The Biology of Brassica carinata (A.) Braun (Abyssinian cabbage)

Biology Document BIO2017-02: A companion document to Directive 94-08 (Dir94-08), Assessment Criteria for Determining Environmental Safety of Plant with Novel Traits

Plant and Biotechnology Risk Assessment Unit
Plant Health Science Division, Canadian Food Inspection Agency
Ottawa, Ontario

Table of Contents

1. General Administrative Information

1.1 Background

The Canadian Food Inspection Agency's Plant and Biotechnology Risk Assessment (PBRA) Unit is responsible for assessing the potential risk to the environment from the release of plants with novel traits (PNTs) into the Canadian environment. The PBRA Unit is also responsible for assessing the pest potential of plant import and plant species new to Canada.

Risk assessments conducted by the PBRA Unit require biological information about the plant species being assessed. Therefore, these assessments can be done in conjunction with species-specific biology documents that provide the necessary biological information. When a PNT is assessed, these biology documents serve as companion documents to Dir94-08: Assessment Criteria for Determining Environmental Safety of Plants with Novel Traits.

1.2 Scope

This document is intended to provide background information on the biology of Brassica carinata, its identity, geographical distribution, reproductive biology, related species, the potential for gene introgression from B. carinata into relatives, and details of the life forms with which it interacts.

Such information will be used during risk assessments conducted by the PBRA Unit. Specifically, it may be used to characterize the potential risk from the release of the plant into the Canadian environment with regard to weediness/invasiveness, gene flow, plant pest properties, impacts on other organisms, and impact on biodiversity.

2. Identity

2.1 Name

Brassica carinata A. Braun

2.2 Family

Brassicaceae (alt. Cruciferae), commonly known as the mustard family.

2.3 Synonym(s)

Synonym for Brassica carinata is Brassica integrifolia (H. West) Thellung var. carinata (A. Braun) O.E. Schulz. (USDA, ARS 2014).

2.4 Common name(s)

Brassica carinata is commonly known as Abyssinian cabbage, Abyssinian mustard, African cabbage, Ethiopian kale, Ethiopian mustard, Ethiopian rape, mustard collard, chou Éthiopien, moutard d'Abyssinie (USDA ARS 2014).

In the peer-reviewed literature, B. carinata has been referred to as gomenzer (Getinet 1996), African sarson (Gill and Bains 2008), TexSel greens (Stephens et al. 1975), karan rai (Chauhan et al. 2011), peela raya (Anwar et al. 1993), raya (Chaudhary and Ullah 1995), and yebesha gomen (Asfaw 1995).

2.5 Taxonomy and genetics

The genus Brassica is a member of the tribe Brassiceae, within the mustard family (Brassicaceae; Warwick et al. 2009). It includes several economically important oilseed crop species: B. juncea (L.) Czern. (brown mustard), B. napus L. (rape, Argentine canola), B. nigra (L.) W.D.J. Koch (black mustard), and B. rapa L. (field mustard, Polish canola). The genus Brassica also includes food crops in B. oleracea L., including cabbage, broccoli, cauliflower, Brussels sprouts, kohlrabi, and kale; B. rapa, including leafy-type, such as Pak Choi and Chinese cabbage, and rapiferous-type, such as turnip; and B. napus, including rutabaga.

Brassica carinata is an amphiploid species (BBCC, 2n = 34; Prakash et al. 2012). It is thought to be derived from interspecific hybridization of two diploid species. The B genome is from Brassica nigra (BB, 2n = 16) and the C genome is from Brassica oleracea (CC, 2n = 18; Prakash et al. 2012). The Triangle of U (U 1935) describes the close genetic relationship between amphidiploid species B. carinata, B. juncea and B. napus, and diploid species B. nigra, B. rapa, and B. oleracea.

Taxonomic position (USDA, NRCS, 2014):

Kingdom: Plantae (plants)
Subkingdom: Tracheobinta (vascular plants)
Superdivision: Spermatophyta (seed plants)
Division: Magnoliophyta (flowering plants)
Class: Magnoliopsida (dicotyledons)
Subclass: Dilleniidae
Order: Capparales
Family: Brassicaceae (mustard family)
Tribe: Brassiceae
Genus: Brassica L. (Mustard)
Species: Brassica carinata A. Braun

2.6 General description

Brassica carinata is a herbaceous annual with a determinate growth habit (Zanetti et al. 2013). Plants have an erect bearing, averaging 1.4 m in height. Plants are highly branched, with a well-developed tap root and extensive rooting system (Barro and Martin 1999).

Seeds are globose, 1–1.5 mm in diameter and finely reticulated (Mnzava and Schippers 2004). They vary from yellow to yellow-brown to brown in colour (Getinet 1986; Rahman and Tahir 2010). The seeds are rich in oil, containing 25–47% depending on the cultivar and growth conditions (Mnzava and Schippers 2007; Cardone et al. 2003; Taylor et al. 2010).

Seeds germinate epigeally, with two 2–3 cm heart-shaped cotyledons (Seegeler 1983; Mnzava and Schippers 2007). Stems are up to 2 cm in diameter, glabrous and usually waxy (Seegeler 1983). Leaves are alternate, glabrous to slightly hairy and often waxy. Lower leaf-blades are large (up to 20 cm long and 10 cm wide) and ovate to oblong with 1–3 deep lobes (Seegeler 1983). Lower leaves are green above and paler or grayish beneath, with veins that may be purple or light-green. Leaves that are higher on the plant are gradually smaller, narrower, less coloured, less waxy, and have fewer lobes. Leaves have a short petiole (Mnzava and Schippers 2007) and trichomes are simple (Al-Shehbaz 2012).

Inflorescenses are highly branched, loose, compound racemes (Seegeler 1983), with flowers that are actinomorphic and perfect (Mnzava and Schnippers 2007). Pedicels are cylindrical and short (5–6 mm long) (Seegeler 1983). There are four light green sepals (4–7 mm long), alternating with four cream to yellow petals (6–10 mm long). Flowers have six stamens (two short outer, and four longer inner). Four floral nectaries are present, two opposite the outer stamens and two alternating with these, between two inner stamens.

Fruits are nondehiscent siliques, which are usually less than 5 cm long, with a 2–7 mm conical beak and may be straight or curved (Seegeler 1983). Siliques contain up to 20 seeds and are remarkably shatter-resistant due to their thick and highly lignified valve margins (Barro and Martin 1999; Banga et al. 2011). Siliques are green when immature and gradually become light brown during maturation.

Seed oil composition varies depending on cultivar and growth conditions, but generally contains: 35–44% erucic acid, 15–22% linoleic acid, 16–20% linolenic acid, 10–12% oleic acid, 7–9% eicosenoic acid, 2–4% palmitic acids (Mnzava and Schnippers 2007). Seeds contain high protein (25–45%) and glucosinolate (150 mmol g-1) content (Getinet et al. 1996; Getinet et al. 1997).

3. Geographical Distribution

3.1 Origin and history of introduction

Brassica carinata is thought to have originated in the highland plateaus of Ethiopia and adjoining parts of East Africa and the Mediterranean coast. Evidence supporting this hypothesis involves the parental species, B. nigra and B. oleracea, being sympatric in these regions during the period that B. carinata was thought to have emerged (Alemayehu and Becker 2002). Cultivation of B. carinata is hypothesized to have started in Ethiopia near 4000 BC (Alemayehu and Becker 2002), although precise information about its domestication is lacking, and cultivation may be more recent (Prakash et al. 2012).

Moderncultivation of B. carinata has experienced marginal growth in southern Europe, Australia, and India (Prakash et al. 2012). However, commercial cultivation remains mostly limited to Ethiopia and neighbouring countries (Marillia et al. 2014), generally taking place on farms of an area less than 2 ha (Seegeler 1983).

Interest in growing B. carinata in Canada, as well as other semi-arid areas throughout the world, began in the mid-1980s. The crop was assessed as a potential alternative to existing oilseed crops in western Canada (Getinet 1986; Getinet et al. 1996; Rakow and Getinet 1998).

3.2 Native range

Africa
Saudi Arabia, Yemen, Ethiopia, Eritrea, Kenya, Rwanda, Uganda, and Tanzania (Warwick et al. 2009; USDA, ARS 2014)

3.3 Introduced range

Africa
Brassica carinata has been reported in Botswana, Cameroon, Côte d'Ivoire, Madagascar, Malawi, Mozambique, Sudan, Democratic Republic of the Congo, Zambia, and Zimbabwe (USDA, NRCS 2014)
Asia
B. carinata has been introduced to India and Pakistan (Malik 1990; Chauhan et al. 2011; Lal et al. 2013; Zada et al. 2013).
Australia
B. carinata has been introduced and is cultivated (Khangura and Aberra 2006).
Europe
B. carinata has been reported in the United Kingdom (Font et al. 2004), Greece (Namatov et al. 2000), Italy (Cardone et al. 2003; Matthäus and Angelini 2005), and Spain (Bouaid et al. 2005; Gasol et al. 2007; Martínez-Lozano et al. 2009; Alcántara et al. 2011)
North America
B. carinata has been cultivated in the Canada (Saskatchewan, Manitoba, Alberta) and the United States (Montana, North and South Dakota, Wyoming, Nebraska, Kansas, Oklahoma, Texas, Louisiana, Mississippi, Alabama, Georgia, Florida) (Getinet 1986; Sask Mustard 2013; NRC 2013).
South America
B. carinata has been grown for experimental purposes in Chile and Uruguay (NRC 2013; Seepaul et al. 2015).

3.4 Potential range in North America

At present,the hardiness of Brassica carinata has yet to be determined. Similar to other Brassica crops, B. carinata grows well in semi-arid environments and is a cool season crop (Marillia et al. 2014). When assessed as a weed, the potential range of B. carinata includes plant hardiness zone 9 (Magarey et al. 2008); however, field tests of B. carinata varieties have been successful across Canada, in Montana and North Dakota, and in southern United States such as Mississippi and Florida (Marillia et al. 2014), which indicates B. carinata can be cultivated in plant hardiness zones 4 through 9 (Magarey et al. 2008).

3.5 Habitat

Brassica carinata grows well in its native habitat, on the highland plateaus of Ethiopia (Seegeler 1983), in cool (14–18°C), moist growing conditions (600–1000 mm average annual rainfall), a long growing season (180 days), and at elevation (2200–2800 m above sea level (Asamenew et al. 1993; Alemayehu and Becker 2002)).

B. carinata grows well in semi-arid climates, on cultivated farmlands, and on marginal lands (Johnson et al. 2011; Canam et al. 2013). In Canada, it is cultivated in the cool, semi-arid prairies during late spring, summer and early fall (Marillia et al. 2014). These areas feature dry summers with extreme seasonal temperature differences (NRCAN 2008) and diurnal temperature variations. B. carinata plants do well in extreme temperatures (Canam et al. 2013) and are heat and drought tolerant (Malik 1990). B. carinata frost tolerance has been widely reported (OMAFRA 2015; Seepaul et al. 2015), although the specific temperature and duration limits have yet to be documented and/or published.

In Canada, B. carinata grows well in soil that is characterized by an organic decomposition layer, cool temperatures, and sufficientbut not necessarily perfect drainage (Cardone et al. 2003). Brassica carinata is able to tolerate low levels of salinity, however, there are severe reductions in plant growth at high levels of salinity (Canam et al. 2013). Its ability to tolerate salinity better than other Brassica species (Ashraf and McNeilly 1990) is hypothesized to be due to improved water use efficiency (Ashraf 2001).

4. Biology

4.1 Reproductive biology

Brassica carinata reproduces sexually, through both cross- and self-pollination, sets seed, and does not demonstrate potential for vegetative reproduction (Warwick et al. 2009; Mnzava and Schippers 2007). B. carinata appears to be photo-insensitive and performs well under the manipulation of seeding date in some climates (Malik 1990). However, seed set is affected by temperature. Higher yields are achieved when flowering occurs before the hottest days of summer (Gan et al. 2004).

B. carinata has been reported to cross-pollinate 30% of the time (Velasco and Fernandez-Martinez 2009; Cheung et al. 2015), due to its flower structure and delayed anthesis (Cheung et al. 2015). While sporophytic self-incompatibility exists in Brassicaceae (Howard 1942), amphidiploid Brassica species, such as B. carinata, are self-compatible (Misra 2010; Niemann et al. 2014). Self-pollination has been reported to occur from 46–88% of the time in 39 analyzed B. carinata accessions (Labana et al. 1987).

B. carinata pollen, like other Brassicaceae, is heavy, sticky, and is not dispersed well by the wind; dispersing only 10 m from the plant (Adeniji and Aloyce 2012).

The flowering of B. carinata has been described by Downey (1983). Flowering begins at the lowest bud, on the main raceme, and continues upward with 3–5 new flowers opening per day. Flowering at the base of secondary racemes is initiated approximately three days after floral initiation on the main raceme. Following pollination, the petals are shed and the pistil elongates to form a silique. Seeds are predominantly embryonic tissue, and embryos are bright yellow at maturity. The embryo consists of an inner and a larger outer cotyledon, arranged in a conduplicate fashion. The cotyledons are attached to a short hypocotyl and radicle. The position of the radicle within the seed can be observed as a distinct ridge on the surface of the seed.

4.2 Breeding and seed production

Brassica carinata was assessed to have potential as an alternative oilseed crop in western Canada if time to maturation and yields were improved (Getinet 1986; Getinet et al. 1996).

B. carinata breeding programs have pursued mainly selective breeding protocols (Alonso et al 1991; Getinet et al. 1994; Fernandez-Escobar et al. 1988; Velasco et al. 1995; Jadhav et al. 2005; Nabloussi et al. 2006; Valasco et al. 2003; Nabloussi et al. 2009; Cheng et al. 2010; Xin and Yu 2014; Márquez-Lema et al. 2006, 2008, 2009 ; Taylor et al. 2010). However, transformation and the recovery of transgenic plants are well established in Brassica species (Palmer and Keller 2002) and transgenic trait development for improved seed quality and agronomic performance has been reported for B. carinata (Taylor et al. 2010).

The major goals of recent breeding programs for B. carinata included increasing seed size, oil content and modification of seed oil composition to increase the proportion of erucic acid and nervonic acid for industrial and pharmaceutical applications (Taylor et al. 2010). Thus far, B. carinata breeding programs have yielded improved seed size (4.5–6.5 g per thousand seed) and seed oil content in excess of 48% (K. Falk, personal communication). Seed oil profile can vary. Erucic acid content traits range from zero (Alonso et al. 1991; Getinet et al. 1994) to low (< 2%; Fernandez-Escobar et al. 1988; Velasco et al. 1995) to high (~50%; Jadhav et al. 2005). In addition, varieties with high oleic acid (~85%, Nabloussi et al. 2006) and low linolenic acid (~6%; Valasco et al. 2003; Nabloussi et al. 2009) have been developed. B. carinata varieties have been developed for industrial applications such as biofuels, plastics, lubricants and specialty fatty acids. These include 5,13-docosadienoic acid and 5-eicosenoic acid (Jadhav et al. 2005), eicosapentaenoic acid (Cheng et al. 2010) and nervonic acid (Taylor 2010).

B. carinata varieties intended for seed meal products have also been produced. Examples include high protein (Xin and Yu 2014) and low glucosinolate (Getinet et al. 1997; Márquez-Lema et al. 2006, 2008) varieties for animal feed and high glucosinolate varieties for biofumigation purposes (Márquez-Lema et al. 2009).

B. carinata's ability to grow and seed western Canadian environments varies considerably; some Ethiopian accessions mature late, and yield less (Getinet et al. 1996), while others perform similarly to B. napus (Falk 1999).

B. carinata is susceptible to clubroot caused by Plasmodiophora brassicae Woronin (Peng et al. 2013). In contrast, it is resistant to blackleg (Leptosphaeria maculans (Desmaz.) Ces. et De Not.; Rimmer and van den Berg 1992) and has been used in attempts to introgress blackleg resistance into other Brassica crops (Secristan and Gerdemann 1986; Rimmer and van den Berg 1992). B. carinata is resistant to white rust (Albugo candida (Pers.) Kunze; Kole et al. 2002). Some accessions of B. carinata were found to be highly susceptible to Alternaria leaf spot (Alternaria brassicae (Berk.) Sacc.; Sharma et al. 2002), while others were partially resistant (Bansal et al. 1990).

The Canadian Seed Growers Association has developed varietal purity standards for pedigree seed production of Foundation, Registered and Certified seed (Canadian Seed Growers Association 2005). However, as of this writing, there were no seed production standards for B. carinata, and this crop is not currently subject to varietal registration in Canada.

4.3 Cultivation and use as a crop

Seeding is recommended from mid-April to early May in the Northern Plains to accommodate Brassica carinata's longer growing season relative to other Brassica oilseed crops, such as Brassica napus (Taylor et al. 2010), and to avoid flowering during the hottest days of summer (Sask Mustard 2013). However, it is planted in late October and the month of November in the south-east United States as a winter crop and information on production methods for winter (Seepaul et al. 2016).

B. carinata seed should be directly sown at a consistent depth of 1.3–2.5 cm in undisturbed stubble (summer fallow, tilled fallow or chemical fallow) when there is adequate soil moisture in the top inch of soil and temperature is 5°C or above. Sowing rates are generally adjusted to establish stand densities between 85–180 plants per square meter (Sask Mustard 2013), which is slightly higher than the recommended stand density for B. napus.

Southern Canadian prairie soils lack sufficient nitrogen for optimum production of B. carinata (Johnson et al. 2013). Maximum yields are achieved when soil nitrogen supplementation is performed at a rate of 108–135 kg N ha-1. This fertilization rate is comparable to B. napus (Johnson et al. 2011).

Based on soil analysis and nutrient requirements of B. carinata, supplemental nitrogen can be applied to fields in late fall, early spring or during seeding. Fertilizer should be applied during seeding using a mid-row or side. Separation of at least one inch between the seed and fertilizer is reported to improve B. carinata performance (Sask Mustard 2013).

Crop rotation, pre-seeding tillage, and/or chemical burn-off are important agronomic practices for reduction of weed levels. B. carinata is best grown in rotation after a cereal or pulse crop where weeds were effectively controlled (Sask Mustard 2013).

Quick seedling emergence and good stand establishment of B. carinata can prevent or minimize weed competition. B. carinata is highly branched, which can result in canopy closure earlier in the growing season relative to canola (Marillia et al. 2014). Therefore, as long as initial weed pressures are minimized, B. carinata can effectively outcompete weeds with minimal herbicide inputs (Marillia et al. 2014).

B. carinata populations have been assayed to evaluate potential natural herbicide tolerance traits. Thus far, unpublished reports have identified some level of dicamba (group 4) tolerance within available B. carinata germplasm (Johnson et al. 2014). B. carinata has tolerance to dinitroaniline (group 3) herbicides (soil applied), including trifluralin (Johnson et al. 2014).

Insect pests and diseases affecting cultivated B. carinata are discussed in Section 6.

B. carinata is harvested in the fall as one of the last crops of the season (Sask Mustard 2013). Siliques of B. carinata generally don't shatter, except in severe weather, and consequently can be directly combined when the seed has reached maturity and seed moisture is less than 9%. In the event that combining occurs late in maturation, it is recommended that the crop is swathed by cutting below the lowest seed pods. Furthermore, if seed moisture exceeds recommended levels, desiccants such as diquat can be used to accelerate the drying process (Sask Mustard 2013).

Traditionally, B. carinata is used in Africa as a leafy vegetable, providing micronutrients in the human diet (Chadha et al. 2007). Young tender leaves are eaten raw, and older leaves and stems are cooked and eaten like collards (Prakash et al. 2012). B. carinata is occasionally grown as an oilseed crop. The oil is used for cooking, illumination and in traditional medicine (Giday et al. 2010).

B. carinata may also be used as livestock fodder, or its meal may be used as a high protein animal feed when mixed with other protein sources. In Spain and Italy, seed oil is used for biofuel (Bouaid et al. 2005; Cardone et al. 2002, 2003; Gasol et al. 2007, 2009) and for bio-industrial purposes (eg. lubricant, paint, cosmetics, plastics). B. carinata has also been used in heavy metal phytoremediation (Ahmed et al. 2001; Cestone et al. 2012).

In Canada, B. carinata has been assessed as a biofuel (Blackshaw et al. 2011), but is currently grown as a cover crop to reduce soil erosion and herbicide use and to promote water conservation in orchards (Alcántara et al. 2011). The cover crop is plowed into the soil for use as a green manure soil additive or as a bio-fumigant (Núñez-Zofío et al. 2010; Porras 2011; Morales-Rodríguez et al. 2012; Guerrero-Diaz et al. 2013; Pane et al. 2013). Furthermore, allyl isothiocyanate from B. carinata seed is used as a bio-fumigant and bio-pesticide (MPT Mustard Products & Technologies Inc. 2015).

4.4 Gene flow during commercial seed and biomass production

Brassica carinata is self-compatible (Sihag 1986) and has been estimated to outcross 30% of the time in the absence of pre-pollination barriers (Labana et al. 1987; Velasco and Fernandez-Martinez, 2009). At present, little is known about intraspecific gene flow in B. carinata besides that it is possible; intraspecific breeding has been used with limited success to reduce the glucosinolate content in B. carinata (Velasco et al. 1999).

There are no documented cases of interspecific or intergeneric gene flow occurring in the field for B. carinata. There is potential for gene flow to occur given that the species appears to outcross and there are close relatives within the genus, tribe, and family that will share the environment with B. carinata, providing opportunities for hybridization and gene flow (see Section 5).

4.5 Cultivated Brassica carinata as a volunteer weed

Little information exists concerning seed dormancy and soil seed bank persistence of Brassica carinata. One previous study demonstrated that seeds exhibit some primary dormancy for a few weeks after maturation (Tokumasu et al. 1985), however it is unclear how readily seeds may enter secondary or environmentally-induced dormancy and how long they may persist in the soil. Seeds of Brassica rapa and Brassica napus can survive for several years in the soil, however their seedbanks have been observed to decline rapidly in agroecosystems (Hall 2005; USDA 2014).

4.5.1 Cultural/mechanical control

Volunteer Brassica carinata can be minimized by preventing pod shatter during harvest. Effective strategies to minimize harvest losses, such as properly setting combines and sealing any leaks will also help to minimize the number of potential B. carinata volunteers. Although some seed will be lost during harvest, it is likely that any volunteers can be easily controlled through implementation of crop rotation to a crop carrying an herbicide-tolerant trait.

4.5.2 Chemical control

Brassica carinata can be controlled with 2,4-D, or any broadleaf herbicide registered to control wild mustard or volunteer canola. No glyphosate-tolerant B. carinata varieties have been developed to date. However, there may be some level of tolerance to dicamba (Johnson et al. 2014) and dinotroaniline herbicides (soil applied) including trifluralin (Johnson et al. 2014).

4.5.3 Integrated weed management

Integrated weed management (IWM) employs a combination of cultural, mechanical and chemical approaches to managing weed populations and maximize crop yields. This may include management of stand densities and proper timing of herbicide application. At the time of writing this biology document, there have not been any IWM strategies developed for Brassica carinata volunteers.

4.5.4 Biological control

Biological control methods for Brassica carinata volunteers have not been developed.

4.6 Means of movement and dispersal

Brassica carinata reproduces and disperses by seed, but not vegetatively.

Environmental dispersal through human intervention occurs occasionally from transport trucks, railcars and improperly cleaned harvesters – similar to Brassica napus (Légère 2005).

While B. carinata dispersal through animal intervention has been proposed to occur through bird feeding, observed feeding rates have been reported to be low (Zanetti et al. 2013).

Furthermore, B. carinata seed does not possess wing or feather-like structures, so wind-mediated dispersal is expected to be negligible. Similar observations have been made with regard to water movement of B. carinata seed; only 5.5% and 0.2% of seeds float in non-turbulent and turbulent water respectively (E. Johnson, unpublished).

5. Related species of Brassica carinata

Brassica carinata is capable of interbreeding with congeners of the Brassica genus found in Canada: B. juncea, B. napus, B. nigra, B. rapa and B. oleracea (Warwick et al. 2013):

B. juncea is an introduced annual that can be found throughout Canada, except for Nunavut, Labrador, and the Yukon (Brouillet et al. 2010). It is regularly identified in cultivated wheat, oat, potato, rape fields, orchards, and as escape weeds in irrigation ditches and spring runoff areas, near grain elevators, and on road margins.

B. napus is an introduced annual that can be found in all provinces and territories, with the exception of Nunavut, and the Yukon (Brouillet et al. 2010). It is rarely observed in the proximal sub-arctic region, and is found in cultivated and abandoned wheat, barley, oat, corn, and rape fields. It is also observed as a weedy escape in dry talus, gravel slopes, river shores, railways and waste spaces.

B. nigra is also an annual and found in British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, New Brunswick, Nova Scotia, and Newfoundland (Brouillet et al. 2010). As a weed, it can be found in fields, orchards, gardens, riverbanks, roadsides, waste spaces and ballast.

B.rapa is an introduced annual found in all regions of Canada except for Nunavut and is ephemeral in the North West Territories (Brouillet et al. 2010). It can be found in open woods, meadows, ballast, on riverbanks, slopes, and beaches, alongside roadways and in waste spaces.

B. oleracea is ephemeral in Ontario and Quebec and extripaded from Saskatchewan, New Brunswick and Newfoundland (Brouillet et al. 2010). It is rare for it to escape from cultivation, and is mainly found in agricultural plots, near driftwood in British Columbia, roadsides and waste spaces.

Outside of the Brassica genus, Brassica carinata may potentially cross with species in other genera within the Brassiceae tribe (Couvreur et al. 2010). Examples of plants within the Brassiceae tribe present in Canada are Cakile edentula (Bigelow) Hook., Cakile maritima Scop., Diplotaxis muralis (L.) DC., Diplotaxis tenufolia (L.) DC., Eruca vesicaria (L.) Cav. subsp. sativa (Mill.) Thell., Erucastrum gallicum (Willd.) O.E. Schulz., Raphanus raphanistrum L., Raphanus sativus L., Rapistrum rugosum (L.) All., Sinapis alba L. and Sinapis arvensis L. Of these Brassiceae members, D. erucoides, D. tenufolia, E. gallicum, R. raphansitrum, S. alba and S. arvensis are considered weeds. The following details the distribution of these species in Canada, according to Brouillet et al. 2010 and Warwick et al. 2013

Cakile edentula is native to Ontario, Quebec, New Brunswick, Prince Edward Island, Nova Scotia, Newfoundland, and Laborador. It has also been introduced in British Columbia (Brouillet et al. 2010). It typically grows on coastal beaches and lakeshores.

Cakile maritima is an introduced annual or perennial. It is found in coastal areas of British Columbia (Brouillet et al. 2010), growing on sandy beaches and dunes among the driftwood.

Diplotaxis muralis is an introduced annual/biennial that grows on disturbed prairies, parklands, gardens, grain fields, shores, harbours, ditches and around fish houses. It grows in ballast, sand, gravel, clay, loam and can be found along roadsides, railways and waste places. It is considered weedy and is found in all areas of Canada except British Columbia, Newfoundland, Labrador, Yukon, Northwest Territories, and Nunavut (Brouillet et al. 2010).

Diplotaxis tenuifolia is an introduced perennial in British Columbia, Ontario, and Quebec, while it is ephemeral in New Brunswick and Nova Scotia (Brouillet et al. 2010). It is considered weedy and can be found in fields, river, and lakeshores, gravel pits, along roadsides, railways, waste spaces and around ports. It grows in ballast, cinders, sand, gravel, clay, and grass.

Eruca vesicaria subsp. sativa is an introduced annual found in British Columbia, Alberta, Saskatchewan, Manitoba, and Ontario. It is ephemeral in Quebec (Brouillet et al. 2010). It is found in cultivated alfalfa fields as a rare escape and seed contaminant, and occasionally along roadsides and waste spaces.

Erucastrum gallicum is an introduced, naturalized annual, winter annual in British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, New Brunswick, Prince Edward Island, Nova Scotia, Newfoundland, and the Northwest Territories (Brouillet et al. 2010). It is considered weedy and can be found in gardens, orchards, grain, mustard and sunflower fields, pastures, woods, thickets, shores, and flats. It grows in ballast, along grain elevators, roadsides and in waste spaces.

Raphanus raphanistrum is an introduced, naturalized annual, or biennial which can be found in British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, New Brunswick, Prince Edward Island, Nova Scotia, Newfoundland and Laborador (Brouillet et al. 2010). It is a weed in grain, rape, potato, cabbage, hay, clover, pea, bulb and hop fields, in gardens, orchards, woods, cliffs, outcrops, beaches, and dunes. It grows in sand, grass, gravel, clay, sandy loam, and can be found along wharves, roadsides railways and waste spaces.

Raphanus sativus is an introduced annual in British Columbia and Saskatchewan, and ephemeral in Mantioba, Ontario, Quebec, New Brunswick, Nova Scotia and Newfoundland (Brouillet et al. 2010). It is found in gardens, grain, rape and corn fields, orchards, riverbanks, flats, by wharves and roadsides. It grows on loamy, sandy soil.

Rapistrum perenne is occasionally found in Saskatchewan a weed in agricultural fields, despite its listing as extirpated (Brouillet et al. 2010). It grows in sandy loam.

Rapistrum rugosum is introduced in Ontario and Quebec (Brouillet et al. 2010). It is occasionally found on gravel shores, waterfront and ballast. It is also found near urban roadsides and waste spaces.

Sinapis alba is an introduced annual found in all areas of Canada except Northwest Territories, Nunavut, Nova Scotia and Newfoundland and Laborador (Brouillet et al. 2010). It is considered a weed and can be found in fields, farmyards, disturbed prairies, irrigated land, ballast, talus wharf, roadsides, railways and waste spaces.

Sinapis arvensis is an introduced found in all areas of Canada except for Nunavut (Brouillet et al. 2010). It is considered weedy and is found in grain, hay, rape, potato, and fruit fields, gardens orchards, clearings, river valleys and shores. It also grows on ballast, gravel, sand and can be found near grain elevators, roadsides, railways and waste spaces.

While reports attempting to cross Brassica crops with non-Brassieae have reportedly been unsuccessful (FitzJohn et al. 2007; Séguin-Swartz 2008), certain Brassicaceae are agricultural weeds and require further consideration because of their prevalence. Camelina sativa (L.) Crantz, Camelina microcarpa Andrz. Ex DC., Camelina alyssum (Mill.) Thell., Arabidopsis thaliana (L.) Heynh., Capsella bursa-pastoris (L.) Medik., Neslia paniculata (L.) Desv., Erysimum asperum (Nutt.) DC., Erysimum cheiranthoides L., Erysimum hieracifoliumL., Erysimum inconspicuum (S. Watson) MacMill. and Turritis glabra L., Alliaria petiolata(Bieb.) Cavara & Grande, Barbarea vulgaris W.T. Aiton., Berteroa incana(L.) DC., Bunias orientalisL., Conringia orientalis(L.) Dumort., Descurainia incana (Bern. Ex Fisch. & C.A. Mey.) Dom, Descurainia pinnata(Walter) Britton, Descurainia sophia (L.) Webb ex Prantl, Hesperis matronalis L., Lepidium appelianum Al-Shehbaz, Lepidium campestre(L.) W.T. Aiton, Lepidium densiflorum Schrad., Lepidium draba L., Lepidium perfoliatumL., Lepidium virginicum L., Nasturtium officinale W.T. Aiton, Rorippa austriaca (Crantz.) Besser, Rorippa sylvestris (L.) Besser, Sisymbrium altissimum L., Sisymbrium loeselii L., Sisymbrium officinale (L.) Scop. and Thlaspi arvense L. The following details the distribution of these species in Canada, according to Brouillet et al. 2010 and Warwick et al. 2013.

Camelina sativa is introduced in all areas of Canada except for Nunavut, Prince Edward Island, Newfoundland and Laborador, and has been excluded, but cultivated in Yukon. Camelina microcarpa is introduced in all areas of Canada except for the North West Territories, Nunavut and Laborador, and has been excluded, but cultivated in Prince Edward Island. Camelina alyssum is introduced in Alberta, Saskatchewan and Manitoba (Brouillet et al. 2010).

Arabidopsis thaliana is introduced in British Columbia, Ontario and Quebec, and ephemeral in Newfoundland (Brouillet et al. 2010).

Capsella bursa-pastoris is introduced in all areas of Canada (Brouillet et al. 2010).

Neslia paniculata is introduced in all areas of Canada except for Nunavut and Laborador, and has been excluded, but cultivated in Prince Edward Island (Brouillet et al. 2010).

Erysimum asperumis native to British Columbia, Alberta, Saskatchewan and Manitoba and has been introduced to Ontario and Quebec. Erysimum cheiranthoides is introduced in all areas of Canada. Erysimum hieracii folium is introduced in Ontario, Quebec, New Brunswick, Nova Scotia and is ephemeral in Saskatchewan. Erysimum inconspicuum native to all parts of Canada except New Brunswick, Price Edward Island, Newfoundland and Laborador (Brouillet et al. 2010).

Turritis glabra is native to British Columbia, Alberta, Saskatchwean, Manitoba, Ontario, Quebec, New Brunswick, Nova Scotia, and has been introduced to the Yukon and the North West Territories (Brouillet et al. 2010).

Alliaria petiolata is introduced in British Columbia, Ontario, Quebec, New Brunswick, Nova Scotia and Newfoundland (Brouillet et al. 2010).

Barbarea vulgaris is introduced across Canada except for Yukon, North West Territories and Nunavut (Brouillet et al. 2010).

Berteroa incana is introduced in all areas of Canada except for Prince Edward Island, Newfoundland and Laborador, Yukon, North West Territories and Nunavut (Brouillet et al. 2010).

Bunias orientalis is introduced in British Columbia, Quebec, ephemeral in New Brunswick and Nova Scotia (Brouillet et al. 2010).

Conringia orientalis is introduced in all areas of Canada except for Laborador, Yukon, North West Territories and Nunavut (Brouillet et al. 2010).

Descurainia incana is native to all areas of Canada except for Prince Edward Island, Nova Scotia, Newfoundland and Nunavut. In Laborador it is introduced. Descurainia pinnata is native to all areas of Canada except New Brunswick, Prince Edward Island, Newfoundland and Laborador. Descurainia sophia is introduced in all areas of Canada except for Laborador and Nunavut (Brouillet et al. 2010).

Hesperis matronalis is introduced in all areas of Canada except for Laborador, Yukon and Nunavut (Brouillet et al. 2010).

Lepidium appelianum is introduced in British Columbia, Alberta, Saskatchewan and Manitoba. Lepidium campestre is introduced in British Columbia, Alberta, Ontario, Quebec, New Brunswick, Prince Edward Island, Nova Scotia, Newfoundland and Laborador. Lepidium densiflorum is native to Alberta, Saskatchewan, Manitoba and North West Territories. It is introduced in British Columbia, Ontario, Quebec, New Brunswick, Prince Edward Island, Nova Scotia, Newfoundland and Laborador and Yukon. Lepidium draba is introduced in British Columbia, Alberta, Saskatchewan, Manitoba, Ontario and New Brunswick. It is ephemeral in Nova Scotia. Lepidium perfoliatum is introduced in British Columbia, Alberta, Saskatchewan and Manitoba. It is ephemeral to Ontario. Lepidium virginicum is native to British Columbia and Ontario. It is introduced in Quebec, New Brunswick, Prince Edward Island, Nova Scotia and North West Territories. It is ephemeral to Newfoundland (Brouillet et al. 2010).

Nasturtium officinale is introduced in all areas of Canada except Prince Edward Island, Newfoundland and Laborador, Yukon, North West Territories and Nunavut (Brouillet et al. 2010).

Rorippa austriaca is introduced in Alberta, Saskatchewan and Manitoba. Rorippa sylvestris is introduced in all areas of Canada except for Prince Edward Island, Laborador, Yukon, North West Territories and Nunavut(Brouillet et al. 2010).

Sisymbrium altissimumis introduced in all areas of Canada except for Laboraor and Nunavut. Sisymbrium loeseliiis introduced in all areas of Canada except for New Brunswick, Nova Scotia, Newfoundland and Laborador, Yukon, North West Territories and Nunavut. Sisymbrium officinale is Native to the Yukon and North West Territories, introduced to British Columbia, Alberta, Manitoba, Ontario, Quebec, New Brunswick, Prince Edward Island, Nova Scotia and Laborador (Brouillet et al. 2010).

5.1 Inter-species/genus hybridization

Brassica carinata can successfully hybridize with other Brassicaceae, such as B. napus, B. nigra, B. oleracea, B. rapa, B. tournefortii (Joshi and Choudhary 1999), B. juncea, Orychophragmus violaceus (L.) O.E. Schulz (Li et al. 1998), Raphanus sativus (Richharia 1937) and Sinapsis arvensis (Mizushima 1950; see summary in FitzJohn et al. 2007). Hybridization of B. carinata with S. arvensis has been possible through hand pollination (Cheung et al. 2015), artificial induction of polyploidy (Mizushima 1950) and in vitro techniques (eg. protoplast culture and embryo rescue; FitzJohn et al. 2007). While in vitro approaches failed to generate plants that developed to maturity and set seed, hand pollination of female B. carinata with S. arvensis pollen succeeded 6.45% of the time. When outcrossing, success rates were highest when female B. carinata accepted pollen (Cheung et al. 2015).

The results of experimental crosses between B. carinata and related plant species or genera are summarized in Table 1. The results of in vitro attempts at hybridization are summarized in Table 2.

Table 1. Reports of experimental crosses between Brassica carinata and related species.
Cross Female Cross Male Description Reference
B. carinata B. juncea Successful; some F1 seeds formed without an embryo Rahman, 1976
B. carinata B. juncea 0.02 seeds per pollination (11 F1 seeds harvested from 375 pollinations) Getinet et al. 1997
B. carinata B. juncea 0.04 seeds per pollination; F1 seeds were viable and demonstrated strong matromorphy Gosh et al. 1999
B. carinata B. juncea 6 F1 seeds from unrecorded number of crosses La Mura et al. 2010
B. carinata B. maurorum Unsuccessful; 0.02 seeds per pollination Yao et al. 2012
B. carinata B. napus Successful Fernandez-Escobar et al. 1988
B. carinata B. napus 0.08 seeds per pollination Getinet et al. 1997
B. carinata B. napus 1 F1 seed produced from unreported number of crosses La Mura et al. 2010
B. carinata B. napus 1.56 seeds per pollination Niemann et al. 2014
B. carinata B. nigra Successful Mizushima 1950
B. carinata B. nigra Successful Chang et al. 2011
B. carinata B. oleracea var. alboglabra 7.8 seeds per pollination Rahman 2001
B. carinata B. oleracea var. alboglabra 7.9 fertilized ovules per silique Rahman 2004
B. carinata B. oleracea Successful Mizushima 1950
B. carinata B. oleracea Unsuccessful Tonguç and Griffiths 2004
B. carinata B. oleracea 2 F1 seeds from 27 flowers; No viable pollen produced from F1 plants Chang et al. 2011
B. carinata B. rapa Successful Mizushima 1950
B. carinata B. rapa 6 F1 hybrids were produced from an unknown number of crosses Struss et al. 1991
B. carinata B. rapa Successful Struss et al. 1992
B. carinata B. rapa 0.23 seeds per pollination; 4 F1 plants recovered Choudhary et al. 2000
B. carinata B. rapa 3.3 seeds per pollination Rahman 2001
B. carinata B. rapa Successful Rahman 2002
B. carinata B. rapa 11.1–11.3 fertilized ovules per silique Rahman 2004
B. carinata B. rapa 0.05–0.16 plants per pollination; F1 pollen viability was 4.4–7.6% Li et al. 2005
B. carinata B. rapa Interspecific crosses with 107 B. carinata accessions yielded between 0–80+ F seeds per 100 buds Jiang et al. 2007
B. carinata B. rapa 642 F1 seeds obtained from an unspecified number of crosses Liu et al. 2009
B. carinata B. rapa 2 F1 seeds produced from undetermined number of crosses Lu Mura et al. 2010
B. carinata B. tournefortii Unsuccessful Lokanadha and Sarla 1994
B. carinata B. tournefortii 0.21 seeds per pollination; 1 F1 plant recovered, which had 2.3% pollen viability Choudhary and Joshi 2012
B. fruticulosa B. carinata Hybrids demonstrated high frequency of multivalent associations Bijral et al. 1994
B. juncea B. carinata Successful Rahman 1976
B. juncea B. carinata Successful Anand et al. 1985
B. juncea B. carinata 108 F1 seeds from undisclosed number of pollinations; 5 F1 plants recovered that were male sterile; backcross with B. carinata was unsuccessful Getinet et al. 1994
B. juncea B. carinata 48 siliques from 60 pollinated flowers Gupta 1997
B. juncea B. carinata Successful Singh et al. 1997
B. juncea B. carinata 0.75 seeds per pollination; F1 seeds were viable GhoshDastidar and Varma 1999
B. juncea B. carinata 0.7 seeds per pod; F1 pollen viability was 22.2% Chang et al. 2007
B. juncea B. carinata 3 F1 seeds from unrecorded number of crosses La Mura et al. 2010
B. juncea B. carinata Successful in only 1 or 2 B. carinata genotypes; 15–20% of F1 demonstrated male sterility Sheikh et al. 2014
B. juncea B. carinata Selfed progenies (A6 generation) of plants derived from 2 out of 9 crosses resulted in plants with B. napus genome (AACC, 2n=38); fertile hybrids resulted from crosses with natural B. napus Chatterjee et al. 2016
B. maurorum B. carinata Unsuccessful; 0.01 seeds per pollination Yao et al. 2012
B. napus B. carinata Successful Fernandez-Escobar et al. 1988
B. napus B. carinata 2.8–6.1 seeds per F1 silique; all F1 plants were sterile Chen and Haneen 1992
B. napus B. carinata 90 siliques from 110 pollinated flowers Gupta 1997
B. napus B. carinata 0–0.6 seeds per pollination; no pollen in F1 Chang et al. 2007
B. napus B. carinata 4 F1 seeds from an undetermined number of crosses La Mura et al. 2010
B. napus B. carinata Hybridization rate of 0.005% in adjacent field; 0.002% in separated field Séguin-Swartz et al. 2013
B. napus B. carinata 3.44 seeds per pollination Niemann et al. 2014
B. napus B. carinata Successful in only 1 or 2 carinata genotypes; 15–20% of F1 demonstrated male sterility Sheikh et al. 2014
B. oleracea B. carinata 7.2–8.2 fertilized ovules per silique Rahman 2004
B. rapa B. carinata 2 F1 hybrids were produced from an unknown number of crosses Struss et al. 1991
B. rapa B. carinata 1.17 seeds per pollination; Highly successful with 2 of 9 female genotypes Meng et al. 1998
B. rapa B. carinata Unsuccessful Choudhary et al. 2000
B. rapa B. carinata 5.2 seeds per pollination Rahman 2001
B. rapa B. carinata Successful Rahman 2002
B. rapa B. carinata Successful Li et al. 2005
B. rapa B. carinata 1 F1 seed from unknown number of crosses La Mura et al. 2010
B. tournefortii B. carinata Unsuccessful Choudhary and Joshi 2012
B. carinata Enarthrocarpus lyratus Unsuccessful Gundimeda et al. 1992.
B. carinata Orychophragmus violaceus 0.67–1.56 F1 hybrids per 100 pollinations Li et al. 1998
B. carinata Orychophragmus violaceus 8 F1 hybrids produced Li et al. 2003
B. carinata Raphanus sativus 1 F1 seed from undocumented number of hybridizations La Mura et al. 2010
B. carinata Sinapis alba 0.175 seeds per pollination; 26 F1 germinations Sridevi and Sarla 2005
B. carinata Sinapis arvensis Successful Mizushima 1950
B. carinata Sinapis arvensis 6 F1 seeds from undocumented number of crosses La Mura et al. 2010
B. carinata Sinapis arvensis 731 hybrids from 997 crosses; Hybridization rate of 6.4% Cheung et al. 2015
Erucastrum abyssinicum B. carinata Unsuccessful Rao et al. 1996
Orychophragmus violaceus B. carinata Unsuccessful Li et al. 1998
Raphanus sativus B. carinata Unsuccessful La Mura et al. 2010
Sinapis alba B. carinata 0.08 seeds per pollination Sridevi and Sarla 2005
Sinapis arvensis B. carinata Unsuccessful La Mura et al. 2010
B. carinata 0.01% hybridization rate Cheung et al. 2015
Table 2. Reports of in vitro hybridization between Brassica carinata and related species.
Cross Description Reference(s)
B. carinata x B. fruticulosa Successful; Embryo culture Harberd and McArthur 1980
B. carinata x B. fruticulosa Successful; Embryo culture Hybrids were male sterile Chen et al. 2012
B. carinata x B. maurorum Embryo culture; nine hybrid plantlets regenerated from 642 pollinated flowers, crossability was 1.39%, F1 hybrids had ~25% pollen viability Yao et al. 2012
B. carinata x B. napus Ovary, ovule culture; two F1 seeds from 44 pollinated flowers, F1 hybrids were male sterile Sabharwal and Doležel 1993
B. carinata x B. napus Polyethylene glycol mediated protoplast fusion; 13 plants were regenerated Klíma et al. 2009
B. carinata x B. nigra Embryo culture; successful Attia et al. 1987
B. carinata x B. oleracea Embryo culture; 6.0–7.2 embryos per pollination were rescued, survival rate of embryos was 57–96% Rahman 2004
B. carinata x B. oleracea Embryo culture; five embryos developed into plantlets from 45 pollinations, four were found to be true hybrids using RAPD analysis, all were male sterile Tonguç and Griffiths 2004
B. carinata var. botrytis x B. oleracea Asymmetric protoplast fusion; 31 hybrids found from 374 regenerated plants Scholze et al. 2010
B. carinata var. capitata x B. oleracea Symmetric protoplast fusion; five hybrids found from 21 regenerated plants Scholze et al. 2010
B. carinata x B. rapa Embryo culture; successful Quiros et al. 1985
B. carinata x B. rapa Embryo culture; successful Busso et al. 1987
B. carinata x B. rapa Embryo culture; successful, rate of natural chromosome doubling was very low Meng et al. 1998
B. carinata x B. rapa Embryo culture; 6.6–8.0 embryos per pollination were rescued, survival rate of embryos was 73–96% Rahman 2004
B. carinata x B. rapa Polyethylene glycol mediated protoplast fusion; 58 calluses, 14 shoots were regenerated, 60% of plantlets were confirmed to be hybrids by flow cytometry Beránek et al. 2007
B. carinata x B. tournefortii Embryo culture; successful with irradiated pollen Lokanadha and Sarla 1994
B. fruticulosa x B. carinata Embryo culture; successful, hybrids were male sterile Chen et al. 2012
B. juncea x B. carinata Embryo culture; successful, between 13–17 bivalents in 27 cells Harberd and McArthur 1980
B. juncea x B. carinata Ovary culture; 91 seeds formed from 226 ovaries cultured, F1 seeds poorly developed and shrivelled Sharma and Singh 1992
B. maurorum x B. carinata Ovary and ovule culture; unsuccessful, four seedlings formed from 51 cultured ovules, all F1 hybrids were pollen sterile Chrungu et al. 1999
B. maurorum x B. carinata Embryo culture; seven hybrid plantlets regenerated from 368 pollinated flowers, crossability was 1.90% Yao et al. 2012
B. napus x B. carinata Embryo culture; successful, nine bivalents observed in 50 cells Harberd and McArthur 1980
B. napus x B. carinata Ovule culture; 17.0–64.1% hybrid yield after varying days of pollination, pollen viability of sample of F1 plants ranged from 0–30%, with most hybrids between 10–20% Sacristan and Gerdemann 1986
B. oleracea x B. carinata Embryo culture; successful Attia et al. 1987
B. oleracea var. alboglabra x B. carinata Embryo culture; 12 hybrid plants obtained from 249 cross-pollinations, pollen fertility in F1 plants was 5.8% Rahman 2001
B. oleracea x B. carinata Embryo culture; 0.02–0.35 embryos per pollination were rescued; survival of the rescued embryos was 16.7% Rahman 2004
B. oleracea x B. carinata Embryo culture; unsuccessful, no hybrid plants obtained from 30 cultured pistils Tonguç and Griffiths 2004
B. rapa x B. carinata Embryo culture; successful Busso et al. 1987
B. carinata x Camelina sativa Polyethylene glycol mediated protoplast fusion; 227 calluses, three shoots were regenerated, no plants could be grown to maturity Narasimhulu et al. 1994
B. carinata x Diplotaxis assurgens Embryo culture; successful, 
3–10 bivalents seen in 55 cells
Harberd and McArthur 1980
B. carinata x Diplotaxis tenuisiliqua Embryo culture; successful,
1–10 bivalents seen in 83 cells
Harberd and McArthur 1980
B. carinata x Diplotaxis virgata Embryo culture; successful,
4–11 bivalents seen in 36 cells
Harberd and McArthur 1980
B. carinata x Enarthrocarpus lyratus Ovule culture; unsuccessful Gundimeda et al. 1992
B. carinata x Erucastrum gallicum Embryo culture; successful,
5–12 bivalents from 77 cells
Harberd and McArthur 1980
B. carinata x Raphanus sativus Embryo culture; successful,
0–4 bivalents found in 142 cells
Harberd and McArthur 1980
B. carinata x Sinapis alba Ovary and ovule culture; successful Sridevi and Sarla 1996
B. carinata x Sinapis alba Ovule culture; eight ovules cultured from 45 pollinations, no hybrid plants obtained Momotaz et al. 1998
B. carinata x Sinapis alba Ovary and ovule culture; 27 ovules cultured from 249 ovaries, only four ovules germinated, two plants formed and were confirmed as matromorphs Sridevi and Sarla 2005
Brassica carinata x Sinapis arvensis Embryo culture; successful,
0–9 bivalents found in 72 cells
Harberd and McArthur 1980
Brassica carinata x Sinapis arvensis Ovule culture; 269 ovules cultured from 96 pollinations, 29 hybrid plants formed, hybrids had no pollen fertility Momotaz et al. 1998
Brassica carinata x Sinapis turgida Ovule culture; 166 ovules cultured from 41 pollinations, eight hybrid plants formed, hybrids had no pollen fertility Momotaz et al. 1998
Enarthrocarpus lyratus x Brassica carinata Ovule culture; one hybrid obtained from 54 pollinated ovaries; F1 hybrid showed 2% pollen fertility Gundimeda et al. 1992
Erucastrum abyssinicum x Brassica carinata Ovary culture; successful, F1 hybrids pollen sterile Rao et al. 1996
Sinapis alba x Brassica carinata Ovary and ovule culture; successful Sridevi and Sarla 1996
Sinapis alba x Brassica carinata Ovule culture; six ovules cultured from 45 pollinations, no ovule development and no hybrid plants obtained Momotaz et al. 1998
Sinapis alba x Brassica carinata Ovary and ovule culture; 11 ovules cultured from 153 ovaries, only one F1 hybrid formed Sridevi and Sarla 2005
Sinapis arvensis x Brassica carinata Ovule culture; 32 ovules cultured from 33 pollinations, no ovule development and no hybrid plants obtained Momotaz et al. 1998
Sinapis turgida x Brassica carinata Ovule culture; 11 ovules cultured from 21 pollinations, no ovule development and no hybrid plants obtained Momotaz et al. 1998

5.2 Potential for introgression of genetic information from Brassica carinata into relatives

Brassica carinata is the least studied brassicacea crop in terms of interspecific hybridization (FitzJohn et al. 2007; Cheung et al. 2015). Attempts to hybridize B. carinata with B. maurorum (Chrungu et al 1999), B. tournefortii (Joshi and Choudhary 1999), E. lyratus (Gundimeda et al. 1992), E. abyssinicum (Rao et al. 1996), O. violaceus (Li et al. 1998), R. sativus (Gupta 1997) and S. alba (Sridevi and Sarla 1996) have failed when B. carinata is the pollen donor (reviewed FitzJohn et al 2007).

There is potential for crossing and therefore gene introgression from B. carinata into some of its cogeners in Canada, however. The creation of hybrids of B. carinata with major, Canadian brassica crops (eg. B. napus, B. juncea, B. rapa, B. oleracea) have been documented in the literature. Attempts to hybridize B. carinata (♂) with B. napus (♀) have been reported seven times in the literature and have always been successful (Nagaharu 1935; Roy 1980; Wahiduzzaman 1987; Fernandez-Escobar et al. 1988; Chen and Heneen 1992; Rashid et al. 1994; Getinet et al. 1997; Pu et al. 2005; Séguin-Swartz et al. 2013; reviewed FitzJohn et al 2007). The hybridization frequency was low, with F1 hybrids being sterile (Getinet et al. 1997). Similarly, attempts to hybridize B. carinata (♂) with B. juncea (♀) have been reported eleven times and have always been successful (Nagaharu 1935; Rahman 1976; Rahman 1978; Anand et al 1985; Katiyar and Gupta 1987; Subudhi and Raut 1994; Katiyar and Chamola 1995; reviewed FitzJohn et al 2007). Attempts to hybridize B. carinata (♂) with B. rapa (♀) appears five times, succeeding 80% of the time (Howard 1942; Struss et al 1991; Meng et al 1998; Choudhary et al 2000; Rahman 2001; reviewed FitzJohn et al 2007). Hybridizations between B. carinata (♂) with B. oleracea (♀) have been reported four times, succeeding half of the time (Morinaga 1933; Nagaharu 1935; Barcikowska et al. 1983; Rahman 2001; reviewed FitzJohn et al 2007).

B. carinata (♂) hybridization with S. arvensis (♀), a self-incompatible wild mustard with persistent seed banks (Warwick et al. 2000), occurs at a rate of 0.01% in the absence of pre-pollination barriers (Cheung et al. 2015). In 1109 crosses a single hybrid was produced and it generated less than 1% of the B. carinata (♂) parent's pollen.

5.3 Summary of the ecology of relatives of Brassica carinata

Brassica species can be found as weeds across Canada, with canola varieties - B. napus and B. rapa - mainly volunteering throughout Alberta, Saskatchewan and Manitoba (Leeson et al. 2005; Gulden et al. 2008; Warwick et al. 2013). Between 2003 and 2014, canola volunteers were promoted from 16th to 4th most prevalent weed in western Canada (Beckie 2015).

Of non-canola varieties of Brassica, B. nigra can be found in old fields, along roadsides, and in waste spaces as weeds and B. oleracea is found as a rare escape from cultivated plots in British Columbia, Alberta, Ontario, and Quebec. B. juncea has not become a problematic or abundant weed despite its presence across Canada (Leeson et al. 2005).

B. napus volunteer populations with herbicide resistance, acquired by means of hybridization with cultivars containing resistant traits has been documented (Hall et al. 2000), including multiple resistances to glyphosate, glufosinate, bromoxynil and imidazolinone (Hall et al. 2000; Beckie et al. 2003). Furthermore, triazine resistance in feral B. rapa has also been reported, and transferred to cultivated B. rapa and B. napus (Beversdorf et al. 1980). Herbicide resistance in Brassica weed populations makes their control in agricultural settings challenging. This is particularly true for weeds acquiring multiple resistances (Hall et al. 2000; Beckie et al. 2003).

Outside Brassicaceae, S. arvensis is considered a weed and grows in a wide variety of habitats including cultivated fields, alongside grain elevators, roadsides, railways and in waste places. It is a primary colonizer of disturbed areas. It is readily killed by frost and generally grows in habitats with high light intensity (Warwick et al. 2000). It is controlled through deploying herbicides (Warwick et al. 2000; Leeson et al. 2005. In annual weed surveys, S. arvensis ranked 24th out of 148 agricultural weeds.

Co-occurence of canola (B. napus and/or B. rapa) with S. arvensis is reported to happen at a frequency of 12.6% in prairie provinces (Leeson et al. 2005). As the cultivation range of B. carinata is likely to overlap with that of B. juncea and B. napus, it is expected that B. carinata will grow in close proximity to S. arvensis. While seed of S. arvensis is classed as a primary noxious weed in Canada at the publication of this biology document (Government of Canada 2005), it has been proposed for recategorization as a secondary noxious weed in the Weed Seed Order update (Canada Gazette 2016).

6. Potential Interaction of Brassica carinata with Other Life Forms

Little is known of Brassica carinata and its environmental interactions in its center of origin (i.e. Ethiopia). Table 3 lists disease interactions from outside B. carinata's center of origin, focusing on Canada.

B. carinata is reported to be resistant to many diseases affecting crucifers in Canada; such as blackleg (Leptosphaeria maculans; Gugel et al. 1990), Verticillium longisporum (Zeise and Buchmuller 1997) white rust (Albugo candida; Naresh 2014), and alternaria (Alternaria brassicae; Chavan and Kamble 2014) and sclerotinia stem rot (Sclerotinia sclerotiorum (Lib.) Massee) and aster yellows (Candidatus phytoplasma asteris) (Sask Mustard 2013).

However, B. carinata is susceptible to clubroot (Plasmodiophora brassicae), a soil-borne fungus-like pathogen (Kingdom: Chromista; Infrakingdom: Rhizaria). Clubroot resistant cultivars of B. carinata may be generated through hybridization B. rapa (Peng et al. 2013). Until clubroot resistant B. carinata cultivars become available, producers must carefully clean equipment to limit the movement of clubroot-infested soil.

Like other Brassica species, B. carinata has been investigated extensively for its ability to reduce soil-borne plant pathogens. It contains glucosinolates that produce bio-fumigants such as isothiocyanate when they breakdown. Allyl isothiocyanate harvested from B. carinata seed has been formulated as biofumigants, biopesticides and bionematicides in Canada (MPT Mustard Products & Technologies Inc. 2015).

The range of insects that B. carinata interacts with in Canada is reported to be similar to other brassicaceous oilseed crops (summarized in Table 3), though B. carinata is reported to be less suspeptable than canola to insect herbivores (Palaniswamy et al. 1997; Ulmer et al. 2001, 2002; Cárcamo et al. 2007). According to a producer survey, the five most serious economic insect pests of canola in western Canada are flea beetles (Phyllotreta spp.), bertha armyworm (Mamestra configurata Walker), diamondback moth (Plutella xylostella L.), Lygus spp. plant bugs, and aphids (suborder Sternorrhyncha, formerly Homoptera; Koch Paul Associates 2000).

The crucifer flea beetle (Phyllotreta cruciferae Goeze) actively feeds on B. carinata, though certain accessions have feeding levels that are reportedly less than on B. juncea, B. napus, or B. rapa (Palaniswamy et al. 1992, Bodnaryk 1992, Palaniswamy et al. 1997). Studies investigating the underlying reasons for these differences point to the absence of stimulatory chemical feeding cues in B. carinata rather than the presence of repellents (Palaniswamy et al. 1997), or an augmented waxy leaf surface in B. carinata (Bodnaryk 1992).

Polyphagous bertha armyworms (Mamestra configurata Walker) feed and oviposit on B. carinata at rates lower than on B. napus and B. juncea (Ulmer et al. 2001, 2002).Likewise, diamondback moths (Plutella xylostella L.) feed on B. carinata at rates lower than on some other Brassica species (Andrahennadi and Gillott 1998, Sarfraz et al. 2005, Sarfraz et al. 2007). In contrast, the generalist tarnished plant bug Lygus lineolaris (Palisot de Beauvois) readily feeds and oviposits on B. carinata (Gerber 1996, 1997).

Crucifer-specialist aphids (e.g. Brevicoryne brassicae L, Lipaphis erysimi Kalt.) and generalist aphid feeders (e.g. Myzus persicae Sulzer) are occasionally found on oilseed crops in Canada, but rarely cause economic losses (Canola Council of Canada 2014a). B. carinata was observed to be highly susceptible to infestation by Brevicoryne brassicae in greenhouse trials in the United States (Jarvis 1982).

Cabbage root maggot (Delia radicum L.) is the most common crucifer-feeding root maggot (Delia spp.) in Canada (Griffiths 1991, Soroka and Dosdall 2011). B. carinata was found to be less resistant to D. radicum than Sinapis alba (Jyoti et al. 2001). Delia spp. flies were found to infest and damage B. carinata at intermediary ratings when compared to of the nine tested crucifer species (Soroka et al. 2014). There are no effective chemical based management options for cabbage root maggot (Sask Mustard 2013).

Leafhopper (Macrosteles quadrilineatus Forbes) has also been observed on B. carinata plants in Saskatchewan, and could potentially serve as a vector for the aster yellows phytoplasma (Olivier, unpublished data).

Swede midge (Contarinia nasturtii Kieffer) is not a well-documented pest of B. carinata, however,recent experiments in Saskatchewan found Swede midge feeding on B. carinata (Andreassen and Soroka, unpublished data).

Cabbage seedpod weevil (Ceutorhynchus obstrictus Marsham) is an invasive alien species that has become a serious pest of canola in Canada (Dosdall and Cárcamo 2011). B. carinata has intermediate susceptibility when compared to cultivars of B. rapa, B. napus and S. alba (Cárcamo et al. 2007). Several other native and non-native Ceutorhynchus species are specialist crucifer feeders (Dosdall et al. 2007, Mason et al. 2014), yet their ability to feed on B. carinata has not been documented.

Other Lepidoptera that occasionally feed on B. napus in Canada includesimported cabbageworm (Peiris rapae L.) and cabbage looper (Trichoplusia ni Hüner). No specific reports of these two pests of B. carinata are available, although they are both listed as potential pests of B. carinata in Florida, USA (Bliss et al. 2015). Congeners P. brassicoidesi Guerin-Meneville and T. orichalcea (Fab.) are listed as insect pests of potential importance of oilseed Brassica in Ethiopia (Gebre-Medhin and Mulatu 1992). In a study of the biology of the related butterfly P. brassicae, which is not present in North America, Chahil and Kular (2013) found that B. carinata was the most susceptible among B. napus, B. juncea, B. rapa and B. carinata lines to feeding by larvae of the butterfly.

Brassicogethes (= Meligethes) pollen beetles, including B. aeneus (Fab.) (= M. aeneus (Fab.)) and B. viridescens (Fab.) (= M. viridescens (Fab.)), are widespread and serious pests of oilseed rape (Brassica napus and B. rapa) in Europe. Brassicogethes viridescens has recently become established in Atlantic Canada, where it is flourishing and is now found as far west as Quebec (Mason et al. 2003). Although degrees of resistance and/or susceptibility to the pollen beetle B. aeneus have been found among B. napus, B. rapa, B. juncea, and S. alba entries (Kaasik et al. 2014), the suitability of B. carinata as a host of Brassicogethes beetles is unknown.

The generic term cutworm refers to larvae of Hymenoptera in the Family Noctuidae, which sever the stems of their host at or above soil level. The most economically important species of cutworm on canola in western Canada include redbacked (Euxoa ochrogaster Guenée), pale western (Agrotis orthogonia Morrison), darksided (Euxoa messoria Harris), army (Euxoa auxiliaris Grote) and dingy cutworms (Feltia jagulifera Guenée; Canola Council of Canada 2014b). These and other, less common, cutworm species would likely feed on B. carinata as well as they do on canola, for most cutworm species are polyphagous feeders.

Surveys of beneficial insects on B. carinata in Canada have not been conducted, but it is likely that the insect species that feed on or parasitize insect pests of canola (B. napus and B. rapa)would behave the same on these pests should they occur in B. carinata. Beneficial insects commonly found in canola fields in Canada include predators such as larval and adult ladybird beetles (Family Coccinelidae), lacewing larvae in the Family Chrysopidae, larvae of hover flies (Syrphidae), nymphal and adult minute pirate bugs (Anthocoridae), ground beetles (Carabidae), beneficial thrips (Thysanoptera), rove beetles (Staphylinidae) and many species of beneficial mites and spiders (Canola Council of Canada 2014c). These predators are generalist feeders, consuming any insect that they can capture.

Parasitoids, primarily Hymenopteran wasps and a few fly (Dipteran) species, are generally host-specific. Principal parasitoids of canola insect pests in Canada include Perlitus brevipetiolatus Thomson (=Microctonus vittatae Muesback) (Braconidae)parasitizing the flea beetles Phyllotreta cruciferae and P. striolata (Soroka 2013), and Banchus flavescens Cresson (Ichneumonidae) and Athrycia cinerea (Coquillett) (Tachinidae) parasitizing bertha armyworm populations (Turnock 1984). The main parasitoids of P. xylostella in canola in western Canada are Microplitis plutellae (Muesbeck) (Braconidae), Diadegma insulare (Cresson) and Diadromus subtilicornis (Gravenhorst) (both Ichneumonidae; Braun et al. 2002; Bahar et al. 2013). Parasitoids of Lygus spp. in Canada include several species in the braconid genera Peristenus and Leophron (Broadbent et al. 2013). Diaretiella rapae (Mcintosh) (Braconidae) is a generalist parasitoid of a wide group of aphids, including Brassica feeders Brevicoryne brassicae, Lipahis erysimi, and Myzus persicae (CABI 2015); it has been reported as parasitizing B. brassicae in oilseed Brassica species in Ethiopia (Gebre-Medhin and Mulatu 1992). D. rapae is also listed as a parasitoid of diamondback moth, Plutella xylostella (CABI 2015). The principal parasitoids of the Delia spp. root maggot complex in Canada is the larval-pupal parasitoid Trybliographa rapae (Westwood) (Hymenoptera; Figitidae) and pupal parasitoids Aleochara bilineata (Gyllenhall) and A. verna (Say) (Coleoptera: Staphylinidae; Hemachandra et al. 2007). Biological control agents of other canola pests may be found in B. carinata crops if the corresponding pest is also present.

Although many oilseed Brassica species such as B.napus and B. carinata are self-compatible and self-pollinated (Eisikowitch 1981), pollen transfer by invertebrate vectors can increase Brassica seed yield (Wescott and Nelson 2001; Steffan-Dewenter 2003; Sabbahi et al. 2005). Mishra and Kaushik (1992) reported that seed yield was higher in honeybee (Apis mellifera L.) associated open-pollinated B. carinata and five other Brassica lines than in self–pollinated lines. Canola flowers secrete large amounts of nectar and are very attractive to bees, including wild species of Bombus (Turnock et al. 2006), Osmia (Steffan-Dewenter 2003), Andrena, Halictus, and others, as well as leafcutting bees Megachile rotundata (Fabricius; Soroka et al. 2001). These and other pollen vectors of canola such as hover flies (Diptera: Syrphidae; Jauker and Wolters 2008) might be equally effective pollinators of B. carinata.

There are several potential animal pests that may be of concern for B. carinata production, including moose (Alces alces), white-tailed deer (Odocoileus virginianus Zimmerman), mule deer (O. hemionus Rafinesque), Richardson's ground squirrel (Spermophilus richardsonii Sabine), birds (R. Bennett, pers. comm. 2014) and cattle (D. Males, pers. comm. 2014; J. Marois, pers. comm. 2014). In related Brassiceae species, such as Camelina sativa, animal pests include white-tailed deer (O. virginianus), pronghorn antelope (Antilocapra americana Ord), slugs and birds.

For a list of species associated with or potentially associated with B. carinata please refer to Table 4.

Table 4. Examples of potential interactions of cultivated Brassica carinata with other life forms present in Canada during its life cycle.

Fungi
Life Form Interaction with B. carinata* (pathogen; beneficial organism; consumer; gene transfer) Presence in Canada Reference(s)
Alternaria brassicae (Berk.) Sacc. pathogen present Sharma et al. 2007; Chavan and Kamble 2014
Erysiphe cruciferarum Opiz ex. Junell (powdery mildew) pathogen present, widespread Naresh 2014
Erysiphe polygoni D.C. (powdery mildew) pathogen present, widespread Tonguç and Griffiths 2004
Hyaloperonospora parasitica (Pers.:Fr) Fr. pathogen present Naresh 2014
Leptosphaeria maculans (Desmaz.) Ces. & De Not. (blackleg) pathogen present Plieske et al. 1998; Fredua-Agyeman et al. 2014
Pseudocercosporella capsellae (Ellis & Everh.) Deighton (grey stem and white leaf spot); teleomorph: Mycospaerella capsellae A.J. Inman & Sivan. pathogen present, widespread Gunasinghe et al. 2014
Sclerotinia sclerotiorum (Lib.) de Bary (sclerotinia stem rot) pathogen present, widespread Barbetti et al. 2014
Chromista
Life Form Interaction with B. carinata* (pathogen; beneficial organism; consumer; gene transfer) Presence in Canada Reference(s)
Albugo candida (Pers.) Kuntze (white rust) pathogen present, widespread Naresh 2014
Plasmodiophora brassicae Woronin (clubroot) pathogen present Peng et al. 2013
Bacteria
Life Form Interaction with B. carinata* (pathogen; beneficial organism; consumer; gene transfer) Presence in Canada Reference(s)
Xanthomonas campestris pv. campestris (Pammel) Dowson (black rot) pathogen present, widespread Vicente and Holub 2013
Phytoplasma
Life Form Interaction with B. carinata* (pathogen; beneficial organism; consumer; gene transfer) Presence in Canada Reference(s)
Candidatus Phytoplasma asteris (aster yellows) pathogen present, widespread Azadvar et al. 2011
Viruses
Life Form Interaction with B. carinata* (pathogen; beneficial organism; consumer; gene transfer) Presence in Canada Reference(s)
Turnip yellow mosaic virus (TYMV) pathogen present, Ontario Babu et al. 2013
Insects
Life Form Interaction with B. carinata* (pathogen; beneficial organism; consumer; gene transfer) Presence in Canada Reference(s)
Bombus spp. (bumblebees) symbiont or beneficial organism present Turnock et al. 2006
Brassicogethes (=Meligethes) viridescens Fab. (pollen beetle) consumer Recently introduced into eastern Canada Mason et al. 2003
Brevicoryne brassicae L. consumer present, widespread Cole 1997; Razaq et al. 2011; Maremela et al. 2013
Ceutorhynchus americanus Buch. consumer present, widespread Mason et al. 2014
Ceutorhynchus erysimi F. consumer present, widespread Borg 1952a; Majka et al. 2007; Mason et al. 2014
Ceutorhynchus neglectus consumer present – weed biocontrol agent Mason et al. 2014
Ceutorhynchus obstrictus Marsham (cabbage seed pod weevil) consumer present, widespread Cárcamo et al. 2007; Ulmer and Dosdall 2006
Ceutorhynchus pallidactylus Marsh. = C. quadridens Panz. (cabbage stem weevil) consumer Recently introduced into North America Borg 1952b; Majka et al. 2007
Ceutorhyncus rapae Gylh. (cabbage curculio) consumer present, widespread Borg 1952a; Mason et al. 2014
Ceutorhynchus subpubescens consumer widespread Dosdall et al. 2007
Contarinia nasturtii Kieffer consumer present Hallett and Heal 2001; Hallett 2007
Delia spp. (root maggots) consumer present, widespread Soroka and Dosdall 2011; Soroka et al. 2014; van Dam et al. 2012
Entomoscelis americana Brown consumer present Canada Department of Agriculture 1951
Grasshoppers consumer present, widespread Gavloski 2003
Leptinotarsa decemlineata Say (Colorado potato beetle) B. carinata bio-oil a source of insecticide present, widespread Suqi et al. 2014
Lipaphis erysimi Kalt. consumer present Kular and Kumar 2011, Razaq et al. 2011, Kumar et al. 2011, Singh and Lal 2012
Liriomyza leafminers, incl. Liriomyza trifolii Burgessand Liriomyza brassicae Riley consumer present Beirne 1971; OMAFRA 2009
Lygus spp. consumer present, widespread Kelton 1980
Macrosteles quadrilineatus Forbes (aster leafhopper) consumer (vector of aster yellows) present, widespread Maw et al. 2000; Hamilton and Whitcomb 2010
Macrosteles fascifrons Stål. (aster leafhopper) consumer (vector of aster yellows) present, widespread Westdal et al. 1960
Mamestra configurata Walker (bertha army worm) consumer present, widespread in western Canada Ulmer et al. 2001, 2002
Myzus persicae Sulzer (green peach aphid) consumer present, widespread Beirne 1972; Canola Council of Canada 2014a
Phyllotreta spp., principally P. cruciferae Goeze and P. striolata Fab. (flea beetle) consumer present, widespread Palaniswamy et al. 1992, Bodnaryk 1992, Palaniswamy et al. 1997, Soroka and Grenkow 2013
Pieris brassicae L. consumer India Chahil and Kular 2013
Pieris rapae L. consumer present Beirne 1971
Plutella xylostella L. (diamondback moth) consumer present; does not overwinter in Canada, migratory Harcourt 1957; Harcourt 1963
Psylliodes punctulata Melsh. consumer present Burgess 1977
Animals
Life Form Interaction with B. carinata* (pathogen; beneficial organism; consumer; gene transfer) Presence in Canada Reference(s)
Bos primigenious f. taurus (cattle) consumer present D. Males, pers. comm. 2014; J. Marois, pers. comm. 2014
Odocoileus virginianus Zimmerman (white-tailed deer); Odocoileus hemionus Rafinesque (mule deer) consumer present R. Bennett, pers. comm. 2014
Spermophilus richardsonii (Richardson's ground squirrel, gopher) consumer present R. Bennett, pers. comm. 2014
Birds consumer present R. Bennett, pers. comm. 2014
Plants
Life Form Interaction with B. carinata* (pathogen; beneficial organism; consumer; gene transfer) Presence in Canada Reference(s)
Brassica napus gene transfer present Getinet et al. 1997; Séguin-Swartz et al. 2013; Niemann et al. 2014
Brassica nigra gene transfer present Chang et al. 2011
Brassica oleracea gene transfer present Meng et al. 1998
Brassica rapa gene transfer present Jiang et al. 2007
Sinapis arvensis gene transfer present Cheung et al. 2015

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