Crop Adaptation to Climate Change (Yadav/Crop Adaptation to Climate Change) || Breeding Oilseed Brassica for Climate Change

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Chapter 22

Breeding Oilseed Brassica for ClimateChangePhillip A. Salisbury and Martin J. Barbetti

Introduction

Oilseed Brassica species are among the mostimportant oilseed species worldwide. There arethree different Brassica species widely usedfor oilseed production, namely Brassica na-pus, Brassica rapa, and Brassica juncea. Afourth species, Brassica carinata is also usedfor oilseed production in Ethiopia, while sev-eral other species are utilized commercially to asmaller degree. These Brassica species are partof the triangle of U (UN 1935), which showsthe relationship between the six major cultivatedBrassica species (Fig. 22.1). It comprises threediploid species, B. rapa (genome AA, 2n = 20),Brassica nigra (BB, 2n = 16), and Brassica ol-eracea (CC, 2n = 18), plus three amphidiploidspecies, B. napus (AACC, 2n = 38), B. juncea(AABB, 2n = 36), and B. carinata (BBCC,2n = 34). The amphidiploid species originatedthrough interspecific hybridization between twoof the three diploid species. Worldwide, around45 million tons of oilseed Brassica are pro-duced annually. The main production areas in-clude Europe, North America, China, India, andAustralia.

Oilseed Brassica crops are grown primarilyfor oil, with oil content typically around 35–48%

of the seed weight. Once the oil is extracted, theremainder is a high-quality protein feed mealused primarily in pig and poultry rations in de-veloped countries. The oil is the most valuablecomponent of the seed, typically accounting for65–80% of the seed value, with the meal account-ing for the balance. The properties and uses ofBrassica oils are determined primarily by theirfatty acid composition. The term canola qualityrefers specifically to oilseed Brassica cultivarswith less then 2% erucic acid (C22 : 1) in the oiland less than 12 μmoles of total glucosinolatesper gram of seed at 8.5% moisture (equivalent toapproximately 20 μmoles of total glucosinolatein seed meal). The level of oleic acid in the oil ofcanola cultivars is around 60%. Any considera-tion of the impact of climate change on oilseedBrassica production must therefore consider notonly yield but also oil content, oil quality (i.e.,fatty acid composition), meal protein content,and glucosinolate content. In addition, the im-pact of climate change on key diseases, insects,and weeds must also be considered.

Global atmospheric concentrations of CO2,CH4, and N2O have increased markedly as aresult of human activities since 1750. The pri-mary source is the increase of CO2 as a resultof fossil fuel use (Torrey 2007). With the advent

Crop Adaptation to Climate Change, First Edition. Edited by Shyam S. Yadav, Robert J. Redden, Jerry L. Hatfield,Hermann Lotze-Campen and Anthony E. Hall.c© 2011 John Wiley & Sons, Ltd. Published 2011 by John Wiley & Sons, Ltd.

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BREEDING OILSEED BRASSICA FOR CLIMATE CHANGE 449

n = 10AA

B. rapa

n = 17BBCC

B. carinata

n = 19AACC

B. napus

n = 9CC

B. oleracea

n = 18AABB

B. juncea

n = 8BB

B. nigra

Fig. 22.1. Triangle of U of Brassica species (redrawn fromUN 1935).

of climate change, production of oilseed Bras-sica crops is expected to face higher growingtemperatures and CO2 levels and more variableclimates, including increased water stress andsalinity in some regions (Torrey 2007; Watkins2008). The impact of these changes may varysomewhat in the different growing regions, fromthe Australian Mediterranean climate, the coolwinter regions of Europe to the hot regionsof India. Most oilseed crops are rainfed crops,making the expected high degree of variabil-ity of rainfall a challenge in every region inthe world.

Any consideration of the impact of climatechange on oilseed Brassica production must con-sider not only yield but also oil content, oil qual-ity, and meal quality. An understanding of theimpact of increased temperatures, moisturestress, and increased CO2 levels on oil and mealquality is crucial for the development of strategicbreeding programs for climate change. Duringthe period of seed growth, a number of qual-ity changes occur in the seed to enable it toreach its final quality at maturity (Mendham andSalisbury 1995).

Quality changes during seeddevelopment

The length of the canola growing season variesfrom around 120 days in Canada to around330 days in Europe. In each environment, the du-ration of the seed development phase is largelydetermined by temperature, with seed taking ap-proximately 50–60 days to reach maturity from aCanadian spring sowing and 90–110 days froma European winter sowing. During this period ofseed development, a number of quality changesoccur in the seed to enable it to reach its fi-nal quality at maturity (Mendham and Salis-bury 1995). The general pattern of developmentwithin each oil and meal trait is similar, althoughthe time frame varies.

Oil content

During seed development, the rate of oil synthe-sis and deposition in the seed follows a sigmoidcurve. In spring grown B. napus, solid dropletsof storage oil are first seen in developing seedabout 18 days after pollination. They essentiallyincrease in size and number between about 20and 35 days after pollination, when the oil con-tent reaches a virtual plateau, with relatively lit-tle change in oil content registered during thelater stages up to maturity at 55 days after pol-lination (Fowler and Downey 1970; Rakow andMcGregor 1975; Pomeroy and Sparace 1992).At maturity, about 80% of the seed oil is con-centrated in liquid droplets in the cells of thecotyledons of the embryo (Stringam et al. 1974).

Oil quality

Fatty acids occur in seeds mainly as triacyl-glycerols. In a low erucic acid B. rapa, thefatty acid composition in the developing seedchanges drastically during the period 15–36 daysafter pollination (Romero 1991). The percent-age of oleic acid (C18:1) increases rapidly, withpalmitic (C16:0), stearic (C18:0), and linolenic(C18:3) acids continually reducing. Linoleic acid



450 CROP ADAPTATION TO CLIMATE CHANGE

(C18:2) decreases initially, then stabilizes at itsmature level. During the last stage of matura-tion, from 36 days after pollination onward, nofurther significant changes to fatty acid composi-tion occur. Pomeroy and Sparace (1992) reporteda similar pattern of fatty acid biosynthesis inB. napus.

Protein content

Synthesis of storage proteins takes place in thedeveloping seed (Finlayson and Christ 1971).The two major storage proteins in B. napus, 12Sand 1.7S (Crouch and Sussex 1981), tend to ap-pear in the second half of the seed growth period.Over 70% of the protein in the mature seed isfound in the cotyledons (Kind et al 1977), withthe rest in the embryo and the seed coat.

Glucosinolate content

Trace amounts of seed glucosinolates are ob-served in spring canola eight days after pol-lination. Seed glucosinolate content increasesrapidly between about 15 and 30–35 days afterpollination, with increase continuing at a muchslower rate after this time (Kondra and Downey1970; De March et al. 1989). Merrien et al.(1991) reported that glucosinolate accumulationin winter canola virtually stopped 30 days before

physiological maturity in the low glucosinolatecultivar Samourai. They also reported changes inthe relative concentrations of the individual glu-cosinolates during seed development. The em-bryo contained most of the glucosinolates in themature seed, while the seed coat had only a rel-atively small amount of it (Josefsson 1970).

Effect of climate change on seeddevelopment

The stresses associated with climate change, par-ticularly increased temperatures, drought stress,and increased CO2 levels, will have a signifi-cant effect on the yield and quality of oilseedBrassica crops. While the effects of high tem-peratures and drought stress are reasonably welldefined, effects of increased CO2 levels are lessunderstood.

Increased temperatures

The effects of increased temperature on yieldand quality of oilseed Brassica are predom-inantly negative. Selected examples are pre-sented in Table 22.1. One exception is the yieldin cooler countries like Sweden and Denmark,where higher temperatures could be beneficialfor vegetative growth and yield. In general, en-vironmental factors that cause a change in oil

Table 22.1. Effects of increased temperature on yield and quality of oilseed Brassica.

Trait Effect References

Biomass Decreased Qaderi et al. (2006)Maturity More rapid Aksouh et al. (2001)Yield-warm countries Decreased Aksouh et al. (2001)Yield-cool countries Potential increase Peltonen-Sainio et al. (2009)Oil content Decreased Canvin (1965), Mailer and Cornish (1987), Hocking

and Strapper (1993), Walton et al. (1999), andAksouh et al. (2001)

Oil quality Increased oleic acid content(when adequate moisture)

Canvin (1965) and Pritchard et al. (2000)

Protein content Increased Mailer and Cornish (1987) and Aksouh et al. (2001)Glucosinolate content Increased Salisbury et al. (1987) and Aksouh et al. (2001)Chlorophyll Decreased Ward et al. (1992)




content tend to cause a corresponding oppositechange in seed protein content.

Each 1◦C rise in average temperature duringseed fill reduces oil content by 1.2–1.5% (Canvin1965, Hocking and Stapper 1993). In Australia,the payment for canola is dependent on the oilcontent of the crop. Oil contents below 42% incura price penalty, while oil contents above 42% re-ceive a bonus. The increased temperatures asso-ciated with climate change could, therefore, po-tentially significantly reduce returns to farmers,both from reduced yields and lower oil contents.In addition, increases in glucosinolate content as-sociated with higher temperatures could furtherreduce crop quality.

Drought stress

As with high temperatures, the effects of in-creased temperature on yield and quality ofoilseed Brassica are predominantly negative.Some examples are presented in Table 22.2.

A comparison of Table 22.1 and Table 22.2illustrates the similarity of the effects of in-creased temperatures and moisture stress. Thetwo factors often occur together, further reduc-ing the quality of the crop.

Increased CO2 levels

In contrast to the good understanding of hightemperature and moisture stress on oilseed

Brassica plant development, Franzaring et al.(2008) indicate that little is known about the yieldand quality responses of oilseed Brassica cropsto increased levels of atmospheric CO2. This canbe seen in the limited data available in Table 22.3.It is difficult to directly compare some studies,as they vary from short term to full season ap-plication of treatments, from glasshouse to fieldtrials and the treatments applied are at variablelevels. In general, studies of increased CO2 haveused CO2 levels ≥700 μmol/mol, compared withcurrent levels of around 380 μmol/mol.

Further work is required to understand the in-teractions between different climate change fac-tors (Qaderi et al. 2006). Where drought stressand high temperature stress occur together, theoverall effect is likely to be much more severethan the sum of the individual effects. There issome indication in the literature that increasedCO2 levels may partially reverse the effects ofhigh temperature and drought. However, aftercarrying out a season-long field study under con-ditions of elevated CO2, Franzaring et al. (2008)suggested that the beneficial effect of CO2 onplant growth tended to diminish after flowering,and may not translate into a significant increasein oil content or oil yield. In addition to under-standing the impact of increased CO2 on yield,a thorough understanding of the impact of theseinteractions on all key quality components is alsorequired.

Table 22.2. Effects of increased moisture stress on yield and quality of oilseed Brassica.


Biomass Decreased Qaderi et al. (2006)Maturity Faster Pritchard et al. (2000)Yield Decreased Jensen et al. (1996), Champolivier and Merrien (1996),

and Walton et al. (1999)

Oil content Decreased Canvin (1965) and Mailer and Cornish (1987)Jensen et al. (1996), Champolivier and Merrien (1996),and Walton et al. (1999)

Oil quality Decreased oleic acid Pritchard et al. (2000) and Aslam et al. (2009)Protein content Increased Mailer and Cornish (1987)Glucosinolate content Increased Mailer and Cornish (1987)

Jensen et al. (1996)




Table 22.3. Effects of increased CO2 on yield and quality of oilseed Brassica.


Vegetativedevelopment

Increased Johannessen et al. (2002), Qaderi et al. (2006),and Franzaring et al. (2008)

Maturity Faster Franzaring et al. (2008)Yield Uncertain increase Franzaring et al. (2008)Oil content No significant change Frick et al. (1994) and Franzaring et al. (2008)

May increase Sator (1999)

There is also an indication that the response ofdifferent cultivars within a species and also dif-ferent species within a family will not responduniformly to elevated CO2. For example, pre-liminary results of the studies of Karowe et al.(1997) and Reddy et al. (2004) suggest differentresponses in the seed chemistry of different taxaand cultivars within species to elevated CO2.

Effect of climate changeon diseases and insects

Disease

Temperature and rainfall affect both the devel-opment of a pathogen (Huang et al. 2006) andthe resistance response of the host (Huang et al.2006). Projected temperature increases from cli-mate change will have profound effects on dis-eases of oilseed Brassica. For example, blackleg(Phoma) stem canker (Leptosphaeria maculans)is the most important disease of oilseed Brassicaworldwide (West et al. 2001). It causes yieldlosses of millions of tons across Europe, NorthAmerica, Australia, and Africa (Fitt et al. 2006).It has spread across North America and East-ern Europe in the last 20 years and now threat-ens China (Fitt et al. 2008). Multisite data col-lected over a 15-year period were used to developand validate a weather-based model forecast-ing severity of blackleg epidemics on oilseedBrassica across the United Kingdom. This wascombined with climate change scenarios to pre-dict that epidemics will not only increase inseverity, but also spread northward by the 2020s(Butterworth et al. 2010).

The faster development of blackleg diseasesymptoms at higher temperature regime iswell documented, with higher temperaturesincreasing symptom severity on cotyledons andleaves. The temperature regime is important indetermining both the rate and severity of crowncankers (Barbetti 1975). Evans et al. (2008) indi-cated that climate change will not only increaseseverity but also the range of over which stemcankers will be found. In Australia, the mostsevere blackleg disease epidemics are associatedwith temperatures ranging from 25–30◦C(Helms and Cruickshank 1979). Therefore, itis not surprising that currently the most severestem canker epidemics are in oilseed Brassicagrowing regions with Mediterranean climateslike Australia or France. Future expected risesin temperatures in association with climatechange, such as those predicted increases acrossthe oilseed-growing areas of Western Australia(Salam et al. 2009a, 2009b), will further increasedisease severity in all regions facing warmingtrends. It is fortunate that currently much ofthe world’s oilseed Brassica is grown in coolerclimates (Evans et al. 2008), but severity inthese countries is expected to also rise withwarming trends. Temperature increases can alsohave profound effect on the expression of hostresistance. It is now widely accepted that warmertemperatures can result in host resistance(s)being less effective (Badawy et al. 1991; Plieskeet al. 1998). Host resistance to stem canker caneven fail as temperatures increase (e.g., Hua Liet al. 2006, Salam et al. 2009a). Both the reducedresistance and more rapid and severe diseasesymptom development have been reported at




18/24◦C by Hua Li et al. (2006), at 27◦C byBadawy et al. (1991), and at 30◦C by Plieskeet al. (1998). Temperatures during the latter partof the growing season in many of the oilseedBrassica production regions of the world fre-quently exceed these temperatures. It is clear thatat least some genes associated with disease re-sistance are temperature dependent and becomeineffective at increased temperatures (Huanget al. 2006). This reduction in effectiveness ofresistance could be another reason why blacklegepidemics are more severe in the Mediterraneanclimates of Australia and France compared withthe United Kingdom (Barbetti and Khangura1999; Howlett et al. 2001; Sivasithamparamet al. 2005). Evans et al. (2008) showed that,as would be expected, the predicted effects ofclimate change on severity of blackleg were lesson canola cultivars with resistance to blacklegthan on susceptible cultivars. Clearly, one partof the strategy is to adapt to climate changeneeds to involve breeding canola cultivars withincreased resistance to blackleg, and to ensurethat this resistance can operate at the highertemperatures predicted from climate change.

Seasonal shifts in rainfall patterns as a con-sequence of climate change may not have thesignificant effects on the severity of some dis-ease levels as originally anticipated. For exam-ple, for blackleg stem canker, Salam et al. (2009a,2009b) showed that the onset of release of black-leg ascospores across Western Australia by 2030is likely to be delayed by about 2 weeks com-pared with release under current climate condi-tions. However, the opening seasonal rains to al-low commencement of the cropping season willalso be affected. In this scenario, it was delayedby about 1 week. Hence, the onset of blackleg as-cospores, relative to the break of the season, wasshifted to be only a week later than currently oc-curs. This means the risk of synchronization ofmajor blackleg ascospore showers with seedlingestablishment remains relatively unchanged un-der future climates.

Increases in CO2 can differentially affectdiseases of canola. For example, Evans et al.

(2010) examined the effect of five climate changescenarios for the United Kingdom in relationto varying CO2 emission levels and differenttimescales. Yield losses from stem canker andlight leaf spot (Pyrenopeziza brassicae) werecompared. Their modeling predicted increasesin severity and range of stem canker with cli-mate change, while losses because of light leafspot were predicted to decrease. The two diseasesactually compensated for each other so that thenet UK losses from climate change on untreatedoilseed B. napus would, in fact, be small underthe scenarios they tested.

Insects

Climate change factors, such as elevated CO2 andtemperature, typically affect carbon and nitrogendynamics of crop plants and the performance ofinsect herbivores (Himanen et al. 2008). Reddyet al. (2004) observed a negative larval growthrate at elevated CO2 for the crucifer special-ist Plutella xylostella (diamond back moth).Bezemer (1998) studied the impact of elevatedCO2 on two aphid pest species, Myzus persicaeand Brevicoryne brassicae. B. brassicae rearedon plants grown in elevated CO2 were larger andaccumulated more fat, while there was no changein M. persicae traits.

It is evident even from these limited ex-amples that climate change factors can affecthost–pathogen and host–insect relationships incomplex ways. A greater understanding of theseinteractions will be an important component ofadapting to climate change.

Breeding for climate change

The Brassicaceae family is genetically very di-verse and provides a large potential gene pool forbreeders to utilize in their efforts to develop newcultivars and species adapted to the changing en-vironments associated with climate change. Inaddition, oilseed Brassica is highly amenable togenetic engineering, providing breeders with theoption of genetically modified (GM) solutions.




Table 22.4. Potential breeding strategies for breeding oilseed Brassica species for climate change.

Breeding method Current status Future

Within species variation Incremental improvements identifiedfor climate change traits.

Limited variability within species due tocanola quality bottleneck

Mutation/tilling Limited use at present Tilling offers the possibility of a moredirected mutation strategy

Interspecific variation with acommon genome

Major genes for disease resistanceand quality have been transferred

Will continue to be an importantstrategy

Interspecific variation withouta common genome

Some B genome traits have beentransferred into B. napus

Continued interest in incorporating Bgenome traits into B. napus

Wild relatives Many useful traits identified Pre- and postfertilization barriersremain a major challenge

Resynthesis Increasing interest as a way toexpand current gene pool

Offers considerable potential toincorporate new variability

Protoplast fusion Limited use for nuclear traits Most likely use is in the development ofCMS systems

Changing species Canola quality B. juncea is replacingB. napus in some marginal areas

Canola quality B. juncea expected toincrease in importance

GM Available traits have been highlyeffective

Offers major potential to provide newkey genes for adaptation from wildrelatives or unrelated organisms

Oilseed Brassica breeders will need to use acombination of methods to most effectively de-velop highly adapted cultivars. The potentialbreeding options for oilseed available to Bras-sica breeders to utilize in breeding for climatechange are summarized in Table 22.4.

Utilizing existing within speciesgenetic variability

The initial focus on breeding oilseed Brassicaspecies for climate change is on identifying andutilizing existing variability within the oilseedBrassica species of interest. In general, breedingfor oilseed Brassica types low in erucic acid andin glucosinolates has led to a narrowing of thegene pool in elite cultivars (Gomez-Campo 1999;Cartea et al. 2005). Breeders and researchersare looking beyond the “canola quality pool” tosource variation for key traits. Singh and Chandra(2005) extensively screened germplasm of B. na-pus and other Brassica species for sources of re-sistances to climate change traits such as drought,

high temperature tolerance, salinity, frost, andearliness, as well as disease resistance and otheragronomic traits. Incremental variation was evi-dent in all traits. Gene pool diversity in B. napushas been shown with SSR markers to cluster intofour main groups: spring oilseed/fodder, win-ter oilseed, winter fodder, and vegetable types(Hasan et al 2006). The European Union Work-ing Brassica Group is working to phenotypicallycharacterize the morphology and phenology of acore collection of B. napus (Poulsen et al. 2002).It has conducted a preliminary evaluation of oilquality traits with wide diversity in erucic con-tent, resistance to clubroot disease, flea beetles,and stem weevils (Luhs et al. 2002).

The response of 25 strains of Ethiopian mus-tard (Brassica carinata) to drought stress wasassessed by Ashraf and Sharif (1998) in a potexperiment under glasshouse conditions. Four-week old plants of all the lines were subjectedto zero or two cycles of drought (twice wilt-ing and rewatering). Five strains were superiorto the remaining lines in production of shoot




fresh and dry masses under drought conditions.Only two lines, C90-1205 and 4007-A, were cat-egorized as moderately resistant on the basis oftheir performance in the two growth variables.Osmotic adjustment and water retention capa-bility were not successful selection criteria fordiscriminating the drought resistance of differ-ent lines. Incremental improvements for droughttolerance were evident. Similarly, Chauhan et al.(2007) screened 14 B. juncea lines for enhanceddrought tolerance and identified two lines, PSR-20 and JMMWR-941, as more tolerant acrosstwo sites. Salisbury and Berry 1983) screenedB. napus lines under a range of environments toidentify those lines best able to maintain higherlevels of oil content under stress.

Aksouh et al. (2001) used a series of hightemperature stress treatments differing in timingand duration to screen three B. napus cultivars fortolerance to high temperature stress. The cultivarOscar clearly demonstrated higher tolerance thanthe other cultivars, with significantly smaller re-ductions in yield and oil content compared to theother cultivars.

Screening trials specifically against high lev-els of greenhouse gasses, such as CO2, have beenlimited to date. Most of the oilseed Brassica tri-als with CO2 have been aimed at understand-ing the impact of CO2 on Brassica production,rather than screening for genetic differences. Be-cause of the difficulties of such studies, most ofthem have evaluated or modeled only very lim-ited numbers of cultivars or breeding lines, usu-ally only one or two (e.g., Qaderi et al. 2006;Franzaring et al. 2008). One exception was thestudy of Johannessen et al. (2002), who reporteddifferences between the six lines screened in re-sponse to the elevated CO2. Three lines increasedin yield, while the others decreased. As the linesthat increased in yield were genetically diverse,the authors indicted the potential for recombi-nation and further progress. It will be importantto develop screening protocols that allow muchlarger numbers of lines to be evaluated for ge-netic differences in response to increased levelsof CO2.

Breeders will continue to screen availablegermplasm for use within species variation inkey traits of interest. This will then be utilized inbreeding and selection programs to make incre-mental advances in these traits.

Mutation/tilling

Rahman et al. (1995) identified microspore-derived embryos of B. napus cultivar Topas thatsurvived salt stress after selection against oth-erwise lethal doses (0.6% and 0.7%) of NaClafter mutagen treatment. One of the lines (PST-2) accumulated less sodium and retained morepotassium, and was therefore able to maintaina more favorable Na : K ratio as compared tothe controls under salt stress. In general, the ran-dom nature of mutagenesis has meant that it isa somewhat less preferred approach in breedingfor climate change.

However, the synteny between Arabidopsisand Brassica offers the possibility of a more di-rected mutation strategy, known as reverse ge-netics or tilling. Where the location of genes thatcontrol key climate change traits are known inArabidopsis, synteny between the species can beused to identify potential loci of interest in Bras-sica. Variation at these loci, either natural or in-duced through mutation, can then be assessed inthe field.

Cultivated Brassicas

B. napus (AACC) has the C genome in com-mon with a B. carinata (BBCC) and B. oler-acea (CC) and the A genome in common withB. juncea (AABB) and B. rapa (AA). Hav-ing a common set of chromosomes significantlyenhances the likelihood of successful interspe-cific hybridization and gene flow between thetwo species. There are several successful exam-ples of gene flow between the cultivated species(Salisbury and Kadkol 1989). To date, these traitshave primarily been those associated with dis-ease and insect resistance and quality. Interspe-cific hybridization among B. napus, B. juncea,




and B. carinata (Getinet et al. 1997, Saal andStruss 2005) can be achieved with loss of chro-mosomes from the noncommon genomes in sub-sequent generations, and the transfer of desiredgenes for blackleg resistance through backcross-ing. The transfer of canola quality from B. napusto B. juncea has been reported by Burton et al.(2003) and Cowling and Wroth (2005). Trans-fer of blackleg resistance from B. juncea to B.napus has been reported by Saal et al. (2004).Many released cultivars have derived from inter-specific crosses between cultivated species with acommon genome (Salisbury and Kadkol 1989).From a climate change perspective, Singh andChandra (2005) have identified B. carinata andseveral B. juncea lines as excellent sources ofthermotolerance. This trait would be highly valu-able during seed development in many regions ofthe world.

Introgression from wild relatives

The wild relatives in the Brassicaceae familyare a potentially rich source of many usefulnuclear genes that offer potential for enhanc-ing adaptation of oilseed Brassica to climatechange conditions. Traits including drought tol-erance, high temperature tolerance, and salin-ity tolerance would be beneficial in breedingoilseed Brassica species adapted to the chang-ing climate (Salisbury and Kadkol 1989, Prakashand Bhat 2007). For example, Prakash and Bhat(2007) have reported drought tolerance in Bras-sica tournefortii, Diplotaxis acris, Diplotaxisharra, Eruca sativa, and Lesquerella species. B.tournefortii is also a likely source of thermo-tolerance. However, if genes from wild Bras-sicaceae species are to be successfully trans-ferred to cultivated species, a number of barriersneed to be overcome. These include pre- andpostfertilization barriers including issues of sex-ual compatibility, hybrid viability, and fertility,viability and fertility of progeny through sev-eral generations of backcrossing and successfulintrogression (incorporation of the gene) fromthe wild species into the chromosomes of the

cultivated species (Scheffler and Dale 1994; Binget al. 1991; Jorgensen et al. 1994, 1996a, 1996b;Salisbury (1989); Salisbury and Wratten 1997;Rieger et al.1999a; Salisbury 2002).

Despite the large number of crosses reportedbetween cultivated Brassica and the wild rel-atives, there is very little published informa-tion on the successful introgression of usefulnuclear genes from wild species to the cul-tivated species (Salisbury and Kadkol 1989).Where crosses have been made with the prac-tical purpose of transferring characters, sterilityin the F1 and subsequent generations have oftenlimited the transfer (Heyn 1977; Kumar et al.1988; Prakash and Bhat 2007). The vast ma-jority of these species belong to secondary andtertiary gene pools, rendering them virtually in-accessible to breeding programs (Prakash andBhat 2007).

One successful example involved the transferof aphid resistance from B. tournefortii to B. rapa(Prakash and Hinata 1980). Whether or not thisgene was successfully incorporated into a com-mercial cultivar is unknown. However, a morerecent successful exception comes from the workof Garg et al. (2010), who are the first to reporthigh levels of resistance against Sclerotinia scle-rotiorum in introgression lines derived from Eru-castrum cardaminoides, Diplotaxis tenuisiliqua,and Erucastrum abyssinicum with B. napus andB. juncea. These novel sources of resistance cannow be used in oilseed Brassica breeding pro-grams to enhance resistance in future B. napusand B. juncea cultivars against Sclerotinia stemrot. Where key genes for adaptation to climatechange are identified in secondary and tertiarygene pools, they may also be able to be accessedthrough GM technology.

In contrast to the limited success withtransferring nuclear genes, the most reward-ing utilization has been in the developmentof alloplasmics combining cytoplasm of wildspecies with the crop nuclei for expressing malesterility (Salisbury and Kadkol 1989; Bangaet al. 2003; Chandra et al. 2004; Prakash andBhat 2007).




Resynthesis

The development of synthetic allopolyploid linesof B. napus, B. juncea, and B. carinata using adiverse range of one or both respective diploidprogenitors would be expected to broaden thegenetic base of the allopolyploid species. Thisin turn will facilitate oilseed Brassica breedingprograms. It is likely that the wide genetic di-versity in the diploid crop gene pools has onlypartially been transferred to the allopolyploidcrops (Ladizinsky 1985). There is considerableopportunity to expand this. Negi et al. (2004) hasidentified diversity among B. nigra accessions.Identification of diverse germplasm has signifi-cant implications for planning new approachesto plant breeding. The diverse parental combina-tions can be used in generating segregated proge-nies with maximum genetic variability for breed-ing and selection.

B. napus resynthesis has been obtainedthrough use of doubled haploids of crosses be-tween B. rapa and B. oleracea (Zhang et al.2004; Abel et al. 2005). Success was highlydependent on the maternal genotype (B. rapa)with 0–1.18 embryos per pollinated bud. In an-other approach, resynthesis of B. napus with theB. rapa A genome and the C genome from B.carinata resulted in partial substitution of theA and C genomes in B. napus to achieve im-proved yield corresponding to the level of thissubstitution (Li et al. 2006). The doubled hap-loid technique was used by Gowers and Christey(1999) with hybrids between B. oleracea andB. napus and backcrossing the hybrid B. napol-eracea to B. napus. This enabled transfer ofaphid resistance, self-incompatibility, and newcombinations of glucosinolates for B. napus.Kahlon et al. (1999) screened both the B. rapaand B. oleracea landraces for tolerance of man-ganese toxicity, selecting superior accessionsfor potential use in resynthesis of B. napus.The resynthesis approach offers considerable po-tential for new breeding lines with enhancedtraits needed for adaptation to changing environ-ments. Prakash et al. (2009) reported that while

synthetic cultivars tend not to compete with re-leased cultivars for yield, they provide a richdiversity of traits for productivity, quality, anddisease resistance that can be backcrossed intocurrent lines.

Protoplast fusion

There are many reports of somatic hybridizationin the Brassicaceae (Salisbury and Kadkol 1989,Prakash et al. 2009). While these somatic hybridshave crossed intergeneric and tribal barriers,there is a high general degree of sterility or severeintergenomic incompatibility, leading to manyabnormalities (Prakash et al. 2009). Asymmet-ric hybrids appear to be more promising option,tolerating only a fraction of alien genetic con-tent. Many of these somatic hybrids developedhave been aimed at incorporating new sourcesof nematode or disease (Sclerotinia, Alternaria,blackleg) resistance (Prakash et al. 2009). Oneof the limiting factors when aiming to developsomatic hybrids involving one wild relative isthe resultant very low level of, or complete ab-sence of, pairing between the wild and cultivatedgenomes. One recent highly successful examplewas the fusion of protoplasts using two canolalines. This allowed the incorporation of the twocytoplasmic traits—triazine tolerance and cyto-plasmic male sterility (CMS). Triazine tolerantCMS hybrids are now commercially available inAustralia.

Several CMS systems in B. napus and B.juncea have been obtained following protoplastfusion. They are based on Raphanus, B. tourne-fortii, Diplotaxis catholica, E. sativa, Morican-dia arvensis, Sinapis arvensis, Orychophragmusviolaceus, Trachystoma ballii, and Arabidopsisthaliana (Prakash et al. 2009). The most widelyused system currently is the Ogura system fromRaphanus. The most effective use of this pro-toplast technology from a climate change per-spective is likely to be in the continued devel-opment of new CMS systems, leading to higheryields.




Utilizing GM traits

Oilseed Brassica species are benefiting from be-ing at the forefront of GM technology. Thistechnology provides potential access to bene-ficial genes outside the traditional barriers ofsexual compatibility. GM traits such as droughttolerance, high temperature tolerance, salinitytolerance, and enhanced disease resistance thatwill be beneficial in coping with climate changeare currently being evaluated in oilseed Bras-sica species. Wan et al. (2009) reported break-throughs in the understanding of key molecularcomponents that regulate homeostasis and sens-ing of abscisic acid and their potential applica-tions in genetic engineering for drought-tolerantcanola. In particular, the α and β subunits of theprotein farnesyltransferase have been identifiedas negative regulators of abscisic acid (ABA)-mediated stomatal responses. Their effective-ness as the targets for engineering drought toler-ance and yield protection has been confirmed incanola in the field. Further development of thistolerance to drought stress in canola is expectedto significantly enhance productivity in many re-gions of the world.

The growth of transgenic canola (B. na-pus) expressing a gene for the enzyme 1-aminocyclopropane-1-carboxylate (ACC) deam-inase was compared to nontransformed canolaexposed to flooding and elevated soil Ni concen-tration in situ by Farwell et al. (2007). The trans-genic canola had greater shoot biomass com-pared to nontransformed canola under low flood-stress conditions. This is the first field study todocument the increase in plant tolerance utiliz-ing transgenic plants exposed to multiple stresses(Farwell et al. 2007).

Prasad et al. (2000) used Agrobacterium me-diated transformation to transform B. juncea cv.Pusa Jaiksan with the codA gene for biosynthe-sis of glycinebetaine from Arthrobacter globi-formis. The seeds of the transgenic lines showedenhanced capacity to germinate under salt stressas compared to that of the wild type. In addition,the seedlings of transgenic plants that expressed

codA gene showed significantly higher growththan the wild type under salt-stress conditions.These results demonstrated that the introductionof a biosynthetic pathway for glycinebetaine intoB. juncea significantly enhanced their salt toler-ance. Homozygous genotypes of selected trans-formed lines can be exploited for improving thesalt tolerance of desirable cultivars of B. junceathrough breeding programs.

The potential to reach beyond sexual barri-ers to access useful genes with GM technologycould play an important role in breeding adaptedoilseed Brassica for climate change. Many ma-jor genes from the secondary or tertiary genepools of the Brassicaceae family or from totallyunrelated species could be accessed.

Changing species

In some environments, the changing climate maymean that a different species is required. For ex-ample, summer B. rapa is currently the dominantoilseed crop in Finland. It has lower productionrisks compared with summer oilseed B. napus.However, climate-change-induced increases intemperature could see increased use of oilseed B.napus at the expense of B. rapa (Peltonen-Sainioet. al. 2009). Similarly, in higher altitude regionswhere B. rapa has traditionally been grown, B.napus may become the preferred option.

Similarly, in lower rainfall marginal canola-growing areas of Australia and Canada, there isincreased interest in the use of canola qualityB. juncea to deal with climate change. WhileB. napus and B. rapa have been the traditionalspecies used in the production of canola qualityBrassica oil, B. juncea has also recently beenconverted to canola quality. B. juncea is betteradapted to the high temperatures and lower rain-fall conditions expected with climate change. B.juncea has superior heat and drought tolerance,disease resistance, and pod-shatter resistance rel-ative to B. napus (Kirk and Oram 1981, Woodset al. 1991, Burton et al. 1999). Wright et al.(1995) confirmed the significant advantage of B.juncea over B. napus in situations of soil water




deficit. This was due to the ability of B. juncea tomaintain higher leaf turgor than B. napus undersoil water deficits, resulting in greater yield ofB. juncea under such conditions (Wright et al.1997). Climatic changes may mean that overtime, a higher proportion of the canola produc-tion comes from B. juncea at the expense of B.napus. Canola quality B. juncea has shown asignificant yield advantage over B. napus in sev-eral low-rainfall environments, especially underconditions of water stress and high temperaturesduring seed fill. Canola quality B. juncea culti-vars have been commercialized in Australia andCanada in recent years (Burton et al. 2003, Oramet al. 2005).

Other technologies

Molecular markers are increasingly being used toenhance traditional breeding and selection pro-grams in oilseed Brassica. They are being usedto select parental genotypes in breeding pro-grams, eliminate linkage drag in backcrossing,and select for traits that are difficult to measureusing phenotypic assays (Edwards and Batley2010). When used at the seedling stage in con-junction with doubled haploid technology, theycan be used to eliminate many progeny priorto taking the progeny to the field. For exam-ple, Burton et al. (2004) utilized markers at thisseedling stage in their canola quality B. junceabreeding program. Molecular markers can beparticularly useful for selecting for complexpolygenic traits.

The increasing availability of DNA sequenceinformation will enable the discovery of genesand molecular markers associated with diverseagronomic traits. A high-quality genome se-quence for B. rapa is due for release shortly.However, at present the function of many genesidentified by genome sequencing is unknown(Edwards and Batley 2010). When it becomesavailable, the information on genes (and mark-ers) linked to key traits will help to accelerateoilseed Brassica breeding.

Conclusions and future directions

Utilization of variability within existing speciesis likely to provide oilseed Brassica breederswith continued incremental gains in key cli-mate traits. However, if major advances are to bemade, other breeding options will be required.This includes incorporation of key genes fromother cultivated genomes and also from weedyrelatives if sexual barriers can be overcome.The resynthesis of the allopolyploid species us-ing a diverse range of germplasm from bothdiploid parents offers potential access to a rangeof new germplasm. This will require access togermplasm collections from around the world.The use of GM technology to access major genesfrom wild relatives or from completely differ-ent species offers major potential benefits tobreeders.

Changing species will offer significant ben-efits to growers in some regions. Canola qual-ity B. juncea is better adapted than B. napus tothe increasing temperatures and increasing mois-ture stress expected in coming generations. In thedrier regions, B. juncea may gradually replace B.napus as the preferred crop. The increasing avail-ability of genomic information on genes (andmarkers) linked to key traits will also help toaccelerate progress in oilseed Brassica breedingprograms. Breeding for blackleg resistance willremain an important component of most breed-ing programs. The available resistances will needto be managed carefully to ensure their longevity.Oilseed Brassica breeders will likely utilizeseveral different methodologies to successfullydevelop cultivars adapted to climate changeconditions.

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Crop Adaptation to Climate Change (Yadav/Crop Adaptation to Climate Change) || Breeding Oilseed Brassica for Climate Change