
maturing class, the relative proportion of gluconapin
was greater than that of glucobrassicanapin. The
opposite was true for medium- and late-maturing types.
(c) Legislation * Bird rape (as B. campestris) is
classified as a noxious weed in Quebec weed legislation
(Anonymous 1977) and in Saskatchewan (Anonymous
1999). However, B. napus is not listed in the noxious
weeds acts in any province or territory in Canada.
4. Geographical Distribution
Bird rape*Bird rape has been reported from every
province and territory in Canada (NT YK NF NS PE
NB QC ON MB SK AB BC) (Fig. 2 upper map;
Warwick et al. 2000; Darbyshire 2003), but is only
established as a naturalized weed in the Maritime
Provinces, the Eastern Townships of Quebec, and the
Fraser Valley and Vancouver Island of British Colum-
bia (Mulligan 2002b; Simard et al. 2007). In Eastern
Canada, bird rape is most common in Quebec, notably
weedy in the Estrie region (Sabourin et al. 1992;
Simard et al. 2007), where it was reported in 15.6
and 11.3% of the cereal fields in the Nicolet and La
Pocatière Regions, respectively (Doyon et al. 1982),
and in some parts of the east supplanted wild mustard
over large areas (Frankton and Mulligan 1987). Bird
rape is less frequent in Ontario, reported as occurring
in a few grain fields and waste areas in southern
Ontario (Alex 1992), but was not found in recent
surveys conducted in 2000
2005 in this region
(P. Mason, personal communication). It is scattered
 958 CANADIAN JOURNAL OF PLANT SCIENCE
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22
For personal use only.

Fig. 2. Distribution of Brassica rapa (upper map) and B. napus (lower map) in Canada from 218 and 47 herbarium specimens,
respectively, examined by the senior author in CAN, DAO, GH, NFLD, QFA, and VPI. Herbarium abbreviations as in Holmgren
et al. (1990).

throughout temperate North America, and has been
reported from almost every US state, including Alaska,
Greenland, and highland areas of Mexico and Central
America (Rollins 1981, 1993; Warwick et al. 2000;
Darbyshire 2003). Maps of B. rapa distribution are
available for eastern Canada (Sabourin et al. 1992;
Simard et al. 2007), Europe (see Jalas et al. 1996) and
the world (Hultén and Fries 1986; Holm et al. 1997).
Brassica rapa occurs as a weed in nearly every country
(i.e., more than 20 crops in over 50 countries) in the
northern hemisphere with temperate climates, several
Asian countries, through the Andean zone of South
America and Argentina and Brazil, and in Australia and
New Zealand (Holm et al. 1997, p. 117).
Staniforth and Scott (1990) compared the abundance
and distribution of 106 introduced species around the
northern port of Churchill, Manitoba, based on floral
surveys conducted in 1959 and 1989, and categorized
B. rapa as a persistent introduction requiring a contin-
uous supply of introduced seed, and not established
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L. 959

Table 2. Frequency, density and relative abundance rank of Brassica napus and/or B. rapa in non-canola crops in the major ecoregions in the Prairie
Provinces based on the most recent weed surveys conducted in Alberta in 2001, Manitoba in 2002 and Saskatchewan in 2003 (Leeson et al. 2005). Weed
populations were assessed in July and August, after the time of in-crop herbicide application and before harvest

Ecoregionz Predominant soil type Number Frequencyy Densityx Relative Rank

of fields (%) (no. m 2) abundancew

Peace Lowland Grey Luvisol 127 26.7 4.6 9.4 12

Boreal Transition Grey Luvisol, Dark Grey Chernozem 372 23.6 4.9 10.2 12
Interlake Plain Dark Grey Chernozem 48 14.2 0.9 4.4 15
Lake Manitoba Plain Black Chernozem 167 16.0 5.4 8.3 9
Aspen Parkland Black Chernozem 1058 14.6 4.7 7.0 8
Fescue Grassland Black Chernozem 73 11.0 1.5 6.4 14
Moist Mixed Grassland Dark Brown Chernozem 701 6.4 4.2 4.6 12
Mixed Grassland Brown Chernozem 586 1.5 3.1 1.2 22

z
Prairie ecoregions are described and mapped in Ecological Stratification Working Group (1995).
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22

y
Percentage of surveyed fields in which the species was present.
x
Mean number of plants in fields in which the species was present.
w
Relative abundance is a synthetic index based on frequency, field uniformity and density and is used to rank species abundance. A low number
indicates a high rank relative to other species (Thomas 1985).

(i.e., able to set viable seeds and maintain their popula-
tions) in the area.
Brassica napus (rapeseed, canola) was also introduced
from Eurasia. The species has been collected as a
sporadic weedy escape from cultivation in cultivated
and abandoned fields, roadsides, railways and waste
places in all provinces and territories (NT NF NS PE
For personal use only.
Canada. Brassica rapa requires approximately 10%
fewer growing degree-days to reach maturity.
(b) Substratum * Volunteer canola is not restricted to
any particular soil type and has been found on Brown,
Dark Brown, Black and Dark Grey Chernozemic and on
Grey Luvisol soils in the Prairie Provinces (Table 2). Soil

NB PQ ON MB SK AB BC), except the Yukon and
almost every US state (Fig. 2 lower map; Rollins 1981,
1993; Warwick et al. 2000; Mulligan 2002b). Volunteer
B. napus is particularly common in canola-growing
areas of western Canada, which is not reflected in the
lower map of Fig. 2.
5. Habitat
(a) Climatic Requirements * Volunteer B. napus and/or
B. rapa have been found in most of the annual field
crops grown in the major ecoregions of Manitoba,
Saskatchewan and Alberta (Table 2). The continental
climate of this region has long, cold winters and short,
warm to hot, summers (Ecological Stratification Work-
ing Group 1995). Mean summer temperatures range
from 13.08C in the Peace Lowland ecoregion to 16.08C
in the Lake Manitoba Plain ecoregion. Mean winter
temperatures range from 14.58C in the Interlake Plain
ecoregion and to 8.08C in the Fescue Grassland
ecoregion. Mean annual precipitation ranging from
250 mm in the Mixed Grassland ecoregion to 700 mm
in the Lake Manitoba Plain ecoregion. Volunteer canola
is least frequent in the Moist Mixed and Mixed Grass-
land ecoregions, where moisture deficits in late summer
are caused by low precipitation and high evapotran-
spiration. Domesticated B. napus requires an average of
1560 growing degree-days (ranging from 1380 to 1718),
with a 08C base temperature, to reach maturity based on
5 yr of trials across western Canada (Thomas 2003).
Longer daylight hours partially compensate for lower
temperatures in northern production areas of western
characteristics most suitable for the production of
domesticated canola are described in the Canola Growers
Manual (Thomas 2003). Medium-textured soils on
the prairies are considered best because of good moisture
infiltration and water-holding capacity and adequate
drainage. Fine-textured soils have a good water-holding
capacity, but can become waterlogged and tend to remain
wet and cold in the spring. Sandy soils are usually not
suitable for canola production because of low moisture-
holding capacity. Yields of domesticated canola are
reduced on acid soils with a pH of less than 5.5, on alkali
soils with a pH of more than 8.3, on saline soils with
electrical conductivity values greater than 6 dS m2, and
on waterlogged soils after 3 d.
(c) Communities in Which the Species Occurs * Alex
(1966) conducted a survey of 45 weeds of cultivated land
in the three Prairie Provinces from 1963 to 1965.
Although B. napus and B. rapa were not included among
the selected weeds, about 1.0% of the survey respondents
added domesticated rapeseed as a weed of concern. Since
then, volunteer canola has been found more frequently in
weed surveys of annual and perennial crops conducted
during the past 30 yr in the prairie region (Table 3).
Unfortunately, the two species were not differentiated in
these surveys but it is very likely that almost all of the
volunteers recorded in the 2000s surveys were B. napus
because this species accounts for 95% of the seeded crop.
Based on relative abundance, volunteer canola ranked as
high as second place in spring wheat and barley fields
 960 CANADIAN JOURNAL OF PLANT SCIENCE
Table 3. Frequency, density, relative abundance and rank of Brassica napus and/or B. rapa in annual field crops and perennial grasses and legumes in the Prairie Provinces based on weed surveys
conducted since 1973. Weed populations were assessed in July and August, after the time of in-crop herbicide application and before harvest

Province/cropz Year Number of fields Frequencyy (%) Densityx (no. m 2) Relative abundancew Rank Source

British Columbia (Peace River Region)

Barley 1978
1980 420 38.6 9.2 12.2 9 Thomas and Wise (1983a)
Spring wheat 1978
1980 157 43.3 15.7 15.0 8 Thomas and Wise (1983a)
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Oats 1978
1980 151 26.5 7.0 6.5 12 Thomas and Wise (1983a)
Forage 1978
1980 372 14.0 7.3 3.7 18 Thomas and Wise (1983b)
Alberta (Fort Vermilion Area)
Barley 1985 55 70.8 7.3 37.0 2 Thomas et al. (1986)
Spring wheat 1985 24 74.5 6.0 40.1 2 Thomas et al. (1986)
Alberta
Barley 1987
1989 383 23.0 3.7 6.0 15 Maurice et al. (1990)
Barley 1997 220 10.5 9.6 5.7 19 Thomas et al. (1998a)
Barley 2001 349 12.9 4.5 6.1 15 Leeson et al. (2002b)
Cereals/oilseeds 1973
1977 3109 10.4 46.6 4.7 12 Thomas and Wise (1985)
Oats 1987
1989 125 12.8 4.8 2.5 27 Maurice et al. (1990)
Oats 1997 36 16.7 18.2 8.5 11 Thomas et al. (1998a)
Oats 2001 86 14.0 7.0 6.4 14 Leeson et al. (2002b)
Peas (dry) 2001 40 50.0 3.4 15.0 7 Leeson et al. (2002b)
For personal use only.

Wheat (spring/durum) 1987
1989 411 15.6 5.8 5.3 14 Maurice et al. (1990)
Wheat (spring) 1997 220 11.8 2.9 5.0 20 Thomas et al. (1998a)
Wheat (spring) 2001 418 16.0 5.2 10.0 10 Leeson et al. (2002b)
Wheat (durum) 1997 29 (species not recorded) Thomas et al. (1998a)
Wheat (durum) 2001 71 1.4 0.4 0.8 20 Leeson et al. (2002b)
Wheat (winter) 1987
1989 22 4.3 0.4 1.7 26 Maurice et al. (1990)
Saskatchewan
Alfalfa (seed) 1989 42 11.9 1.0 2.5 24 Malik et al. (1991)
Alfalfa (seed) 1997 120 7.5 1.5 1.5 28 Myhre et al. (2002)
Barley 1976
1979 600 9.0 4.0 3.1 22 Thomas and Wise (1983c)
Barley 1986 119 10.1 3.3 4.0 17 Thomas and Wise (1987)
Barley 1995 161 17.4 3.1 6.7 11 Thomas et al. (1996)
Barley 2003 355 11.8 11.2 9.9 6 Leeson et al. (2003)
Canary seed 2003 46 15.2 0.8 6.9 11 Leeson et al. (2003)
Cereals (organic) 2002 73 (species not recorded) Buhler (2005)
Flax 1976
1979 75 8.0 1.9 2.1 24 Thomas and Wise (1983c)
Flax 1986 27 18.5 8.3 13.7 9 Thomas and Wise (1987)
Flax 1995 53 17.0 1.4 4.3 17 Thomas et al. (1996)
Flax 2003 93 5.4 0.6 1.9 26 Leeson et al. (2003)
Lentils 1985 83 3.6 1.4 1.2 20 Douglas and Thomas (1986)
Lentils 1995 34 14.7 1.5 5.2 14 Thomas et al. (1996)
Lentils 2003 73 5.5 0.6 2.3 21 Leeson et al. (2003)
Mustard 1985 107 0.9 0.2 0.1 65 Douglas and Thomas (1986)
Mustard 1995 25 (species not recorded) Thomas et al. (1996)
Mustard 2003 45 2.2 0.6 0.8 35 Leeson et al. (2003)
Oats 1976
1979 233 8.2 5.2 2.2 23 Thomas and Wise (1983c)
Oats 1986 24 4.2 0.2 0.8 35 Thomas and Wise (1987)
Oats 1995 47 12.8 9.0 6.0 13 Thomas et al. (1996)
Oats 2003 119 10.9 3.4 3.8 21 Leeson et al. (2003)
Peas (dry) 1985 30 30.0 1.5 5.7 16 Douglas and Thomas (1986)
Peas (dry) 1995 60 40.0 7.2 24.3 4 Thomas et al. (1996)
Peas (dry 2003 104 14.4 2.6 6.3 11 Leeson et al. (2003)
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Table 3 (Continued)
Province/cropz Year Number of fields Frequencyy (%) Densityx (no. m 2) Relative abundancew Rank Source

Rye (spring/fall) 1976
1979 38 (species not recorded) Thomas and Wise (1983c)
Sunflowers 1985 50 2.0 2.4 1.3 26 Thomas and Wise (1986)
Wheat (spring/durum) 1976
1979 3164 4.0 2.7 1.2 28 Thomas and Wise (1983c)
Wheat (spring/durum) 1986 876 4.1 6.2 2.4 21 Thomas and Wise (1987)
Wheat (spring) 1995 407 17.0 5.4 9.0 8 Thomas et al. (1996)
Wheat (spring) 2003 581 7.4 3.2 4.3 14 Leeson et al. (2003)
Wheat (durum) 1995 164 5.5 8.2 4.7 14 Thomas et al. (1996)
Wheat (durum) 2003 274 2.2 3.1 1.9 18 Leeson et al. (2003)
Wheat (winter) 1985
1988 1050 3.9 3.9 2.1 27 Thomas and Wise (1989)
Manitoba
For personal use only.

Alfalfa (seed) 1983 77 (species not recorded) Goodwin et al. (1985)

Barley 1978
1979, 1981 355 2.5 5.2 0.8 32 Thomas and Wise (1984)

GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.

Barley 1986 114 3.5 0.9 0.8 30 Thomas and Wise (1988)
Barley 1997 71 7.0 0.8 1.6 24 Thomas et al. (1998b)
Barley 2002 79 15.2 3.2 7.4 10 Leeson et al. (2002a)
Flax 1978
1979, 1981 160 8.1 2.0 1.9 19 Thomas and Wise (1984)
Flax 1986 60 1.7 2.4 0.6 30 Thomas and Wise (1988)
Flax 1997 33 24.2 2.9 8.5 9 Thomas et al. (1998b)
Flax 2002 37 10.8 4.0 5.3 14 Leeson et al. (2002a)
Oats 1978
1979, 1981 92 1.1 1.0 0.2 55 Thomas and Wise (1984)
Oats 1986 25 (species not recorded) Thomas and Wise (1988)
Oats 1997 71 3.6 2.8 1.2 21 Thomas et al. (1998b)
Oats 2002 112 15.2 2.2 4.9 13 Leeson et al. (2002a)
Wheat (spring) 1978
1979, 1981 629 3.8 4.9 1.0 28 Thomas and Wise (1984)
Wheat (spring) 1986 262 6.9 2.5 2.0 20 Thomas and Wise (1988)
Wheat (spring) 1997 211 15.6 4.2 5.0 16 Thomas et al. (1998b)
Wheat (spring) 2002 248 23.8 2.9 9.8 7 Leeson et al. (2002a)

z
Data are provided for province and crop combinations with more than 20 surveyed fields.
y
Percentage of surveyed fields in which the species was present.
x
Mean number of plants in fields in which the species was present.
w
Relative abundance is a synthetic index based on frequency, field uniformity and density and is used to rank species abundance. A low number indicates a high rank relative to other
species (Thomas 1985).

961
 962 CANADIAN JOURNAL OF PLANT SCIENCE
surveyed in 1985 in the Fort Vermilion area of Alberta, as
low as 55th in oat fields in the late 1970s surveys in
Manitoba, and absent in some crops in some years. The
frequency of occurrence was generally B20% of the
surveyed fields and densities were B10 plants m 2, but
frequencies were as high as 70% and densities were as
high as 18.2 plants m 2 in some surveys. Densities were
much higher (46.6 plants m 2) when the surveys were
conducted prior to the application of in-crop herbicides
from 1973
1977 in Alberta.
A series of surveys in various regions and crops of
southern Ontario between 1960 and 1990 did not record
any occurrences of B. napus or B. rapa (Alex 1964;
Richards 1979; Hamill et al. 1983; Hamill and Thomas
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22
1985; Frick et al. 1990). Brassica rapa was found during
extensive surveys of 4854 fields throughout Quebec in
1980
1984. The species was present in 27% of the cereal
(wheat, barley and oats), 7% of the corn, 6% of the
strawberry, 2% of the hay and pasture, and 2% of the
fruit and vegetable fields (Doyon et al. 1986a, b, c, d, e;
Lemieux et al. 1988a, b, c). Only B. campestris (B. rapa)
was found in 0.5% of the spring cereal (187) fields in
New Brunswick in 1986
1987 (Thomas et al. 1994).
Brassica campestris (B. rapa) was found in 4.1% of the
cereal fields in Prince Edward Island in 1978, but neither
species was recorded during a survey of the same crops
For personal use only.
in 1979 (Ivany 1980; Ivany and Thomas 1983). Weeds
were counted during June, July and August so that a
portion of the observations was made before in-crop
herbicide applications. Neither species was recorded
during a Nova Scotia survey of corn and barley in
July and August and strawberries in October in 1999
(R. Hoeg and A. G. Thomas, unpublished data).
In annual cereal, oilseed and pulse crops in the Prairie
Provinces, 93 weed species occurred in fields with
volunteer canola. Twenty-nine of these species were
present in greater than 4% of the fields (Table 4). These
29 species along with volunteer canola account for 94%
of the total relative abundance in these fields. All of
these species commonly occur in surveyed fields
throughout the prairies and the weed community is
typical of that found in the ecoregions where canola
is grown as a crop (Leeson et al. 2005). Three of the
species associated with volunteer canola were annual
grasses that ranked second (wild oats, Avena fatua L.),
fourth [green foxtail, Setaria viridis (L.) P. Beauv.] and
twentieth [barnyard grass, Echinochloa crus-galli (L.)
P. Beauv.] in terms of relative abundance. Two addi-
tional grasses were the volunteer wheat and volunteer
barley, ranking 21st and 28th, respectively. Six species
were perennial, with Canada thistle [Cirsium arvense (L.)
Scop.] ranking fifth and occurring in 59.2% of the fields
with volunteer canola. Five species were facultative
winter annuals with stinkweed (Thlaspi arvense L.) being
the most frequent among these (29.8%). The remaining
species were spring-germinating annuals with wild
buckwheat (Polygonum convolvulus L.) being the most
frequently occurring (68.5%) of all the associated
species and chickweed [Stellaria media (L.) Vill.]
had the highest density at 19.6 plants m 2. In the
United Kingdom, weedy B. napus has been reported in
woodland communities (Dixon et al. 2006).
6. History
Brassica napus and B. rapa are native of Eurasia and
introduced into North America as either a seed con-
taminant or crop escape (Rousseau 1968; Rollins 1981;
Frankton and Mulligan 1987; Crompton et al. 1988;
Rollins 1993; Mulligan 2002b; Darbyshire 2003). Bras-
sica rapa was reported in Nova Scotia as early as 1829
and first present in Quebec in 1908 (Rousseau 1968),
whereas B. napus was first collected in New Brunswick
in 1874 (Mulligan 2002b). Neither species is listed in
early North American floras (Torrey and Gray 1838;
Gray 1848), but both are listed in later floras (Britton
and Brown 1897; Robinson and Fernald 1908). Eur-
opean origins for both species were suggested in the
latter two floras. Britton and Brown (1897) reported
B. rapa as occurring on cultivated ground, sometimes
persisting for a year or two and occasionally in waste
places eastward in the northern United States and
Canada and that B. napus is sometimes found in waste
places. Robinson and Fernald (1908) stated that both
species tended to escape from or persist after cultivation
and are often noxious weeds.
In contrast, Arnason et al. (1981) suggested an earlier
introduction and described the extensive use of various
Brassica and Sinapis spp. for food and medicine by
native peoples of eastern Canada. Brassica napus bark
was used to treat colds, cough, grippe and small pox by
the Micmac and used for chilbains by the Rappahan-
nock. Brassica rapa was used as medicine by the Bois
Fort Chippewa.
7. Growth and Development
(a) Morphology * Brassica napus and B. rapa have an
indeterminate growth habit. Winter B. rapa crops
exhibit greater frost tolerance than domesticated winter
B. napus. This is thought to be the result of increased
protection of the growing points in winter B. rapa by
the more compact arrangement of the foliage of the
surrounding rosette [Trossell (1959) cited by Mendham
and Salisbury (1995)]. Both species produce seeds in
siliques (pods) that shatter easily at maturity. In general,
seed shatter potential is higher in B. napus than in
B. rapa, B. juncea and S. alba; however, differences
among genotypes within species also exist (Morgan et al.
2000; Squires et al. 2003; Wang et al. 2007). In part, the
difference in shatter susceptibility between these species
may be due to differences in the pattern of water loss
from the pericarp and surrounding tissues (Squires et al.
2003). Water loss occurs from the inside of the pericarp
in B. napus and from the outside of the pericarp
in B. rapa. In B. napus pods, the dehiscent zone
is composed of simple cells without further wall
thickening (Meaking and Roberts 1990). Genes from
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L. 963

Table 4. Frequency, density, relative abundance and rank of species occurring in the 377 fields with volunteer canola (Brassica napus and/or B. rapa) in
the Prairie Provinces. Unpublished data derived from the most recent weed surveys conducted in Alberta in 2001, Manitoba in 2002 and Saskatchewan in
2003 (Leeson et al. 2005). Weed populations were assessed in July and August, after the time of in-crop herbicide application and before harvest

Species Lifez cycle Frequencyy (%) Densityx (no. m 2) Relative abundancew Rank

Brassica napus L. and B. rapa L. Sa 100.0 4.5 34.8 1

Avena fatua L. Ag 57.9 12.0 32.6 2
Polygonum convolvulus L. Sa 68.5 4.5 27.3 3
Setaria viridis (L.) P. Beauv. Ag 36.2 19.2 27.1 4
Cirsium arvense (L.) Scop. P 59.2 2.5 17.5 5
Stellaria media (L.) Vill. Sa 20.8 19.6 16.0 6
Thlaspi arvense L. Wa 29.8 8.2 13.7 7
Galium aparine L. and G. spurium L. Sa 23.7 8.2 11.2 8
Chenopodium album L. Sa 33.3 3.1 10.3 9
Sonchus asper (L.) Hill Sa 20.1 7.9 8.8 10
Taraxacum officinale Weber in F. H. Wigg. P 31.3 1.7 8.0 11
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Galeopsis tetrahit L. Sa 19.1 6.8 8.0 12

Equisetum arvense L. P 18.7 4.4 6.5 13
Sonchus arvensis L. P 24.3 2.1 6.3 14
Capsella bursa-pastoris (L.) Medik. Wa 16.1 5.6 6.2 15
Crepis tectorum L. Wa 18.7 3.2 5.6 16
Polygonum lapathifolium L. Sa 20.6 2.2 5.5 17
Elytrigia repens (L.) Desv. ex B. D. Jacks P 15.2 4.7 5.0 18
Amaranthus retroflexus L. Sa 13.5 6.2 5.0 19
Echinochloa crusgalli (L.) P. Beauv. Ag 7.8 13.7 4.3 20
Triticum aestivum L. Ag 10.3 4.9 3.7 21
Matricaria discoidea D. C. Sa 6.9 10.4 3.1 22
Sinapis arvensis L. Sa 12.6 1.7 3.0 23
Silene noctiflora L. Wa 7.9 1.9 2.1 24
Senecio vulgaris L. Sa 6.3 3.6 2.0 25
Kochia scoparia (L.) Schrad. Sa 7.4 1.8 1.9 26
For personal use only.

Malva pusilla Sm. Sa 6.5 2.2 1.8 27

Hordeum vulgare L. Ag 5.0 2.9 1.6 28
Trifolium spp. P 4.3 2.6 1.3 29
Lappula squarrosa (Retz.) Dumort. Wa 4.8 2.3 1.3 30

z
Life cycles codes: Agannual grass, P perennial, Sa spring-germinating annual broadleaf, Wafall-germinating annual broadleaf.
y
Percentage of surveyed fields in which the species was present.
x
Mean number of plants in fields in which the species was present.
w
Relative abundance is a synthetic index based on frequency, field uniformity and density and is used to rank species abundance (Thomas 1985).

more shatter-resistant Brassicaceae, including B. rapa,
are being incorporated into the domesticated B. napus
genome (Morgan et al. 2000, Wang et al. 2007). These
genes acted recessively, showed high heritability, and did
not appear to be linked to agronomically important
traits (Morgan et al. 2000). Reduced seed shatter would
likely result in decreased volunteer B. napus populations.
In undisturbed populations, seeds from shattered
pods are dropped near the parent plant. When grown
as a crop, large numbers of seeds may be added to the
seed bank at the time of harvest (Section 8), while
combine harvesters also may substantially increase the
distance of seed dispersal within and between fields.
In a laboratory study, domesticated B. napus and
B. rapa formed nodule-like structures when inoculated
with broad host-range Rhizobium strains isolated from
Lablab purpureus (L.) Sweet and Phaseolus vulgaris L.
(Trinick and Hadobas 1995). Bacteria were found on the
root epidermis, but could not be detected inside root
tissue which when ground was toxic to the bacteria.
Dunfield and Germida (2001) investigated the rhizo-
sphere and root interior microbial communities asso-
ciated with domesticated B. napus genotypes using fatty
acid methyl ester (FAME) analysis in a field experiment
in Saskatchewan. Significant differences were found
between and within transgenic (GM) glufosinate- and
glyphosate-resistant and non-GM genotypes indicating
genotype specific differences in the structure of the
microbial communities associated with B. napus. Similar
results were reported by Gyamfi et al. (2002) in GM
B. napus resistant to glufosinate and metazachlor in
Germany. Further study is required as inferences about
functional differences among the microbial communities
cannot be made from data generated by these techniques.
(b) Perennation * Brassica napus and B. rapa are annual
species with one generation per year under Canadian
conditions. In Quebec, rootstocks of B. napus may remain
alive after harvest, overwinter, and regrow as volunteers
the following year (Simard et al. 2002). Similar observa-
tions have not been reported in B. rapa.
(c) Physiological Data * The physiology of volunteer
B. napus and B. rapa and weedy B. rapa (i.e., known self-
 964 CANADIAN JOURNAL OF PLANT SCIENCE
sustaining populations of B. rapa) has not been described.
However, physiological data for domesticated genotypes
of both species are well established. The oil content of
domesticated spring B. napus and B. rapa seed tends to be
similar and ranges from 40 to 45% in western Canada
(Kimber and McGregor 1995) and for spring and winter
B. napus in the United States (Shafii et al. 1992; Butruille
et al. 1999). In domesticated B. napus, maximum seed oil
levels were reached at physiological maturity (Fowler and
Downey 1970; Rakow and McGregor 1975). Seed oil
content and seed yield tend to be greater in western
Canada (Canvin 1965; Kirkland and Johnson 2000) and
Australia (Mailer and Cornish 1987) in B. napus and
B. rapa crops when planted in the autumn or early spring
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compared with the traditional time of seeding (mid- to
late spring) while waterlogging decreased the seed oil
content in domesticated winter B. napus in the United
Kingdom (Cannell and Bedford 1980). For a review of
the environmental effects on seed oil content in domes-
ticated B. napus and B. rapa, readers are directed to
Mendham and Salisbury (1995).
In domesticated B. napus and B. rapa, fatty acids are
present primarily in the form of triacylglycerides. Fatty
acid composition determines the use of the oil and
breeding has resulted in four broad categories including
low erucic acid (LEAR), high erucic acid (HEAR)
For personal use only.
(Kimber and McGregor 1995), low linolenic acid (high
stability varieties) (Anonymous 2008a), and canola oil.
To qualify as canola, seed oil content must be less than
2% erucic acid while the aliphatic glucosinolate content
of seed meal must be below 30 mmol g 1 (Canola
Council of Canada 1990). Currently, much effort is
directed at generating genotypes with specific fatty acid
composition for specialty oil markets including indus-
trial uses and fish food (Opsahl-Ferstad et al. 2003).
This elevates concerns of potential contamination
caused by volunteer populations of B. napus and
B. rapa when genotypes of different quality character-
istics are grown without adequate separation in time and
space. Seed fatty acid composition of 1475 entries from
21 Brassica species, including B. napus and B. rapa was
characterized by Velasco et al. (1998). On average,
B. napus seed contained more C18:1 and less C22:1
than B. rapa, even though a wide range of fatty acids
was observed in both species. Mandal et al. (2002) also
compared seed oil content and fatty acid profiles in nine
Brassicaceae species and found no correlation between
seed oil content and fatty acid profiles in B. napus and
B. rapa.
Similar to seed oil content, fatty acid composition in
domesticated B. napus and B. rapa is influenced by
environmental conditions. In western Canada (Deng
and Scarth 1998) and France (Bouchereau et al. 1996),
an increase in saturated and monounsaturated fatty
acids and a concomitant decrease in polyunsaturated
fatty acids was observed in B. napus when exposed to
high temperatures during seed production. However, a
10-d moisture stress period applied at various stages of
B. napus growth in a greenhouse study did not affect
seed fatty acid saturation (Champolivier and Merrien
1996). Weselake (2000) recently reviewed lipid biosynth-
esis in domesticated B. napus seed. The effects of
environment and soil fertility on domesticated B. napus
and B. rapa fatty acid composition are reviewed in
Mendham and Salisbury (1995).
Domesticated B. napus and B. rapa seeds contain
about 22 to 26% crude protein (Bell 1995). In both
species, seed protein was primarily located in the
cotyledons (76%), with 17% in the remainder of
the embryo and the remaining 7% in the seed coat (King
et al. 1977). In B. napus, seed protein accumulates
mainly during the cell expansion phase of seed forma-
tion (Finlayson and Christ 1971), while the maximum
period of fatty acid deposition occurs during seed
maturation (Fowler and Downey 1970).
In India, Uprety et al. (1995) examined photosynth-
esis in 43 domesticated Brassicaceae and allied genera.
In domesticated B. napus, net photosynthesis was
14.0 mmol m 2 s 1 (SE 0.51) with a stomatal resistance
of 1.9 s cm 1 (SE 0.36). In domesticated B. rapa, net
photosynthesis was 13.0 mmol m 2 s1 (SE 0.36) with a
stomatal resistance of 2.2 s cm 1 (SE 0.07). Similar
values of stomatal conductance were reported in six
B. napus and four B. rapa genotypes by Kumar and
Singh (1998) with lower stomatal conductance on
average in B. rapa. Uprety et al. (1995) found greater
chlorophyll content in B. napus (1.5 mg g1 FW, SE
0.04) than in B. rapa (1.1 mg g1 FW, SE 0.02). This
difference was due to greater chlorophyll a content in
B. napus, as chlorophyll b levels were similar in both
species (0.23 and 0.25 mg g1 FW). Transpiration rates
were similar between B. napus and B. rapa (10.4 mg
cm 2 s1, SE 0.35 and 12.2 mg cm2 s 1, SE 0.27,
respectively).
Mid-season tissue osmotic concentrations in well-
watered domesticated plants of B. napus and B. rapa
tend to be similar, and ranged from 0.8 to 1.1 MPa
(Clarke and McGaig 1982; Good and Maclagan 1993).
Brassica napus possesses only a limited ability for
osmotic adjustment under extended moisture stress
( 0.3 to 0.4 MPa) (Good and Maclagan 1993;
Jensen et al. 1996). Brassica rapa, in contrast, showed
increased cell solute concentration in response to
moisture stress (Clark and McCaig 1982) reaching
osmotic potentials as low as 3.0 MPa (Good and
Maclagan 1993).
No information is available on rooting depths and
moisture extraction of volunteer canola and weedy
populations in Canada. In domesticated B. rapa grown
in the United States, Merrill et al. (2002) reported a
maximum rooting depth of 1.17 m with greater total
root length in dryer seasons. In general, rooting depth
and total root dry matter production were greater in
B. napus than in B. rapa, and Richards and Thurling
(1979) suggested that this may be due to differences in
time to maturity between these species. In India,
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
maximum soil moisture extraction was 1 to 3 cm H2O
per 30 cm depth between 30 and 150 cm soil depth, while
soil moisture extraction between 150 and 180 cm
declined sharply in two B. napus and two B. rapa
genotypes (Kumar and Singh 1998). In contrast, Kumar
et al. (1987) reported lower total moisture extraction in
five domesticated B. napus genotypes (175 to 184 mm
H2O) than in three domesticated B. rapa genotypes (186
to 215 mm H2O).
Brassica napus and B. rapa are species with prominent
tap roots and predictably the majority of root dry
matter (80 to 85%) (Kjellstrom 1991; Kjellstrom and
Kirchmann 1994) and root number distribution (41 to
78%) (Almond et al. 1984) in domesticated winter
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B. napus in Sweden were in the top 20 cm of the soil
profile. Maximum root growth was observed during the
stem elongation phase, while maximum root dry matter
(200 g m 2), length (mean2 km m 2, but as large as
4.9 km m 2) and surface area (5 m2 m 2) were found at
the time of flowering (Kjellstrom 1991; Kjelstrom and
Kirchmann 1994). The root:shoot ratios in winter
B. napus tended to be lower (0.16 to 0.64) in a cool,
wet season than during a warmer, drier year (up to 0.72)
(Kjellstrom and Kirchmann 1994). In Australia, Ri-
chards and Thurling (1979) found a lower root:shoot
ratio in drought-stressed B. rapa (0.199) than in
For personal use only.
drought-stressed B. napus (0.216). Under drought stress,
B. napus may produce short, tuberized roots [Potfer et
al. (1988) cited by Mendham and Salisbury (1995)].
Domesticated B. napus and B. rapa are sensitive to
low sulphur (S) availability in soil. When grown as a
crop in the Parkland ecoregion of western Canada,
supplemental S often increases yield (Malhi and Gill
2002). In a comparative greenhouse study in Australia,
Mailer (1989) found that supplemental S, up to 100 mg
g1, substantially increased seed glucosinolate concen-
trations in B. napus (from 1 to 32 mmoles g1) and
B. rapa (from 7 to 79 mmoles g1). Moreover, seed yield,
1000-seed-weight and seed oil content increased 15, 1.7,
and 1.5 times, respectively, in B. napus, whereas in
B. rapa, seed yield doubled, while 1000-seed-weight and
seed oil content remained unaffected. Higher levels of
supplemental S were toxic to both species.
Phosphorus (P) limiting conditions in soils have been
shown to induce the release of organic acids into the
rhizosphere in domesticated B. napus (Hedley et al.
1982). The resulting reduction of the soil pH in the
rhizosphere solubilizes P. The efficiency of this mechan-
ism for enhancing plant P uptake depends largely on the
availability of acid desorbable P in the rhizosphere
(Hedley et al. 1994). Shi et al. (2004) observed greater P
depletion (up to 13%) between a distance of 3 and
15 mm from the root surface in domesticated B. napus
than in domesticated B. rapa. The opposite effect was
observed for depletion of potassium (K) between 3 and
20 mm from the root surface. No differences in tissue P
and K concentrations and root biomass were observed,
but shoot biomass was about 67% greater in B. rapa.
965
(d) Phenology * In western Canada, seedling recruit-
ment of volunteer B. napus and B. rapa may occur in the
autumn after crop harvest (R. H. Gulden, personal
observations). However, volunteer canola seedlings
from spring genotypes generally do not survive the
winter and therefore fail to reproduce. Lawson et al.
(2006) described the recruitment periodicity of glypho-
sate-resistant volunteer B. napus in Manitoba fields
under three tillage systems (conventional tillage, low
disturbance direct seeding and high disturbance direct
seeding) 1 yr after a canola crop. Maximum seedling
recruitment was reached between early May and the end
of June [100 and 450 growing degree days (GDD) (base
temperature58C)] over the 2 yr of the study. Seedling
recruitment of volunteer B. napus was the same among
tillage systems. In a 3 yr study in Saskatchewan, the
seedling recruitment periodicity for B. napus and B. rapa
was similar and ranged from early May until the middle
of June (Gulden et al. 2003b). In that study, consecutive
flushes of volunteer canola seedling recruitment coin-
cided with spring rainfalls, whereas uncharacteristically
dry conditions during the remainder of the growing
seasons may have precluded seedling recruitment at
other times. In these Canadian studies, volunteer canola
seedling recruitment tended to be early-season, which
contrasts with reports from Europe where seedling
recruitment of volunteer B. napus occurred throughout
the entire growing season (Lutman 1993). It is not clear
whether this is related to the spring genotypes grown in
Canada compared with the primarily winter genotypes
grown in Europe or whether this is due to differences in
environment between these regions.
In a field study in Manitoba, growth rates of B. napus
were 1.21 g g1 d1 at the cotyledon stage (8 to 12 d
after planting) and 1.47 g g1 d1 at the first true leaf
stage (17 to 23 d after planting) and growth rates of
B. rapa were 0.78 and 0.82 g g1 d1, respectively
(Brandt and Lamb 1994). In a greenhouse component of
the same study, differences in growth rates between
these species were less pronounced and ranged from 0.43
to 0.87 g g1 d1 at the same growth stages. Linder and
Schmitt (1995) reported growth rates to the four-leaf
stage ranging between 0.10 and 0.18 g g1 d 1 in
several B. napus lines and one B. rapa line in a field
study conducted in California and Georgia. No sig-
nificant differences in growth rates were found between
B. rapa and B. rapa B. napus hybrids.
In the field in western Canada, domesticated spring
B. napus produces approximately 7 leaves, each taking
about 46 GDD to develop (Morrison and McVetty
1991) and in a growth chamber study, B. napus leaf area
expansion and leaf area duration was stable over a range
of temperatures when expressed based on GDD (Mor-
rison et al. 1989). In India, the rate of leaf appearance in
domesticated B. rapa was consistently more rapid than
in domesticated B. napus over a range of 5 sowing dates
(Oct. 19 to Dec. 09), particularly in the later emerging
leaves (leaves 8 to 14) (Nanda et al. 1995). Final leaf
 966 CANADIAN JOURNAL OF PLANT SCIENCE
number was consistently lower in B. rapa (13 to 21
leaves) compared with B. napus (19 to 27 leaves) and
decreased similarly as the crop was planted later. In B.
napus, not all initiated leaf primordia expand. Smith and
Scarisbrick (1990) reported a total of 23 to 28 initiated
leaf primordia in winter B. napus in the United King-
dom, while only 9 or 10 of these expanded.
In domesticated B. napus, flowering was reached at
about 576 GDD (about 47 d), pod fill was reached at
about 860 GDD (about 66 d) and maturity was reached
at about 1157 GDD (about 87 d) in western Canada
(Morrison et al. 1989; Kimber and McGregor 1995)
with similar thermal time requirements in eastern
Canada (Morrison and Stewart 2002). Domesticated
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B. rapa required about 60 d to reach maturity in western
Canada (Kimber and McGregor 1995). Days to flower-
ing and days to maturity were greater (1.3 and 1.1 d,
respectively) in a group of domesticated hybrid B. napus
genotypes compared with a group of domesticated
open-pollinated B. napus genotypes in western Canada
(Van Deynze et al. 1992). Despite an indeterminate
growth habit, the period of flowering of individual
plants was confined to 20 to 25 d in both B. napus and
B. rapa when grown as a crop in western Canada and
time of emergence appeared to have little influence on
the length of duration of flowering of individual plants
For personal use only.
(McGregor 1981). On any one plant, individual inflor-
escences may flower for as many as 15 d or fewer than
5 d, depending on the time of anthesis relative to other
flowers on that plant (McGregor 1981). Although not
obligatory, many domesticated spring B. napus geno-
types continue to respond to vernalization treatment
which reduces the time for flowering (Salisbury and
Green 1991; Robertson et al. 2002).
Periodicity of flowering is important for inter- and
intra-specific outcrossing to occur. In Canada, little is
known about the flowering periodicity of volunteer
canola. One experiment examining the overlap of
flowering periodicity between volunteer spring B. napus
and R. raphanistrum in wheat in eastern Canada showed
substantial overlap of flowering in these species (up to
88% of the total flowering time) (Simard and Légère
2004). In Europe, contradicting results have been
reported. In one study, Gruber et al. (2004a) reported
some overlap of flowering periodicity between cropped
and volunteer winter B. napus, while in a different study
(Gruber et al. 2005), volunteers flowered one month
later than a winter canola crop sown at the typical time.
The flowering phenology of weedy B. rapa popula-
tions along a cline in California was evaluated over 3 yr
(Franke et al. 2006). The mean flowering date varied by
up to 3 wk among successive years, which included the
influence of an El Niño event. Moreover, populations at
a drier site consistently flowered earlier than those at
wetter locations and greenhouse evaluations confirmed
a strong genetic component to flowering time within
these populations.
(e) Mycorrhiza * There are no known associations
between mycorrhiza and B. rapa and B. napus (Medve
1983).
8. Reproduction
(a) Floral Biology * Brassica napus is a predominantly
(on average 70%) self-pollinating species, while B. rapa,
with the exception of the Indian subsp. trilocularis
(Roxb.) Hanelt, yellow sarson (Specht and Diederichsen
2001), is considered an obligate outcrossing species. Self-
incompatibility in B. rapa is due to a sporophytic
incompatibility system and involves three genes. Two
genes at the S locus, the S locus glycoprotein (SLG) and
the S receptor kinase (SRK), are expressed in the stigma
(Nasrallah et al. 1994) and the third, a cysteine-rich
polypeptide named SCR (Schopfer et al. 1999) and SP11
(Takayama et al. 2000), is expressed in pollen. In the
stigma, the gene product of SRK alone can confer self-
incompatibility, whereas the gene product of SLG alone
was unable to confer complete self-incompatibility,
suggesting a complimentary rather than a unique role
for SLG (Takasaki et al. 2000). Self-incompatibility
occurs when the product of these genes is identical in the
stigma and pollen, causing failure in pollen tube
initiation (Zuberi and Lewis 1988; Takasaki et al.
2000). Several comprehensive reviews on self-incompat-
ibility in the Brassicaceae have been published (Cock
et al. 2000; Watanabe et al. 2000; Kemp and Doughty
2003; Takayama and Isogai 2003). Recently, self-in-
compatibility from B. oleracea was introgressed into the
domesticated B. napus genome in Canada to facilitate
the generation of hybrid cultivars (Ripley and Bevers-
dorf 2003a). In B. napus, differences in S-allele dom-
inance (Ripley and Beversdorf 2003b) and differences in
S-allele expression (Ripley and Beversdorf 2003c) com-
pared with B. oleracea were observed. The significance
of these breeding systems for volunteer B. napus
populations remains unknown.
In weedy B. rapa (bird rape), a comparison by
Warwick and Black (1993) of observed levels of allelic
heterozygosity with expected levels under random mat-
ing provided evidence for significant departures from
random mating, as lower mean values than expected
were observed for both triazine-resistant populations
(0.133 vs. 0.153 mean observed versus mean expected
heterzygosity per locus) and susceptible populations
(0.159 and 0. 184 mean observed versus mean expected
heterzygosity per locus) in Quebec. In China, variation
in outcrossing rates (54 to 100%) has been reported
among natural populations of B. rapa (Liu 1985, cited in
Bing et al. 1991).
Despite being a primarily self-pollinated species, out-
crossing rates in B. napus can be as high as 47% (Becker
et al. 1992; reviewed in Beckie et al. 2003). In western
Canada, average outcrossing rates to adjacent plants
greater than 20% have been reported in domesticated
B. napus (Rakow and Woods 1987; Cuthbert and
McVetty 2001) with rates of outcrossing decreasing as
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
the distance between plants increases (Staniland et al.
2000). In B. napus, the rate of outcrossing is influenced
by the position of the flowers on the plant (Rakow and
Woods 1987). Flowers located near the bottom of the
plant are up to 3.5 times more likely to be cross-
pollinated than flowers located near the top of the plant
(Becker et al. 1992). In domesticated winter B. napus in
the United Kingdom, the rate of outcrossing does not
appear to influence total seed production (Williams
et al. 1986).
Within B. napus and B. rapa, outcrossing can occur by
wind and insect pollinators. In a study in western
Canada, Beckie et al. (2003) used herbicide-resistance
as a marker and reported mean outcrossing rates of
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1.4% between adjacent B. napus fields at the common
margin and 0.04% at a distance of 400 m in western
Canada. Some pollen movement was detected at the
study margin (800 m). Studies in Europe have measured
long-distance movement of domesticated B. napus
pollen by wind. At a distance of 360 m from the field,
airborne pollen density declined to 10% of that found at
the field margin (Timmons et al. 1995). Nevertheless, up
to 22 airborne pollen grains m 3 were measured 1.5 km
from the source field, with pollination of emasculated
bait plants reported at this distance. In male-sterile
B. napus, 23 to 29% of seed production within 6 m from
For personal use only.
the pollen source was attributed to wind pollination,
while at a distance greater than 24 m, seed production
from wind pollination was negligible relative to insect-
assisted pollination (Mesquida and Renard 1982). Out-
crossing percentage was influenced by wind direction in
this study. However, even with insect-assisted pollina-
tion, long distance outcrossing appears to be a relatively
rare event in domesticated B. napus in the United
Kingdom (0.016% at 200 m, 0.004% at 400 m)
(Scheffler et al. 1995). In a similar study conducted at
a smaller scale, Scheffler et al. (1993) reported even
lower rates of outcrossing, indicating that the rate of
outcrossing also is influenced by the magnitude of the
pollen source.
In the United Kingdom, honeybees (Apis mellifera L.)
and bumblebees (Bombus terrestris L.) are among the
most prominent outcrossing vectors of domesticated
B. napus (Scheffler et al. 1993). The duration of B. napus
flower visitation by bumblebees was directly correlated
to the amount of nectar present, but was independent of
the quantity of pollen present (Cresswell 1999). In
Germany, the red mason bee (Osmia rufa L.), a solitary
species, was also identified as an efficient pollination
vector for B. napus (Steffan-Dewenter 2003). Mason bee
species also occur in Canada, although their role in
pollinating Brassicaceae has not been evaluated. In a
study in the United States, exposure to increased UV-B
radiation did not affect flower visitation behaviour of
A. mellifera in rapid cycling B. rapa and B. nigra (Collins
et al. 1997). In this study, UV-B radiation treatments
were applied in growth chambers daily and plants were
taken outside near a bee colony, where bee foraging
967
behaviour was observed. In France, Pierre et al. (1996)
compared honeybee visitation between B. napus lines
with and without petals. Pollinator visitation was high
in both flower types, but individual honeybees prefer-
entially visited only one flower type and therefore cross-
pollination between petalous and apetalous B. napus
was limited. Visitation by honeybees, bumblebees,
solitary bees and Diptera was the same between domes-
ticated conventional and transgenic glufosinate-resistant
winter B. napus (Pierre et al. 2003).
In Manitoba, total sugar content of nectar of
domesticated B. napus ranged from 600 to 800 mg per
flower in 21 different genotypes (Pernal and Currie
1998). Total sugar, composed of about equal quantities
of fructose and glucose, was highest during early
flowering and displayed a diurnal pattern with max-
imum total sugar content per flower observed in the
afternoon. No differences in nectar production were
observed among the three different breeding systems
(open-pollinated, cytoplasmic male-sterile, and domi-
nant self-incompatible) present in this selection of
genotypes. In a comparative study conducted in Poland,
domesticated spring B. napus genotypes produced up to
50% more pollen and up to 50% less nectar than
domesticated winter B. napus genotypes (Kotowski
2001). Seed fatty acid profile did not affect pollen or
nectar production. In contrast, Pierre et al. (1999) found
that among 71 domesticated winter B. napus genotypes,
nectar production tended to be greatest in low erucic
acid and low glucosinolate lines compared with low
erucic acid and double high lines. The presence of the
glufosinate-resistant transgene did not affect nectar and
pollen production in domesticated winter B. napus
grown in France (Pierre et al. 2003).
Apomixis or agamospermy is a common phenomenon
in the Brassicaceae, especially when a generally self-
incompatible species, such as B. rapa, is the pollen
recipient of pollen from another Brassica species
(reviewed in Heyn 1977). Such matroclinous or matro-
morphic plants (resulting through the formation of
diploid gametes), have been reported repeatedly in
inter-specific crossings in Brassica (e.g., Kakizaki 1925;
Olsson 1954, 1960; MacKay 1968, 1972; Heyn 1977).
Fukushima (1931), Chèvre et al. (2000) and Warwick
et al. (2003) have shown the presence of unredu-
ced gametes. Pseudogamy has also been reported
in Brassica species (Noguchi 1928; Mohammud and
Sikka 1940). Production of unreduced gametes is also
reported for several inter-generic crosses in the Brassi-
caceae (Salisbury 1989), Sinapis arvensis B. napus
(Moyes et al. 2002) and R. raphanistrum B. napus
(Warwick et al. 2003). Lelivelt (1993) studied the
fertilization process and seed set in intraspecific
and intergeneric crosses involving B. napus, B. napus 
Brassicoraphanus (raparadish), and S. alba. The occur-
rence of matromorphic individuals in crosses where the
female plant was B. napus suggested that relatively high
rates of matromorphy/parthenogenesis occur in this
 968 CANADIAN JOURNAL OF PLANT SCIENCE
species. This, however, was not confirmed experimen-
tally. A low frequency (B1%) of matromorphy was also
suspected in weedy B. rapa in New Zealand when
fertilized with domesticated B. napus pollen (Jenkins et
al. 2005).
Domesticated B. napus and B. rapa are particularly
sensitive to heat stress during early flowering. In a
greenhouse study in western Canada, domesticated
B. rapa was more sensitive to heat stress treatment (35/
15oC) than domesticated B. napus (Angadi et al. 2000), yet
the optimum temperature for successful flowering was
lower in B. napus than in B. rapa. Both species recovered
quickly from the 7-d heat stress that was imposed early
during flowering. Plants compensated for aborted main
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stem flowers by initiating more lateral flowers once the
heat stress was removed. High temperature stress in B.
napus caused the expression of heat shock proteins in
both, the micro- and the megagametophyte (Young et al.
2004); however, the detrimental effects of heat stress were
greater on the microgametophyte. In domesticated B.
napus in western Canada, heat stress can result in seed pod
deformation during development which may lead to
deformed seed (Morrison 1993; Angadi et al. 2000), which
has not been observed in B. rapa (Angadi et al. 2000).
In the United States, a series of studies examining the
effects of increased ultraviolet-B (UV-B) radiation on
For personal use only.
rapid cycling B. rapa genotypes were conducted. In the
greenhouse study, growth and flowering period were
decreased, while fitness was increased in B. rapa in
treatments where supplemented UV-B radiation levels
were two- to threefold greater (Feldheim and Conner
1996). In an outdoor experiment, B. rapa plants were
artificially supplemented with UV-B from light racks
placed in the field. Visitation by pollinators doubled in
the supplemented UV-B treatments; however, fitness
was not affected by this treatment (Conner and Zangori
1997). Variable ambient light levels precluded an
accurate estimation of the supplemental UV-B radiation
in this experiment. In a greenhouse study, water and
nutrient-stressed B. rapa plants were exposed to ambient
and three-times ambient UV-B levels. Fitness traits
showed no significant interaction among the imposed
moisture and nutrient stresses and supplemental UV-B
radiation, suggesting a high degree of independence
between these stresses in B. rapa (Conner and Zangori
1998).
Bell and Cresswell (1998) described the function of an
inflorescence as primarily female for the first hour after
opening, followed by a primarily male role for the
remainder of anthesis in domesticated B. napus in the
United Kingdom. Pollen removal dictated flower senes-
cence, with high levels of pollen removal, reducing by
half the period of anthesis of individual flowers.
(b) Seed Production and Dispersal * In eastern Canada,
domesticated B. napus that overwintered as a rootstock
after harvest produced more than 3000 seeds plant 1 as
a volunteer during the following season (Simard et al.
2002). No information on seed production in volunteer
B. napus in western Canada was found in the literature.
However, in a recent 2-yr study that included one dry
growing season in Saskatchewan, seed production of
B. napus grown in monoculture at different population
densities ranged from 16 to 25.8 seeds pod1 (Angadi
et al. 2003). The number of seeds pod 1 was not
influenced by plant density (5 to 80 plants m2);
however, pod number was strongly influenced by plant
density and ranged from 44 to 582 pods plant 1,
indicating high phenotypic plasticity in this species.
Thousand-seed weight ranged from 2.70 to 3.58 g with
no effect of plant density on this parameter. Similar seed
yield component values were reported by Morrison et al.
(1990) in a field study on domesticated B. napus in
monoculture in Manitoba. In Saskatchewan, Kirkland
and Johnson (2000) found a negative relationship
between time of emergence and seed size of progeny in
domesticated B. napus where the size of individual seeds
decreased substantially (up to 25% smaller) when the
crop was planted in mid-May compared with late-April
or the previous fall.
There also is a paucity of information on seed
production of volunteer or weedy B. rapa in Canada.
In the United States, fruit production ranged from 19 to
187 pods plant 1 in a naturalized B. rapa population in
California (Agren and Schemke 1994); seed production
was not reported. As a crop in Turkey, B. rapa had 217
and 404 pods plant1 with 10 to 11 seeds pod1,
resulting in stable seed production over planting den-
sities ranging from 50 to 800 seeds m 2 (Bilgili et al.
2003). Seed production in domesticated B. rapa tends to
be lower than in domesticated B. napus (Miller et al.
2003).
In Australia, Richards (1978) found that post-anthesis
growth was an important factor in determining
final seed production in domesticated B. rapa. In
contrast, pre-flowering biomass accumulation was
more important in determining final seed production
in B. napus (Richards and Thurling 1978). Brassica
napus and B. rapa are susceptible to a number of
diseases and pests which may reduce final seed produc-
tion (Section 13).
Seed dispersal*Brassica napus and B. rapa produce
seeds in pods which shatter at maturity. This enables
seed dispersal over a limited distance. Neither species
possesses other specialized dispersal mechanisms. In
Australia, where sheep (Ovis aries L.) are often allowed
to graze stubble of canola crops, viable B. napus seed
was recovered from sheep excrement as many as 5 d
after ingestion (Stanton et al. 2003). This could promote
seed dispersal.
Human-assisted dispersal at the time of harvest can
play an important role in seed dispersal. Success of
breeding efforts to increase resistance to seed shattering
has been limited, particularly in domesticated B. napus,
although incorporation of genes from other Brassica
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
crop species has shown some promise (Morgan et al.
2000; Wang et al. 2007) (Section 7). Increasing shatter-
ing resistance in these species which may increase
difficulty of seed separation during mechanical harvest
could increase volunteerism (Bruce et al. 2001). Long-
distance dispersal of volunteer B. napus to non-agricul-
tural habitats such as roadsides and railroad lines is
thought to have occurred through commercial seed
movement with trucks in Canada (Yoshimura et al.
2006) and the United Kingdom (Crawley and Brown
1995) and trains in Canada (Yoshimura et al. 2006).
Seed Bank Inputs * Addition of canola seed to the soil
seed bank at harvest varies according to many factors
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and can be quite large. A survey of 35 fields over 2 yr in
western Canada revealed large variability in the number
of B. napus seeds added to the soil seed bank among
fields from different producers at the time of harvest
(Gulden et al. 2003a). On average, 3000 viable seeds
m 2 (5.8% of the yield) were added to the
seed bank; however, seed bank additions ranged from
1500 to 7100 seed m 2 with one exception where almost
14 000 seeds m 2 were added to the seed bank. In this
study, most fields were windrowed (i.e., rows of crop left
to dry in the field) prior to harvest and the majority of
the lost seeds appeared to be located where the windrow
For personal use only.
had lain (visual observations). One year after a canola
crop, greater volunteer B. napus populations (up to
sixfold) were found where the windrow from the crop
had lain before harvest in untilled fields in Saskatch-
ewan (Beckie et al. 2003). In a study conducted in the
1970s in Saskatchewan, canola (type not provided)
harvest losses of 9 kg ha 1 and 14.5 kg ha1 were
reported when the crop was windrowed at 37 and 12%
seed moisture content, respectively (Bowren and Pitt-
man 1975). Nevertheless, the relatively small seed size of
B. napus (1000 kernel weightB4 g based on dry matter)
contributes to large seed bank numbers.
In the United States, spring B. napus harvest seed
losses of 350 kg ha 1 or approximately 15% of the
attainable yield have been reported in the Pacific
Northwest (Brown et al. 1995); however, delayed wind-
rowing increased harvest losses to as high as 28.5%. In
the southeastern United States, average harvest seed
losses of 34.5% were reported in winter B. napus
(Thomas et al. 1991). In the United Kingdom, mean
harvest seed losses of winter B. napus during direct
harvest range from 2 to 5% under ideal conditions, but
may be as high as 50% under unfavourable harvest
conditions (Price et al. 1996). In another British study,
harvest losses between 200 and 300 kg ha1 or 5000 to
7000 seeds m 2 were reported in winter B. napus
(Pekrun et al. 1998b).
(c) Seed Banks, Seed Viability and Germination * In
arable soil seed banks, seed persistence has been
observed for at least 3 yr in both B. napus and B. rapa
(Gulden et al. 2003b) and increased autumn soil
969
moisture was associated with decreased seed bank
persistence in B. napus in western Canada (Gulden
et al. 2004b). Testimonials (no data available) suggest
that longer periods of seed bank persistence may be
possible in western Canada, particularly in B. rapa. In
Europe, winter B. napus seed can persist for 10 or more
years in the seed bank (Lutman et al. 2003; Jørgensen
et al. 2007).
Seed burial increases seed bank persistence in B. napus
and B. rapa. In western Canada, all B. napus seeds
buried at a depth of 1 cm lost viability by June in the
year following entry into the seed bank (Gulden et al.
2004b). At the same time, 20 to 30% of the seed bank of
the most persistent B. napus genotype remained viable in
seed buried at a depth of 10 cm during the previous
autumn. Burial of domesticated B. rapa (cv. Tobin) up
to depths of 12 cm increased the proportion of persisting
seed over one winter in Alaska where seed survival was
enhanced significantly when covered with snow (62%
seed survival) compared with without snow cover (39%
seed survival) (Sparrow et al. 1990). The highest levels of
seed bank mortality occurred between March and April.
Most studies on volunteer winter B. napus seed bank
persistence in the United Kingdom (Pekrun and Lutman
1998) and Germany (Schlink 1995) also have reported
increased seed bank persistence with increased depth of
seed burial, although the opposite was reported in one
instance in France (Chadoeuf et al. 1998). Lack of
disturbance following seed burial tends to increase
persistence of the seed bank in B. napus (Schlink 1995;
Lutman et al. 2003; Gulden et al. 2003b; Gulden et al.
2004b). Seed bank persistence of volunteer B. napus and
B. rapa tends to be greater in fine-textured soils in
western Canada (Gulden et al. 2003b) and the United
Kingdom (Lutman et al. 2003). Herbicide-resistance
traits did not affect seed bank persistence of volunteer
B. napus in farm fields in western Canada (Gulden et al.
2003b) and Germany (Gruber et al. 2004b), and natural
habitats in the United Kingdom (Crawley et al. 1993;
Hails et al. 1997).
The maximum depth of seedling recruitment of
volunteer B. napus was less than 10 cm in western
Canada (Gulden et al. 2004b) and the United Kingdom
(Lutman 1993), and was different among the three
genotypes examined in the latter study. In the United
States, Linder and Schmitt (1995) reported seedling
recruitment between 7 and 60% from a depth of 4 cm
among one B. rapa and several B. napus lines. In a long-
term seed burial experiment in Virginia, germination
percentages of seed of B. rapa following exhumation
after burial at depths of 20, 56, and 107 cm were 0, 0,
1% after 1 yr; 0, 0, 2% after 3 yr; 0, 1, 1% after 6 yr; 0,
0, 3% after 10 yr; and none after 20, 30 and 40 yr (Toole
and Brown 1946). In the United Kingdom, Walker et al.
(2004) conducted a seed burial experiment to determine
the seed bank persistence of winter B. napus lines with
different seed oil quality characteristics. Seed bank
persistence tended to be lower in the high stearate and
 970 CANADIAN JOURNAL OF PLANT SCIENCE
high laurate transgenic and parental lines compared
with conventional oil quality genotypes after 18 mo of
seed burial.
In fields, the seed bank and population density of
volunteer B. napus and B. rapa decreases rapidly with
increasing time since the last canola crop in Canada
(Simard et al. 2002; Gulden et al. 2003b; Gulden et al.
2004b) and Europe (Pekrun and Lutman 1998; Lutman
et al. 2003). In Saskatchewan, the seed bank population
was 23 to 40%, B2%, and negligible in the first, second,
and third spring after seed bank establishment, in a
study where seed bank replenishment was not allowed
(Gulden et al. 2003b). In the United Kingdom, the
volunteer winter B. napus seed bank declined to 5%
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within 15 and 39 months after seed bank establishment
(Lutman et al. 2003). However, long-term studies also
indicated that after 3 to 4 yr, seed mortality in
undisturbed, buried seeds becomes negligible and a
small fraction of volunteer winter B. napus seed readily
persisted for up to one decade (Lutman et al. 2003).
When compared with other common weed species in the
United Kingdom, the initial seed bank decline rates of
volunteer winter B. napus were among the most rapid
(Lutman et al. 2002; Lutman et al. 2005).
Dormancy * Neither domesticated B. napus nor domes-
For personal use only.
ticated B. rapa exhibit primary seed dormancy at
maturity (Pekrun et al. 1998b; Gulden et al. 2004a). In
contrast, weedy B. rapa does exhibit some primary seed
dormancy (Hauser et al. 1998a, b). Domesticated
B. napus seed can develop secondary seed dormancy
when exposed to darkness and conditions inhibitory to
germination, such as moisture stress (Pekrun et al.
1997a), hypoxia (i.e., low oxygen concentrations) (Pek-
run et al. 1997a; Momoh et al. 2002), or high tempera-
tures (Zhang and Hampton 1999; Gulden et al. 2004a).
In the laboratory, moisture stress (1.5 to 2 MPa) in
combination with darkness effectively induce secondary
seed dormancy in B. napus over a period of 4 wk, while
hypoxic conditions (3% O2) for the same duration were
less effective (Pekrun et al. 1997a; Momoh et al. 2002).
Alternating temperatures reduced secondary seed dor-
mancy induction in B. napus (Pekrun et al. 1997b).
Secondary seed dormancy in B. napus was released by
gibberellic acid (0.2 mg L 1) (Pekrun et al. 1998a), a
0.002 s flash of white light (Schlink 1994 cited in Pekrun
et al. 1998a), and a short stratification treatment (3 to
5 d at 2 to 48C) (Pekrun et al. 1998a; Gulden et al.
2004a). Germination of dormant domesticated winter B.
napus seed was double that of the water control (48%)
when solutions of gibberellic acid (0.44 kg ai ha 1 GA3)
or 1-(3-chlorophthalimido) cyclohexanecarboxamide
(AC94377) (2 kg ai ha1) were applied to the soil
surface in a greenhouse study in the United Kingdom
(Thornton et al. 1998). Research in the 1950s on B.
napus (type unknown) in Japan found that seed dor-
mancy (presumably primary) was released by low
temperatures in conjunction with washing and pricking
the seeds (Sugiyama 1949, cited by Takahashi and
Suzuki 1980) or by treatment with either thiourea or
urea (Hori and Sugiyama 1954, cited by Takahashi and
Suzuki 1980).
In Germany, a seasonal response of germination to
light exposure was observed in domesticated winter
B. napus where, when exposed to light, the germinable
proportion of the viable seed bank was lower from seed
exhumed during summer compared with any other time
of the year (Schlink 1995). Such a seasonal response in
germinability was not observed in seeds exposed to
darkness after exhumation. In a laboratory study in
Poland, germination of domesticated winter B. napus
was inhibited by the continuous exposure to white light
in combination with moisture stress (Bazanska and
Lewak 1986). Further experimentation led the authors
to suggest the presence of a water-soluble germination
inhibitor.
Secondary seed dormancy potential varies among
domesticated B. napus genotypes (Pekrun et al. 1997c;
Momoh et al. 2002; Gulden et al. 2004a). Among a
group of 16 Canadian spring genotypes, secondary seed
dormancy potential ranged from B20 to90% (Gulden
et al. 2004a) with a high potential in most genotypes
(80%). Among several factors investigated in this
study, genotype contributed most prominently to sec-
ondary seed dormancy potential (69% of total varia-
tion) and seed size was the second most important factor
(21%), with large seeds showing a greater ability to
develop dormancy. Despite a large variation in environ-
mental conditions during the 4 yr of this study, the
contribution of year and location to the total variation
in secondary seed dormancy potential was negligible
(B5%). In Canadian spring B. napus (Gulden et al.
2003b, b) and European winter B. napus genotypes
(Gruber et al. 2004b), secondary seed dormancy poten-
tial has been linked to seed bank persistence in the field.
Conventional and transgenic herbicide-resistant
(glyphosate and glufosinate) genotypes behaved simi-
larly in both studies.
Genotype also influenced seed bank persistence in
domesticated B. rapa where, after 3 yr, seed bank
persistence of one B. rapa genotype was approximately
one order of magnitude greater than that of a second
B. rapa genotype in western Canada (Gulden et al.
2003b). However, seed dormancy induction in domes-
ticated B. rapa could not be achieved using the
combination of moisture stress, darkness, and tempera-
ture that efficiently induced secondary seed dormancy in
B. napus (Gulden et al. 2003b).
Germination and dormancy characteristics of hybrids
between domesticated B. napus and weedy B. rapa were
investigated to assess the effects of possible transgene
escape through hybridization between these species
(Landbo and Jørgensen 1997). Levels of primary
dormancy were high for the parental weedy B. rapa
populations, but low for B. napus and the reciprocal
B. napus weedy B. rapa hybrids.
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
Germination * Little information on germination of
volunteer and weedy populations of B. napus and
B. rapa is available; however, the requirements for
germination of domesticated genotypes of these species
are well documented. In Canadian spring B. napus
genotypes, base temperatures (i.e., the lowest tempera-
tures at which germination occurs) ranged between 0.44
and 28C (Kondra et al. 1983; Vigil et al. 1997), while in
European spring and winter B. napus genotypes, base
temperatures were 38C or lower (Marshall and Squire
1996; Squire 1999). Breeding for lower base tempera-
tures in B. napus has been attempted with limited success
(Acharya et al. 1983; King et al. 1986).
In laboratory and greenhouse experiments, final
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germination percentage and seedling vigour were re-
duced by up to 98% and about 80%, respectively, in
domesticated Canadian B. napus and B. rapa genotypes
at low temperatures (2
58C compared with 158C)
(Acharya et al. 1983). Similar reductions in vigour also
have been observed in the United States and Denmark
(Elias and Copeland 1997; Larsen et al. 1998). In a
different laboratory study, the mean increase in time to
germination at low temperatures (28C) was greater
(1.7-fold) in 12 domesticated Canadian B. rapa lines
than in 11 domesticated Canadian B. napus lines
compared with time to germination at ambient tem-
For personal use only.
peratures (King et al. 1986). In the field, germination
and emergence of domesticated winter B. napus may
take up to 18 d at 58C in western Canada (Blackshaw
1991). In a greenhouse study conducted in the United
States, emergence of winter B. napus genotypes was
slower than that of spring genotypes (Vigil et al. 1997).
At low temperatures, differences in field emergence
percentage were observed among genotypes of forage
B. napus and B. rapa in Nevada, United States (Wilson
et al. 1992) and Scotland (Marshall and Squire 1996).
Freezing temperatures may not kill seeds that have
completed germination (i.e., the protrusion of the
radicle from the seed). Domesticated European B. napus
with radicles protruding by up to 1.5 mm survived
storage at 208C for at least 336 d with only a minor
loss of viability (4%) when air dried to a water content
of 11% prior to exposure to freezing temperatures
(Finch-Savage and McKee 1989). Storage at 1 and
158C increased the loss of viability substantially. Never-
theless, temperatures as low as 208C are within the
range that may be experienced in fields throughout the
winter in western Canada.
In Petri dishes, Schopfer and Plachy (1984) demon-
strated that below -0.6 MPa, the germination rate and
total germination of European winter B. napus dimin-
ished rapidly. Similarly, germination of domesticated
forage B. napus and B. rapa decreased at soil water
potentials of 0.25 and 0.50 MPa at 7.5 oC (Rao and
Dao 1987). Livingston and de Jong (1990) observed an
interaction between matric (the reduction of free energy
971
of water due to adsorption to hydrophilic surfaces) and
osmotic potentials (the reduction of free energy due to
solutes in solution water) during germination of domes-
ticated B. napus. At osmotic potentials of 1.2 MPa,
the germination rate of B. napus was lower in soils with
higher matric potential (0.02 MPa) than soils with
lower matric potential (0.2 MPa) indicating an
interaction between soil salinity and total soil water
potential. High levels of sodium (Na) decreased the rate
and percentage of germination in domesticated Cana-
dian B. rapa (Zheng et al. 1998) and domesticated
Canadian B. napus (Huang and Redmann 1995a, b) in
the laboratory and caused deformation in seedlings
(Zheng et al. 1998). The presence of sulphate appeared
to alleviate these symptoms to some extent (Zheng et al.
1998), while supplemental calcium (Ca) alleviated some
of the symptoms caused by high NaCl (Huang and
Redmann 1995a, b).
(d) Vegetative Reproduction * There is no evidence of
vegetative reproduction in these species.
9. Hybrids
Extensive hybridization studies have been conducted on
B. napus and B. rapa. Interspecific studies which
successfully used conventional sexual crosses to produce
F1 hybrids are summarized in Table 5 and recently
summarized and evaluated with regards to potential for
transgene escape in an extensive review by Fitzjohn et al.
(2007). Many other hybridizations with more distant
relatives have been accomplished using embryo, ovary
or ovule rescue techniques (reviewed in Warwick et al.
2000). Various studies (Olsson 1954; McGrath and
Quiros 1991) have examined fertility levels between the
various subspecies and members of the B. rapa AA
genome.
Gene Flow to Other Brassica Crops
There are two other Brassica crops grown in Canada,
both of which are sexually compatible with B. napus.
Ongoing pollen flow studies from transgenic glyphosate-
resistant B. napus to B. juncea (oriental mustard) and
B. rapa (Polish canola) have documented gene flow to
both crops at distances up to 200 m (Séguin-Swartz,
Beckie, and Warwick, unpublished data in Warwick
2006) (Section 8a). The consequence of gene flow to
domesticated B. rapa, which occupies approximately 5%
of the total Canadian canola acreage, is considered to be
negligible. However, at present there are no established
thresholds for transgene presence in the oriental mus-
tard crop.
Gene Flow to Related Wild/weedy Species in Canada
In Canada, there are four wild relatives that have the
potential to cross with B. napus, including three out-
crossing (i.e. self-incompatible) species: Sinapis arvensis
 972 CANADIAN JOURNAL OF PLANT SCIENCE

Table 5. Interspecific hybridization studies with Brassica napus (Bn) and Brassica rapa (Br). All were conventional sexual crosses that produced F1 first
generation hybrids. F1 and BC backcross hybrid generation production are indicated in square brackets

Female parentmale parent Citation

Brassica napus (n19)

BnBrassica carinata A. Braun (n 17)
BnBrassica juncea (L.) Czern. (n18)
[BnBrassica juncea F1]B. juncea
BnBrassica maurorum Durieu (n8)
BnBrassica nigra (L.) W.D.J. Koch (n 8)
BnBrassica oleracea L. (n 9)
BnBr
BnDiplotaxis catholica (L.) DC. (n9)
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Alam et al. (1992), Gupta (1997)
Alam et al. (1992), Bing et al. (1991, 1996b), Gupta (1997),
Choudhary and Joshi (2001a), Zhao et al. (2003), Heenan et al. (2007)
Kirti et al. (1995)
Bijral et al. (1995)
Bing et al. (1991)
Gupta (1997)
Bing et al. (1991, 1996b), Jorgensen and Andersen (1994), Vijayakumar et al. (1994), Brown and
Brown (1996), Mikkelsen et al. (1996a), Gupta (1997), Landbo and Jørgensen (1997), Lewis et al.
(2001), Lu and Kato (2001), Liu et al. (2002), Zhang et al. (2002), Leflon et al. (2006)
Bijral and Sharma (1998)

BnDiplotaxis muralis (L.) DC. (n21) Bijral and Sharma (1996a)

BnEruca vesicaria (L.) Cav. subsp. sativa Bijral and Sharma (1996b)
(Mill.) Thell. (n11)
BnErucastrum gallicum (Willd.) O.E. Schulz Lefol et al. (1997)
(n15)
BnHirschfeldia incana (L.) Lagr.-Foss. Chadoeuf et al. (1998), Lefol et al. (1995, 1996b), Chèvre et al. (1998b)
(n7)
BnOrychophragmus violaceus (L.) O.E. Cheng et al. (2002)
Schulz (n 12)
BnRaphanus raphanistrum L. (n9) Eber et al. (1994), Baranger et al. (1995), Chadoeuf et al. (1998), Chèvre et al. (1998a), Darmency
et al. (1998), Lefol et al. (1997), Rieger et al. (2001), Guéritaine et al. (2003a, b), Chèvre et al.
(2003, 2007)
[BnRaphanus raphanistrum F1] Chèvre et al. (1997, 1998a)
R. raphanistrum
BnRaphanus sativus L. (n9) Gupta (1997)
For personal use only.

BnSinapis alba L. (n 12) Bijral et al. (1993, 1994), Chèvre et al. (1994)
BnSinapis arvensis L. (n9) Moyes et al. (2002)
[BnSinapis arvensis F1]Bn Inomata (1997)
Brassica carinata A. Braun (n17)Bn Getinet et al. (1997)
Brassica junceaBn Bing et al. (1991, 1996b), Alam et al. (1992), Sharma and Singh (1992), Vijayakumar et al. (1994),
Frello et al. (1995), Rao and Shivanna (1997), Gupta (1997), Sandhu and Gupta (2000),
Choudhary and Joshi (2001a), Zhao et al. (2003), Heenan et al. (2007)
Brassica nigraBn Bing et al. (1991)
Brassica oleraceaBn Gupta (1997)
Br Bn (see below)
Brassica tournefortii Gouan (n 10)Bn Gupta (1997)
Diplotaxis erucoides (L.) DC. (n 7)Bn Ringdahl et al. (1987)
Diplotaxis muralisBn Fan et al. (1985), Ringdahl et al. (1987), Salisbury (1989), Gupta (1997)
Diplotaxis siifolia Kunze (n10)Bn Gupta (1997)
Hirschfeldia incanaBn Lefol et al. (1996b), Darmency and Fleury (2000)
Raphanus raphanistrumBn Eber et al. (1994), Baranger et al. (1995), Darmency et al. (1998), Chèvre et al. (1997, 1998a, 2000),
Guéritaine and Darmency (2001), Rieger et al. (2001), Guéritaine et al. (2002, 2003a, b),
Benabdelmouna et al. (2003), Warwick et al. (2003)
Sinapis arvensisBn Moyes et al. (2002)
Brassica rapa (n10)
BrBrassica barrelieri (L.) Janka (n10) Mattsson (1988)
BrBrassica carinata Rahman (2002), Li et al. (2005)
BrBrassica juncea Prasad et al. (1997), Rhee et al. (1997)
BrBn Bing et al. (1991), Leckie et al. (1993): Jorgensen and Andersen (1994), Bing et al. (1996b), Brown
and Brown (1996), Landbo et al. (1996), Mikkelsen et al. (1996b), Landbo and Jørgensen (1997),
Hauser et al. (1998a, b), Hansen et al. (2001), Halfhill et al. (2001, 2002, 2004), Wilkinson et al.
(2000, 2003), Warwick et al. (2003), Zhu et al. (2004), Ammitzbøll et al. (2005), Jenkins et al.
(2005), Simard et al. (2007), Yoshimura et al. (2006)
[BrBn F1]Br Mikkelsen et al. (1996b), Metz et al. (1997), Hansen et al. (2001), Halfhill et al. (2003)
BrBrassica nigra Mattsson (1988), Bing et al. (1991), Prasad et al. (1997)
BrBrassica oleracea Wojciechowski (1985), Mattsson (1988), Akbar (1989), Cheng et al. (1994)
BrErucastrum gallicum Lefol et al. (1997)
BrRaphanus sativus Ellerström (1978), Gupta (1997), Rhee et al. (1997)
Brassica barrelieriBr Takahata and Hinata (1983)
Brassica carinataBr Choudhary et al. (2000), Li et al. (2005)
Brassica fruticulosa Cirillo (n8)Br Takahata and Hinata (1983)
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L. 973
Table 5 (Continued)
Female parentmale parent Citation

Brassica junceaBr
Brassica maurorum Durieu (n8)Br
BnBr (see above)
Brassica oleraceaBr
Brassica oxyrrhina (Coss.) Willk. (n9)Br
Brassica tournefortii Br
Diplotaxis muralisBr
Diplotaxis tenuifolia (L.) DC. (n11)Br
Diplotaxis virgata (Cav.) DC. (n9)Br
Raphanus sativusBr
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22
Sharma and Singh (1992), Gupta (1997), Katiyar and Chamola (1995, 1998), Rhee et al. (1997),
Choudhary and Joshi (2001a), Choudhary et al. (2002)
Takahata and Hinata (1983)
Ellerström (1978), Wojciechowski (1985), Gupta (1997), Lu and Kato (2001)
Mattsson (1988)
Choudhary and Joshi (2001b)
Salisbury (1989)
Salisbury (1989)
Takahata and Hinata (1983)
Ellerström (1978); Rhee et al. (1997)

(wild mustard), Raphanus raphanistrum (wild radish),
and weedy B. rapa L. (bird rape), and one predomi-
nantly selfing species, Erucastrum gallicum (Willd.) O.E.
Schulz (dog mustard).
Sinapis arvensis. Gene flow from B. napus to S. arvensis
has a low probability of occurrence. Moyes et al. (2002)
is the only study to date to report fertile hybrids when S.
arvensis was the maternal parent, under greenhouse
conditions. In the field, studies have not detected gene
transfer from B. napus to S. arvensis in experiments
conducted in Saskatchewan (Bing et al. 1995; 1996b),
For personal use only.
detected in numerous field experiments in France when
R. raphanistrum served as the maternal parent (Eber
et al. 1994; Baranger et al. 1995; Darmency et al. 1998;
Chèvre et al. 2000). The hybridization rate was esti-
mated at between 10 7 and 10 5 (Chèvre et al. 2000).
In Australia, gene flow studies between R. raphanistrum
and imidazolinone-resistant B. napus growing in experi-
mental field plots (Rieger et al. 2001) indicated even
lower hybridization rates (B4108), with no hybrids
found when R. raphanistrum was the maternal parent. In
Canada, R. raphanistrum co-exists with B. napus only in
Quebec and Alberta. Canadian studies (Warwick et al.

France (Lefol et al. 1996a), and the United Kingdom
(Moyes et al. 2002). In Canada, S. arvensis is the most
common of the four weeds listed above. In recent studies
(Warwick et al. 2003), the absence of gene flow was
inferred by screening seed collected from S. arvensis
populations for the presence of the herbicide resistance
(HR) trait found in adjacent commercial HR B. napus
fields in Saskatchewan. No S. arvensis HR B. napus
hybrids were detected in 42 828 seedlings, suggesting
that the probability of inter-specific gene flow from
B. napus to S. arvensis is very low (B5 10 5) under
commercial field conditions.
2003) confirm that gene flow between R. raphanistrum
and B. napus is also rare. A single R. raphanistrum B.
napus F1 hybrid was obtained in an HR B. napus field
plot experiment in ON, where R. raphanistrum plants
were grown at a density of one plant m 2 with HR B.
napus. This hybrid had an unstable genomic structure
consistent with the fusion of an unreduced gamete of R.
raphanistrum and a reduced gamete of B. napus
(RrRrAC, 2n 37 chromosomes) and B1% pollen via-
bility. No hybrids were detected in commercial HR B.
napus fields in Quebec and Alberta (22 114 seedlings
screened, probability of B2105).

Weedy B. rapa. Numerous studies have indicated a high

Erucastrum gallicum. Gene flow from B. napus to E.
gallicum has not been extensively studied. In one report,
a single B. napus E. gallicum hybrid was obtained
under greenhouse conditions, but no hybrids were
detected when E. gallicum served as the maternal parent
(Lefol et al. 1997). Erucastrum gallicum occurrence in
B. napus growing areas of Canada is limited and found
primarily in Saskatchewan. In the same Canadian field
study described above for S. arvensis (Warwick et al.
2003), no E. gallicum B. napus hybrids were detected
in 21 841 E. gallicum seedlings from commercial HR
B. napus fields in Saskatchewan. These results again
indicate a very low probability of inter-specific gene flow
(B2105).
Raphanus raphanistrum. Studies in France and Australia
have indicated that hybridization between R. raphanis-
trum and B. napus is very rare. Only three hybrids were
potential for hybridization between weedy B. rapa and
B. napus. This is not surprising, as B. rapa (AA genome,
2n 20 chromosomes) is one of the progenitor species of
B. napus (AACC genome, 2n 38 chromosomes).
Spontaneous hybridization and introgression between
weedy B. rapa and B. napus was reported in Danish
studies (Jørgensen and Andersen 1994; Jørgensen et al.
1996; Landbo et al. 1996; Hansen et al. 2001, 2003),Uni-
ted States field studies (Halfhill et al. 2002, 2004),
and United Kingdom studies (Wilkinson et al. 2003);
and between cultivated lines of B. rapa and B. napus in
field experiments in Canada (Bing et al. 1996b). Based
on the distribution of herbarium specimens, weedy
B. rapa has a limited distribution as an agricultural
and/or ruderal weed in B. napus growing areas in
Quebec (Simard et al. 2007). Domesticated B. rapa can
also be a weedy volunteer in western Canada. In recent
Canadian studies (Warwick et al. 2003), which include
 974 CANADIAN JOURNAL OF PLANT SCIENCE
data from experimental field trials and commercial HR
B. napus fields, hybridization between weedy B. rapa
and B. napus occurred at a frequency of ca. 7% in two
field experiments where weedy B. rapa plants were
grown at a density of one plant m 2 with HR
B. napus. Brassica rapa B. napus F1 hybrids were
also detected in two weedy B. rapa populations growing
in or near commercial HR B. napus fields in Quebec.
This represented the first reported case globally of
transgene escape into a natural weed population. A
high frequency of hybridization (13.6%) was observed
in one of the weedy B. rapa populations and was likely
due to greater distance between B. rapa plants (i.e., a
thin stand). All F1 hybrids were morphologically similar
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22
to weedy B. rapa, but hybrids were confirmed by the
presence of the herbicide resistance trait, the presence of
species-specific AFLP molecular markers from both
parental species, and a triploid ploidy level (AAC,
2n 29 chromosomes). The F1 hybrids had reduced
pollen viability (ca. 55%) and segregated for both self-
incompatible and self-compatible individuals, the latter
being a B. napus trait. Other researchers have also
shown that F1 hybrids produced from the hybridization
of weedy B. rapa and B. napus were triploid (Metz et al.
1997; Halfhill et al. 2002). Recent studies have docu-
mented the occurrence of F1 hybrids in an additional
For personal use only.
9 sites in Quebec (Simard et al. 2007). Follow-up studies
have documented the persistence and stable introgres-
sion of the HR trait in a weedy Brassica rapa population
at one of the two original 2001 commercial field
hybridization sites (Warwick et al. 2007). Persistence
occurred over a 6-yr period, in the absence of herbicide
selection pressure (with the exception of possible ex-
posure to glyphosate in 2002) and in spite of the fitness
cost associated with hybridization (see below).
Hybrid Fitness (B. rapa B. napus hybrids)
Previous studies of transgenic glufosinate-resistant F1
weedy B. rapa B. napus hybrids (Snow et al. 1999)
indicated no fitness effect in the F1 hybrid. In similar
studies, Hauser et al. (1998a) found that F1 hybrids had
intermediate fitness between the two parental species
based on several combined characteristics, and they
concluded that F1 hybrids were significantly more fit
than weedy B. rapa. In a subsequent study, Hauser et al.
(1998b), found that a fitness penalty occurred in F2 and
backcrossed individuals, although a small percentage of
hybrids were as fit as the weedy parent. The fitness of F1
hybrids may also be frequency dependent (based on
hybrid versus parent ratio), and the experimental design
in future research may need to include the appropriate
ratio of hybrid to parental weedy B. rapa plants to
simulate selection for the hybrids with the highest fitness
(Pertl et al. 2002; Hauser et al. 2003). Preliminary results
from an experimental field study conducted in Ottawa,
ON, in 2005 on two Canadian glyphosate resistant
weedy B. rapa B. napus BC2F2 backcross hybrid
populations under competitive field conditions (with
vs. without wheat; weedy vs. weed-free) suggested that
the hybrid is less fit than the parental weed population,
regardless of the presence of the transgene (Warwick
2006).
Weedy B. rapa B. napus hybrids with the insect
resistance trangene Bt-GFP [Bacillus thuringiensis (Bt)-
green fluorescent protein (GFP)] also showed reduced
fitness/competitiveness (Halfhill et al. 2005). In a non-
competitive greenhouse experiment, both transgenic and
non-transgenic hybrids showed reduced vegetative
growth and seed production. In a field experiment
conducted under two herbivory levels and high intra-
specific competition, transgenic hybrids also produced
less vegetative dry weight and fewer seeds than weedy
B. rapa. In competition experiments with wheat, the
hybrids were the least competitive as compared with
parental Brassica competitors. Again, reduced hybrid
fitness appeared to be independent of transgene pre-
sence.
10. Population Dynamics
Volunteer B. napus and B. rapa population densities in
fields declined quickly after the last canola crop, from
the first (100 to 210 plants m 2), to the second year (0 to
3 plants m 2), to the third year (B1 plant m 2) after
seed bank establishment when seed bank replenishment
was not allowed (Gulden et al. 2003b). The same was
found in field suveys in Saskatchewan (Thomas and
Leeson 1999) and Quebec (Simard et al. 2002) where
seed bank replenishment could occur and residual (after
in-crop weed control) densities of volunteer canola (B.
napus and B. rapa) were typically less than 1 plant m 2 4
to 5 yr after a canola crop. These data suggest that seed
bank replenishment in volunteer populations in fields is
limited. During the growing season, volunteer B. napus
and B. rapa densities are often variable, but tend to be
greatest in the spring. In a pre-weed control spring
survey where volunteer canola was the most abundant
weed, fewer than one and as many as 812 volunteer
canola plants m2 were counted in fields in Manitoba in
1994 (Thomas et al. 1997). In the 85 fields where
volunteer canola was present, the mean density of 85.7
volunteer canola plants m 2 in the spring decreased to
13.7 plants m 2 by autumn. Similar observations were
made by Harker et al. (2006) in a 4-yr study across
western Canada, which showed highly variable seedling
densities among the locations (0 to 179 plants m 2),
despite identical initial seed bank populations. Volun-
teer B. napus densities declined to B5 plants m 2 after
the first year of the study. At some locations, increased
levels of soil disturbance and continuous cropping
promoted volunteer B. napus persistence, while the
effect of seeding date of the wheat crop was negligible
on seedling recruitment and persistence in volunteer B.
napus.
Other factors are also known to affect populations
densities of volunteer canola. In a long-term rotation
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
study in Saskatchewan, volunteer B. napus seedling
densities in the first spring after a canola crop were
between one and two orders of magnitude greater after
an uncharacteristically dry autumn (Légère et al. 2001).
Volunteer B. napus populations were influenced by
landscape slope position in Saskatchewan with a 22-
fold difference in average pre-spraying volunteer
B. napus populations between the upper slope (34 plants
m 2) and lower slope positions (740 plants m 2) 1 yr
after a canola crop (Beckie et al. 2003). Self-thinning has
been observed in B. napus crops in western Canada with
consistent behaviour among genotypes. In an agronomic
performance experiment, van Deynze et al. (1992)
reported mean survival at maturity of 73% when the
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22
seedling recruitment density was 49 plants m 2, whereas
mean survival at maturity was only 55% when seedling
recruitment was 127 plants m 2. Other studies reported
similar results (Morrison et al. 1990).
A number of different models describing the popula-
tion dynamics of volunteer B. napus have been devel-
oped in Europe. Two independent models based on data
primarily collected in the United Kingdom describe the
seed bank dynamics of volunteer B. napus in cropping
systems (Squire et al. 1997; Pekrun et al. 2005). Two
further models developed in the United Kingdom
describe the population dynamics of winter B. napus.
For personal use only.
The first examines the persistence of transgenic volun-
teer B. napus populations in fields (Begg et al. 2006),
while the second describes the spatial population
dynamics of feral B. napus outside agricultural fields
(Crawley and Brown 2004). A fifth model, developed in
France, simulates the effect of cropping system on
transgene escape through pollen flow in B. napus crops
from volunteer and cropped plants (Colbach et al. 2000)
and a sixth model developed in Germany describes
reproduction in volunteer winter B. napus (Gruber and
Claupein 2007).
Little is known about the population dynamics of
volunteer and weedy (naturalized) canola (B. napus and
B. rapa) in non-agricultural areas in Canada. In a recent
survey of railroad beds and roadsides in Saskatchewan
and British Columbia, the primary source and port
destination of canola crops in Canada, Yoshimura et al.
(2006) found that feral populations (i.e., populations
outside arable fields) of B. napus tended to be greater
along roadsides in Saskatchewan and along railroads in
British Columbia, both reflecting the major means of
transportation of commercial canola seed at each
location. Individual plants resistant to glyphosate or
glufosinate comprised a large proportion of the feral
populations which was a reflection of the large acreage
of HR B. napus canola that is currently grown in
Canada. In Manitoba, 16 escaped populations of B.
napus growing along verges of fields and roadways were
monitored from 2004 to 2006 (Knispel et al. 2008). In
these populations, stacking of herbicide-resistance genes
through intraspecific gene flow was observed. In a
survey in Canterbury, New Zealand, Heenan et al.
975
(2004) documented Brassicaceae outside agricultural
fields. Naturalized B. napus populations tended to be
rare and small with the largest along a roadside
composed of about 100 plants. Weedy B. rapa var.
oleifera also was rare in rural areas and was found to be
most abundant at the periphery of urban areas.
Although naturalized populations of B. rapa are self-
sustaining outside arable fields, feral B. napus popula-
tions tend to go extinct after 3 to 8 yr in Europe
(Crawley and Brown 1995; Scott and Wilkinson 1999;
Pessel et al. 2001).
Blackshaw and Dekker (1988) investigated the rela-
tive competitiveness of domesticated Canadian B. napus
to two common weeds using a replacement study.
Brassica napus was less tolerant to interference than
S. arvensis, but more tolerant to interference than
lambsquarters (Chenopodium album L.). The ability to
withstand drought conditions was directly proportional
to the interference potential of each species. Leaf area
development and light interception were closely linked
to the competitive ability of these species (Blackshaw
et al. 1989), while competition for soil nutrients did not
appear to play a role (Blackshaw and Dekker 1988).
In laboratory studies, intra- and interspecific compe-
tition have been shown to influence reproductive traits
in subsequent generations in rapid-cycling B. rapa
differently, with flower number increasing more
under intraspecific competition than under interspecific
competition after two generations (Miller 1995). High
genetic variation was observed in rapid-cycling
B. rapa, but environmental factors also strongly influ-
enced the phenotype of progeny (Karoly and Conner
2000).
11. Response to Herbicides and Other Chemicals
The occurrence of stacking of herbicide resistance genes
in volunteer B. napus in western Canada (next section)
provided the impetus for research on alternative herbi-
cide control options for volunteer canola in fields.
Beckie et al. (2004) examined the susceptibility of one
conventional non-herbicide tolerant canola cultivar,
four single herbicide-resistant cultivars (glyphosate,
glufosinate, imidazolinone, and bromoxynil), and seven
double- or triple-resistant lines to 2,4-D (amine and
ester), MCPA ester and metribuzin. The susceptibility
(90% mortality) to 2,4-D amine and ester, MCPA
ester and metribuzin applied at the two- to three-leaf
stage to all, but the bromoxynil resistant B. napus
genotypes, did not differ, although the sensitivity of
B. napus to 2,4-D amine was much lower when applied
at the later stage (five- to six-leaf). In Saskatchewan,
Kirkland (1997) reported a residual effect of spring 2,4-
D applications that consistently reduced seedling re-
cruitment in B. napus and B. rapa crops seeded shortly
after application of this herbicide. In response to
testimonials of failed control of volunteer B. napus by
phenoxy herbicides in spring in Canada, Légère et al.
(2006) examined the influence of cold acclimation on
 976 CANADIAN JOURNAL OF PLANT SCIENCE
herbicide efficacy on volunteer B. napus seedlings in
greenhouse and field studies in Saskatchewan and
Quebec. Greenhouse studies showed no appreciable
difference in herbicide efficacy of 2,4-D or MCPA in
cold-acclimated compared with unacclimated B. napus
seedlings. The same was observed in the field trials
where herbicide efficacy was influenced more by growth
stage of volunteer B. napus than by the degree of cold
acclimation. There are many (25
30) registered herbi-
cide options to control volunteer canola in the major
cereal crops in western Canada (Anonymous 2008b).
However, in crops such as sunflower (Helianthus annuus
L.), lentil (Lens culinaris L.), chickpea (Cicer arietinum
L.), and field pea (Pisum sativum L.), there are few in-
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crop herbicide options for controlling volunteer canola
(Anonymous 2008b). Carfentrazone, a triazolinone
herbicide recently registered for use in Canada, may be
useful for control of volunteer B. napus in these
phenoxy-sensitive crops (Légère et al. 2006). Imazamox,
a post-emergence applied imidazolinone herbicide, pro-
vided adequate control of domesticated B. rapa at rates
that did not injure field pea in Alberta (Blackshaw
1998). In common buckwheat (Fagopyrum esculentum
Moench), volunteer B. napus and B. rapa may be
controlled with desmedipham, a phenyl-carbamate her-
bicide commonly used in sugarbeet (Beta vulgaris L.)
For personal use only.
(Wall and Smith 1999). Buckwheat tolerance to this
herbicide is approximately five times greater than that of
domesticated B. napus and B. rapa; however, the high
cost of this herbicide at the required rate may be
prohibitive for this use. In western Canada, pre-harvest
applications of glyphosate are becoming more common
in some crops. The intent is to speed and harmonize
maturation of the crop throughout a field, while also
controlling weeds. Pre-harvest application of glyphosate
in a B. napus crop after the pods were formed did not
influence the rate of maturation or seed characteristics
(Darwent et al. 2000).
In Quebec, Maltais and Bouchard (1978) reported
that 2,4-D, linuron, and amino-triazole effectively con-
trolled triazine-resistant weedy B. rapa (see below) in
greenhouse studies. In B. rapa, triazine-resistance is a
maternally inherited trait (Souza Machado et al. 1978;
Souza Machado and Bandeen 1982) and this trait has
been tranferred from these weedy B. rapa (bird rape)
populations to domesticated B. rapa and B. napus
(Beversdorf et al. 1980).
In the Pacific Northwest, Rainbolt et al. (2004) found
that aside from glyphosate, paraquat plus diuron was
the most effective herbicide combination for controlling
herbicide-resistant volunteer B. napus. Glyphosate plus
glufosinate and glyphosate plus paraquat were less
effective. In the United Kingdom, none of the pre-
emergence herbicides registered for use in woodlands
provided effective control of volunteer B. napus in these
areas; however, atrazine in combination with cyanazine
did result in substantial growth reductions of B. napus in
woodlands (Dixon et al. 2006).
Herbicide Resistance
Currently, B. napus genotypes resistant to herbicides
with three different modes of action are available for
commercial production in Canada. Two of these are
transgenic genotypes (glyphosate- and glufosinate-resis-
tant), while resistance to imidazolinones was generated
through mutagenesis. Brassica napus cultivars resistant
to bromoxynil, the fourth chemistry of herbicide resis-
tance in this species, are no longer available commer-
cially. Herbicide resistant genotypes comprise greater
than 80% of the annual acreage of B. napus currently
grown in Canada with genotypes resistant to glyphosate
comprising approximately 50% of that acreage (Canola
Council of Canada 2001). Due to outcrossing in
B. napus (Sections 8 and 9), volunteer B. napus popula-
tions with multiple herbicide resistance have been
documented. Two years after the commercial introduc-
tion of HR genotypes, Hall et al. (2000) found multiple
resistant volunteer B. napus plants in an Alberta field
resulting from inadequate spatial and temporal separa-
tion of three different HR B. napus crops. Two plants
were resistant to all three herbicides with many more
double resistant volunteers. In a study in Saskatchewan,
Beckie et al. (2003) documented the location of all
double HR B. napus volunteers generated via pollen
flow between 11 paired fields 1 yr after a glyphosate-
resistant cultivar was grown in one field and a glufosi-
nate-resistant cultivar was grown in the adjacent field.
Large numbers of double-resistant plants were found,
indicating that gene stacking via outcrossing was
common in B. napus. The spatial pattern of the
distribution of double resistant individuals was not
consistent among and within paired fields. Similar
experiments showing similar results, albeit on a smaller
scale, have been conducted in Europe (Dietz-Pfeilsetter
and Zwerger 2004).
Simard et al. (2005) generated double resistant
B. napus lines (imidazolinone/glufosinate, glyphosate/
glufosinate, imidazolinone/glyphosate) using Canadian
spring B. napus genotypes and evaluated the fitness of
these lines. No significant fitness penalty was detected in
the double resistant lines compared with the single
resistant lines. The glyphosate/glufosinate double resis-
tant plants produced the greatest biomass. Kumar et al.
(1998) generated transgenic Canadian spring B. napus
lines containing the Pat transgene for glufosinate-
resistance and evaluated these lines in field trials in
Saskatchewan. Comparisons of the 19 transgenic lines
with their isogenic susceptible counterparts showed that
in 11 instances, presence of this transgene affected
agronomic attributes (i.e., total biomass, yield, time to
flowering and maturity, oil content, protein content)
negatively and this effect became more pronounced in
lines with more copies of the transgene. These findings
indicated that transgene location in the genome and
accumulation of transgenes may affect fitness in volun-
teer populations of B. napus. In the United States,
Stewart et al. (1996) produced a series of transgenic
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
B. napus lines containing the insecticidal Bt transgene
and evaluated these in the laboratory and the field
(Stewart et al. 1997). Under insect selection pressure in
the field, transgenic plants had increased fitness due to
reduced defoliation compared with their non-transgenic
counterparts when allowed to naturalize following
cultivation. In a Danish study, no fitness penalty was
found when a single glufosinate-resistance transgene was
introgressed from domesticated transgenic B. napus into
weedy B. rapa (Snow et al. 1999).
In weedy B. rapa in Canada and elsewhere, acquired
resistance to triazines is clearly associated with a fitness
penalty (Mapplebeck et al. 1982; Plowman et al. 1999).
In natural habitats, triazine-resistant weedy B. rapa
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 201.189.166.52 on 11/15/22
germinated slower (Mapplebeck et al. 1982), the max-
imum depth of seedling recruitment was less (Mapple-
beck et al. 1982), and the frequency of triazine-resistant
plants rapidly decreased in a weedy population in
natural habitats (Plowman et al. 1999). Carbon ex-
change rates were lower (up to 28%) in triazine-resistant
domesticated B. napus and B. rapa genotypes compared
with triazine-susceptible genotypes, despite similar
chlorophyll a and b contents (Hobbs 1987). Burke
et al. (1982) observed differences in chlorophyll a:b
ratios and chloroplast fatty acid composition among
triazine-resistant and triazine-susceptible biotypes of
For personal use only.
weedy B. rapa in the United States. An interaction
between temperature and light levels in the competitive
outcome between triazine-tolerant and triazine-suscep-
tible rapid-cycling domesticated B. rapa was observed.
At low light and high temperatures, the resistant biotype
was a better competitor than the susceptible biotype
(Plowman and Richards 1997).
Friesen et al. (2003) examined 27 commercially
available certified seedlots of conventional B. napus for
contamination with herbicide-resistant seed. Some level
of glyphosate-resistance presence (up to 4%) was found
in almost all seedlots and 14 seedlots exceeded the
0.25% impurity threshold set for certified seedlots.
12. Response to Other Human Manipulations
Volunteer canola is associated with crop management
systems in western Canada. Leeson et al. (2000)
investigated the distribution of residual weeds in fields
classified into one of seven management systems in
Saskatchewan. The management systems were categor-
ized according to crop rotation (fallow, diversified-
annual grain, diversified-grain-forage) and intensity of
external inputs (organic, moderate, high). Over the 3-yr
study, volunteer canola (B. napus and B. rapa) was most
prevalent in high-input fallow and moderate input
diversified-annual grain management systems. These
systems also are those in which canola is most likely
grown as a crop.
In eastern Canada, volunteer B. napus seedling
recruitment was greater under zero-tillage (4 plants
m 2) compared with using a chisel or a moldboard plow
(B2 plants m 2) in the first spring following a canola
977
crop (Simard and Légère 2003). In all treatments,
however, volunteer B. napus seedling density was quite
low. In a study in western Canada, tillage did not affect
total volunteer B. napus and B. rapa seedling recruit-
ment, although the main flush of seedling recruitment in
zero-tillage was delayed until after the first substantial
spring rain (Gulden et al. 2003b). Other reasons for
these apparent regional differences include much lower
tillage intensities in conventional tillage in western
Canada compared with eastern Canada and inherent
differences in climate between these regions. In a
synthesis of results from 10 tillage studies on the
Canadian prairies, Thomas et al. (2004) concluded that
volunteer canola was associated with reduced- and zero-
tillage systems and not with conventional-tillage sys-
tems. Significant associations were only found in 21% of
the site-years, but the trend in the remaining site-years
provided support for an association with reduced and
zero tillage. Crop rotation was not considered in the
analysis. In Europe, Bowerman (1993) reported as early
as 1993 that management practices influence volunteer
B. napus seedling recruitment after a canola crop.
Burning the residue and/or shallow incorporation of
the residue of the canola crop resulted in fewer volunteer
seedlings (46 and 33 plants m 2, respectively) than
ploughing (223 plants m 2) 1 yr after seed bank
initiation. Studies on volunteer winter B. napus have
demonstrated greater seed bank persistence when seeds
were buried immediately after seed shed compared with
delayed or no tillage (Pekrun and Lutman 1998). Even
delayed ploughing resulted in a significantly lower
winter B. napus seed bank the following spring com-
pared with ploughing immediately after seed shed
(Pekrun and Lutman 1998), indicating increased seed
mortality or predation when seeds were left on the soil
surface. A Swiss study examined the combined effects of
preceding crop, tillage system and time of herbicide
application on weed populations using a winter wheat-
oilseed rape-winter wheat-maize rotation (Streit et al.
2003). Volunteer B. napus was one of the dominant
species in the weed community in winter wheat follow-
ing oilseed rape and was significantly related to the no-
tillage treatment.
Kirkland (1993) investigated the influence of barley
(Hordeum vulgare L.) competition on biomass produc-
tion of domesticated volunteer B. rapa in Saskatchewan
and showed biomass reductions of 95% or more when
barley was planted at 220 kg ha1 in 11-cm rows
compared with when barley was planted at 50 kg ha1
in 46-cm rows. In an interference study between
domesticated B. rapa and tartary buckwheat (Fago-
pyrum tataricum L.) where B. rapa was grown as the
crop, O’Donovan (1994) found that B. rapa reproduc-
tive output was least affected at high B. rapa densities
(200 plants m 2) compared with lower densities (50 and
100 plants m 2).
Blackshaw et al. (2003) compared shoot and root
growth of 23 agricultural weeds and the crop B. napus to
 978 CANADIAN JOURNAL OF PLANT SCIENCE
a range of N fertilizer rates from 0 to 240 mg kg1 soil
in a controlled environment study. Shoot biomass
production of B. napus and of 10 other species [e.g.
wild mustard, redroot pigweed (Amaranthus retroflexus
L.) and lambsquarters] responded to increasing amounts
of N. Root biomass production of B. napus was less
responsive than shoot biomass. Brassica napus and six
other species had greater than 90% uptake of available
N. The responses of the same species to P fertilizer levels
from 0 up to 60 mg kg1 soil were studied in a similar
controlled environment study (Blackshaw et al. 2004).
Brassica napus, along with wild mustard, kochia [Kochia
scoparia (L.) Roth] and Russian thistle (Salsola pestifer
A. Nels.), exhibited the least response in shoot biomass
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to added P. Root biomass in B. napus was more
responsive than shoot biomass. Brassica napus, redroot
pigweed and wild mustard were in the top group of
species that took up greater than 40% of the available
P. Similar controlled environmental studies have not
been conducted on S and K, but S is the most limiting
plant nutrient after N and P as Brassica napus has a high
S requirement. Application of sulphate at about
15
30 kg S ha 1 is usually sufficient for a B. napus
crop on S-deficient soils (Malhi et al. 2005). The balance
between S and N is also important. Although it is
recommended to fertilize with N and S in a 7:1 ratio
For personal use only.
(Thomas 2003) to achieve optimum yield and seed
quality, this may not be appropriate for the production
of new high-yielding canola hybrids. Karamanos et al.
(2005) found that the N requirement for optimum yield
of hybrid canola cultivars were higher than those of
conventional cultivars and that once S deficiency was
corrected there was little need to balance N and S at any
particular ratio. Potassium is required in large amounts
by canola but fertilization trials in western Canada have
rarely shown a response to the addition of K even when
grown on soils with very low extractable K levels (Doyle
and Cowell 1993). The lack of response is due to ample
soil K reserves in western Canada and the crops ability
to absorb K from the soil (Thomas 2003). The growth,
competitive ability and reproductive potential of volun-
teer canola plants in subsequent crops will be influenced
by fertility levels, but this subject has not been
researched.
Harker et al. (2005) investigated the effects of varying
frequencies of glyphosate-resistant wheat and B. napus
canola in a wheat-canola-wheat-field pea rotation (five
different rotations) on weed populations at six sites in
western Canada. Crops were grown under either a
conventional tillage or a low soil-disturbance system
with early or late seeding dates. Weed populations were
assessed in the non-resistant pea crop in the final year of
the study. The response of volunteer canola was
inconsistent; populations were not significantly asso-
ciated with any rotation at three of the sites, were
associated with the rotation using no glyphosate-resis-
tant crops at two of the sites and, at the sixth site, were
associated with the rotation using both glyphosate-
resistant crops. Volunteer canola densities were signifi-
cantly higher in the conventional tillage systems at two
sites in some of the five rotations and in early-seeded
treatments at one site. Rotations including glyphosate-
resistant crops did not increase the risk from volunteer
canola. Harker et al. (2006) compared the density of
canola volunteers in a wheat-field pea-barley rotation
and a fallow-field pea-fallow rotation with five different
seeding system combinations comprised of seeding date
and soil disturbance level at seven sites across western
Canada. Treatment effects were not consistent among
sites and only one comparison was significant in 1 yr
where greater volunteer canola densities tended to be
associated with increasing soil disturbance in the con-
tinuous cropping rotation.
13. Responses to Herbivory, Disease and Higher
Plant Parasites
(a) Herbivory
(i) Mammals * Deer graze on canola plants in Canada.
Canola was tested in Nova Scotia as a lure crop to
protect carrots from grazing by white-tailed deer
(Odocoileus virginianus). Staggered plantings of canola,
which continuously produced flowers, was the most
effective crop that attracted deer away from carrots
(Schwab et al. 2001). The Saskatchewan Crop Insurance
program expanded the crop establishment benefit to
include damage from Richardson’s ground squirrels
(Spermophilus richardsonii) in 2007. Under the 2007
pilot program, 176 producers in southwestern Saskatch-
ewan filed claims and were paid damages totalling
$200 000 in canola (D. Wotherspoon, personal commu-
nication).
(ii) Birds and/or Other Vertebrates * There is no
published evidence of birds feeding in canola fields in
Canada; however, in 2006 the Saskatchewan Crop
Insurance Corporation paid out $86 000 in damages
caused by migratory waterfowl in canola (P. Screpnek,
personal communication). The use of canola fields by
sandhill crane, Grus canadensis (L.), in Saskatchewan
was too low to be included in an analysis of foraging
preferences during their southward migration in the
autumn (Sugden et al. 1988). Hall (2007) indicates that
canola in Ontario may be grazed by wild turkeys.
American goldfinches (Carduelis tristis L.) fed so
intensively on mature canola plants in experimental
plots in St-Davi-de-Lévis that nets were required to
salvage crop yields (M.-J. Simard, personal communica-
tion). In the United Kingdom, yield losses in autumn-
sown oilseed rape due to mute swans (Cygnus olor L.)
were as high as 33.7% at high grazing intensity (Parrott
and McKay 2001), to brent geese (Branta bernicla
bernicla L.) as high as 27.5% (McKay et al. 1993), and
to woodpigeon (Palumba columbus L.) averaged 9%
(Inglis et al. 1989). These birds overwinter in the United
Kingdom and feed from December to April.
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
(iii) Insects * A number of insect pests attack the
cotyledons, leaves, stems, apical and flower buds,
flowers, pods, seeds and roots of canola and rape plants
(Thomas 2003). Pests in the order Lepidoptera include
alfalfa looper (Autographa californica Speyer), beet
webworm (Loxostege sticticalis L.), bertha armyworm
(Mamestra configurata Walker), cabbageworms (Pieris
species), clover cutworms (Dicestra trifolii Hufnagel),
diamondback moth (Plutella xylostella L.), painted lady
butterfly (Vanessa cardui L.) pale western cutworm
(Agrotis orthogonia Morrison), and redbacked cutworm
(Euxoa ochrogaster Guenée). Insect pests in the order
Coleoptera include cabbage seedpod weevil (Ceutor-
hynchus obstrictus Marsham), crucifer flea beetle (Phyl-
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lotreta cruciferae Goeze), striped flea beetle (Phyllotreta
striolata Fabricius), red turnip beetle (Entomoscelis
americana Brown); in the order Homoptera: aphids
(Brevicoryne brassicae L. and other species); in the order
Hemiptera: plant bugs (Lygus species); and in the order
Diptera: cabbage root maggot (Delia radicum L.) and
turnip root maggot (Delia floralis Fallén).
The cabbage seedpod weevil (C. obstrictus), first
reported in Alberta in 1995, is predicted to become a
major pest throughout the entire canola production area
in western Canada (Dosdall et al. 2002). The lesser
migratory grasshopper (Melanoplus sanguinipes Fabri-
For personal use only.
cius) has been shown in controlled experiments to
reduce the yield of both B. napus and B. rapa (Olfert
and Weiss 2002). The bronzed or rape blossom beetle
(Meligethes viridescens Fabricius) is a common pest of
spring-seeded B. napus and B. rapa in Europe that is
known to be present in eastern Canada and has the
potential to spread westward to the prairies (Mason
et al. 2003). Recently swede midge (Contarinia nasturtii
Kieffer), a pest of cruciferous species including canola,
was accidentally introduced into North America and is
now found in a few areas in Ontario and Quebec (Olfert
et al. 2006) and in Saskatchewan in 2007 (J. Soroka:
personal communication). Bioclimatic models indicated
that this pest could potentially become established in all
provinces of Canada but the greatest risk would be in
southern Ontario and southern Quebec.
Burgess and Weegar (1988) surveyed insects in the
order Thysanoptera in domesticated canola fields in
Saskatchewan and Alberta. They found flower thrips
(Frankliniella tritici (Fitch)), onion thrips (Thrips tabaci
Lindeman), and white flower thrips (T. vulgatissimus
Haliday) in these fields, however, the extent of the
damage to seed production caused by these species is not
known.
Kaminski (2002) compiled a comprehensive list of
pest and beneficial insects found during extensive
surveys of canola fields in the prairie provinces but
over 50% of the taxa were identified only to the genus
level. At least 30 insects have been identified feeding on
canola plants in Australia but only a few are considered
major pests (Gu et al. 2007). Although the species
diversity is high, little is known about many of these
979
insect populations and their relationship with arthropod
communities in adjacent crops and other habitats (Lamb
1989).
It is not known if volunteer canola plants contribute
to the diversity and structure of insect populations in
agricultural habitats, but a recent study has shown that
maintaining small populations of weeds in the canola
crop has the potential to reduce root maggot infesta-
tions (Dosdall et al. 2003). The incidence and damage of
the crucifer and striped flea beetles, cabbage seedpod
weevil and lygus plant bugs were similar among
herbicide-tolerant and conventional cultivars of canola
(Cárcamo and Blackshaw 2007).
(iv) Nematodes and/or Other Invertebrates * Nematodes
are not known to cause yield loss in B. napus and B. rapa
crops grown in Canada, although under greenhouse
conditions in Quebec B. napus (cv. HL99) was a good
host for the root-lesion nematode Pratylenchus pene-
trans (Cobb) Filipjev and Schuurmans Stekhoven,
increasing nematode numbers 9.3 times (Bélair et al.
2002). Nielsen et al. (2003) tested many B. napus (both
spring and winter types) and B. rapa (winter) cultivars
grown in the north-central region of the United States
and found that none had any resistance to sugar beet
cyst nematode (Heterodera schachtii Schmidt). They
concluded that if canola is grown in rotation with sugar
beet, the nematode problem may increase. The cabbage
cyst nematode, Heterodera cruciferae Franklin, and
root-lesion nematodes, Pratylenchus spp., have been
shown to cause damage to oilseed rape crops in the
United Kingdom (Evans and Webb 1989). Damage to
B. napus seedlings by the oat race of stem nematode,
Ditylenchus dipsaci (Kühn) Filipjev, has been observed
in a few fields in South Australia (Taylor and Szot
2000). Pratylenchus neglectus and P. thornei have been
found to infect B. napus in Iran (Fatemy et al. 2006).
Brassica napus was found to be susceptible to P.
neglectus (Taylor et al. 2000) and moderately resistant
to P. thornei (Hollaway et al. 2000) in field trials in
southeastern Australia.
Slugs and snails have not caused damage to canola
crops in western Canada, but slugs (specific species not
mentioned) have been a problem recently in winter canola
production in Ontario (Hall 2007). In recent years slugs
have been responsible for significant yield reductions in
canola in western Europe (Moens and Glen 2002). The
increase in slug damage was attributed to greater use of
reduced tillage systems, more crop residue left on fields,
planting of winter canola and use of more susceptible
varieties. Seedlings up to the four-leaf stage are very
sensitive to slug grazing. The number of canola seedlings
attacked and the amount of leaf area destroyed by the
grey garden slug, Deroceras reticulatum Müller, is in-
versely proportional to the concentration of glucosino-
lates in the cotyledons of seedlings. According to the 2004
pesticide use survey in Great Britain, 25% of the canola
area was treated with a molluscicides in 2004, down from
 980 CANADIAN JOURNAL OF PLANT SCIENCE
56% in 2002 (Garthwaite et al. 2005). The feeding
preferences of another abundant slug in Europe, Arion
lusitanicus Mabille, were tested in a laboratory study
(Briner and Frank 1998) and B. napus was the most
palatable species of the 78 plant species tested. The grey
garden slug is also a serious pest of canola in the high-
rainfall cropping areas in Australia (Nash et al. 2007).
(b) Diseases
(i) Fungi * Canola is susceptible to several diseases
caused by fungi, as catalogued by Rimmer et al. (2003).
Damping-off and seedling blight are major diseases
across the prairies that are caused by soil-borne fungi
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(Rhizoctonia solani Kühn, Fusarium and Pythium spp.)
that survive mainly on crop residue. Rhizoctonia solani
and Fusarium are also the cause of brown girdling root
rot and foot rot. Brown girdling root rot is the major
disease of canola in the Peace River region of Alberta
and British Columbia and is mainly caused by a strain of
R. solani (Anastomosis Group 2). Brassica rapa cultivars
are highly susceptible while B. napus cultivars have
greater tolerance. Foot rot is a disease of minor
importance in the northern parkland zone of the prairies
and is usually caused by a different strain of R. solani
(Anastomosis Group 4). Alternaria black spot or grey
For personal use only.
leaf spot is a common disease that occurs every year
with varying severity. The disease is caused mostly by
Alternaria brassicae (Berk.) Sacc. and A. raphani Groves
& Skolko but A. alternata Keissl. is also associated with
the disease. Brassica rapa is more susceptible than
B. napus. Although a weakly aggressive type of blackleg
caused by Leptosphaeria biglobosa R. A. Shoemaker &
H. Brun has been known to infect rape/canola in
western Canada for many years, an aggressive type of
the disease caused by L. maculans (Desmaz.) Ces. and
De Not was first found in Saskatchewan in 1975 and is
now present across the prairies. Most B. napus cultivars
are now at least moderately resistant but almost all
B. rapa cultivars are susceptible. Sclerotinia stem rot
caused by the fungus Sclerotinia sclerotiorum (Lib.) de
Bary occurs in canola crops throughout Canada and is
most prevalent in higher rainfall areas. Most canola
cultivars are susceptible to the disease; however, apeta-
lous cultivars are much less susceptible.
Diseases of lesser importance include grey stem and
white leaf spot that is caused by the fungus Pseudocer-
osporella capsellae (Ellis & Everh.) Deighton. This
disease is widespread across northern areas of the
prairies but rarely affects crop yields because it develops
late in the growing season. Brassica rapa is more
susceptible than B. napus. White rust or staghead is
caused by Albugo candida (Pers.) Kunze and is a
common disease of B. rapa, since most cultivars are
susceptible and B. napus cultivars are highly resistant.
Downy mildew is caused by the fungus Peronospera
parasitica [(Pers.:Fr.) Fr.) and is nearly always asso-
ciated with white rust. Three additional diseases that do
not occur on canola crops in Canada but are important
in other countries are verticillium wilt caused by
Verticillium dahliae Kleb., light leaf spot caused by the
Pyrenopeziza brassicae Sutton & Rawlinson and white
leaf spot caused by Mycosphaerella capsellae A. J.
Inman & Sivanesan (Rimmer and Buchwaldt 1995).
Other fungi that have been found on B. napus and B.
rapa are listed in Ginns (1986).
Clubroot and fusarium wilt are two relatively new
diseases in prairie canola crops. Clubroot is caused by
the soil-borne fungus Plasmodiophora brassicae Woro-
nin. The disease was not found on canola in Canada
until 2003 when it was reported for the first time in fields
in an area north of Edmonton (Tewari et al. 2005).
A survey in 2006 indicated that the disease was more
widely distributed in the area surrounding Edmonton
(Strelkov et al. 2007). Clubroot was added as a declared
pest to the Agricultural Pests Act of Alberta in April
2007 (Government of Alberta 2007) in an attempt to
minimize the spread and build-up of this disease. Two
fungi, Fusarium avenaceum Sacc. and F. oxysporum
Schlechtend, are associated with fusarium wilt that
was first reported in the Peace River area of Alberta
in 1999 (Rimmer et al. 2003). Subsequent surveys
in 2000 and 2001 reported that the disease was common
in east central Alberta and present in a few fields in
Manitoba and Saskatchewan. In the 2002 survey for
diseases in Manitoba the incidence of fusarium wilt
was B2% of the B. napus fields (Lange and McLaren
2002) but increased to 21 and 18% in 2005 and 2006,
respectively, and was restricted to the southwest region
of Manitoba (McLaren et al. 2007). Fusarium wilt was
not found in the 2006 survey of B. napus fields in
Saskatchewan (Pearse et al. 2007).
(ii) Bacteria * Bacterial leaf spot caused by Pseudomo-
nas syringae pv. maculicola (McCulloch) Young, Dye &
Wilkie can cause a leaf blight on B. rapa (Rimmer
et al. 2007). Other bacterial diseases are known to be
associated with canola worldwide (APSnet 2001), but
these diseases are either not present or are of little
concern in Canada. Aster yellows is a phytoplasmal
disease spread by leafhoppers that is rarely a disease of
concern in Canadian canola crops (Pearse et al. 2007).
Surveys in B. napus and B. rapa fields in Saskatchewan
between 2001 and 2005 indicated that aster yellows
phytoplasma were found in asymptomatic plants
throughout the province and the leafhopper Macrosteles
quadrilineatus Forbes was the most commonly found
vector (Olivier et al. 2007). The specific strains of
phytoplasma associated with aster yellows in B. rapa
in Saskatchewan have been identified as belonging to
the 16SrI-A and 16SrI-B subgroups (Olivier et al. 2006).
(iii) Viruses * Viral diseases of canola are of minor
concern in Canada but seed yield losses in canola of 10

15% have been reported from beet western yellows
virus (BWYV) in the United Kingdom, Germany and
 GULDEN ET AL. * BRASSICA NAPUS L. AND B. RAPA L.
Australia (Rimmer et al. 2007). A survey over 2 yr of B.
napus (canola) fields in Australia (Coutts and Jones 2000)
indicated that BWYV was the most commonly occurring
virus (61% of fields), cauliflower mosaic virus (CaMV)
was found less frequently (22%), and turnip mosaic virus
(TuMV) was infrequent (4%). All three viruses were
detected in 2% of the fields. The incidence of infected
plants within a field was generally low with no virus
detected in 31% of fields. BWYV was found to infest
volunteer canola (B. napus) growing in roadside verges at
very low levels in Australia and the aphid vectors for the
disease were also present on volunteer canola (Coutts et
al. 2006). Combined damage from BMYV and Myzus
persicae (green peach aphid) reduced reproductive output
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by up to 50% and altered seed quality in B. napus crops in
Australia (Jones et al. 2007). Nearly all of the Australian
canola cultivars of B. napus tested were found to be
resistant to TuMV (Coutts et al. 2007).
In the United States, weedy B. rapa was a host species
of BWYV in vegetable-sugar beet seed production
regions of western Oregon (Hampton et al. 1998) and
the tobacco streak virus (TSV) in tomato fields in
California (Cupertino et al. 1984). In the northwestern
United States, Brassica napus and B. rapa (winter
rapeseed) served as overwintering hosts of BWYV but
not for potato leafroll virus (PLRV), the two viruses
For personal use only.
involved in potato leafroll disease (Thomas et al. 1993).
Susceptibility to turnip mosaic viruses (TuMV), a
serious disease of rutabaga (B. napus), was surveyed in
171 weed species in southern Ontario (Stobbs and
Stirling 1990). Ten of the 13 crucifer species surveyed,
including weedy B. rapa and B. napus subsp. oleifera
(winter rapeseed), were found to be susceptible to the
common strain, TuMV-S1, and could potentially serve
as an overwintering reservoir of the virus. This strain
was transmissible to both species, through both aphid
and mechanical inoculation. Rao et al. (1977) found
TuMV infrequently in a 1975 survey of B. rapa crop
fields in the Edmonton area of Alberta.
Viral host ranges can be extended by the presence of
other viral systems. For example, Sehgal et al. (1993)
found that replication and encapsidation of the viroid-
like satellite RNA of lucerne transient streak virus
(LTSV) were supported in the presence of turnip rosette
sobemovirus (TROV) in crucifer species S. arvensis,
B. rapa, and R. raphanistrum, but not in T. arvense,
indicating that host species markedly influenced this
interaction.
(c) Higher Plant Parasites * There are no reported
higher plant parasites of B. napus or B. rapa.
Acharya, S. N., Dueck, J. and Downey, R. K. 1983. Selection
and heritability studies on canola/rapeseed for low tempera-
ture germination. Can. J. Plant Sci. 63: 377
384.
Agren, J. and Schemke, D. W. 1994. Evolution of trichome
number in a naturalized population of Brassica rapa. Am. Nat.
143: 1
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Ajwa, H. A., Ba_uelos, G. S. and Mayland, H. F. 1998.
Selenium uptake by plants from soils amended with inorganic
and organic materials. J. Environ. Qual. 27: 1218
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Akbar, M. A. 1989. Resynthesis of Brassica napus aiming for
improved earliness and carried out by different approaches.
Hereditas (Lund) 111: 239
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Alam, M., Ahmad, H., Quazi, M. H. and Khawaja, H. I. T.
1992. Cross compatability studies within the genus Brassica 1.
Amphidiplpoid combinations. Sci. Khyber 5: 89
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Alex, J. F. 1964. Weeds of tomato and corn fields in two
regions of Ontario. Weed Res. 4: 308
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Alex, J. F. 1966. Survey of weeds of cultivated land in the
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