Plant Breeding for Abiotic Stress Tolerance Roberto Fritsche-Neto Aluízio Borém • Editors Plant Breeding for Abiotic Stress Tolerance 123 Editors RobertoFritsche-Neto Aluízio Borém Departamento de Fitotecnia Departamento de Fitotecnia Universidade Federalde Viçosa Universidade Federalde Viçosa Av.PH Rolfss/n Av.PH Rolfss/n Viçosa, MG36570-000 Viçosa, MG36570-000 Brazil Brazil ISBN 978-3-642-30552-8 ISBN 978-3-642-30553-5 (eBook) DOI 10.1007/978-3-642-30553-5 SpringerHeidelbergNewYorkDordrechtLondon LibraryofCongressControlNumber:2012940243 (cid:2)Springer-VerlagBerlinHeidelberg2012 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionor informationstorageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilar methodology now known or hereafter developed. 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While the advice and information in this book are believed to be true and accurate at the date of publication,neithertheauthorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityfor anyerrorsoromissionsthatmaybemade.Thepublishermakesnowarranty,expressorimplied,with respecttothematerialcontainedherein. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface Practical experience shows that abiotic stresses occur at high or low intensity in just about all agricultural growing areas around the globe. In some regions the stressduetoheat,drought,andmineralnutritionorevenduetoaluminumtoxicity may not be present in a specific year, but this could change in the coming years. The consequences of the climate change for agriculture have caused concern for manypolicymakersaroundtheworldandithasbeenahottopicinmanyscientific forums.Themainapprehensionisthattheworldmaynotbeabletosustainitsfood production under the crescent abiotic stresses with global warming. The organizers of this book have pulled together this publication aiming to cluster the most relevant scientific achievements and the state of the art in plant breeding and cultivars development for abiotic stresses. Writtenbyexpertsindifferentareasofabioticstresses,inaneasytounderstand language, this book is an obligated reference for all interested in plant breeding and in the upcoming challenges that agriculture will face with climate change. Roberto Fritsche-Neto Aluízio Borém v Contents 1 Abiotic Stresses: Challenges for Plant Breeding in the Coming Decades. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Aluízio Borém, Magno Antonio Patto Ramalho and Roberto Fritsche-Neto 2 Breeding for Stress-Tolerance or Resource-Use Efficiency? . . . . . 13 Roberto Fritsche-Neto and Júlio César DoVale 3 The Physiology of Abiotic Stresses . . . . . . . . . . . . . . . . . . . . . . . 21 Paulo C. Cavatte, Samuel C. V. Martins, Leandro E. Morais, Paulo E. M. Silva and Fábio M. DaMatta 4 Breeding for Nitrogen Use Efficiency. . . . . . . . . . . . . . . . . . . . . . 53 Júlio César DoVale, Rodrigo Oliveira DeLima and Roberto Fritsche-Neto 5 Breeding for Phosphorus Use Efficiency . . . . . . . . . . . . . . . . . . . 67 Sidney Netto Parentoni, Flávia Ferreira Mendes and Lauro José Moreira Guimarães 6 Breeding for Water Use Efficiency . . . . . . . . . . . . . . . . . . . . . . . 87 Marcelo de Almeida Silva, Claudiana Moura dos Santos, Carlos Alberto Labate, Simone Guidetti-Gonzalez, Janaina de Santana Borges, Leonardo Cesar Ferreira, Rodrigo Oliveira DeLima and Roberto Fritsche-Neto 7 Breeding for Salinity Tolerance . . . . . . . . . . . . . . . . . . . . . . . . . 103 Nand Kumar Fageria, Luís Fernando Stone and Alberto Baêta dos Santos vii viii Contents 8 Breeding for Aluminum Tolerance . . . . . . . . . . . . . . . . . . . . . . . 123 Lauro José Moreira Guimarães, Claudia Teixeira Guimarães, Jurandir Vieira Magalhães, Sidney Netto Parentoni and Flávia Ferreira Mendes 9 Breeding for Heat-Stress Tolerance. . . . . . . . . . . . . . . . . . . . . . . 137 Moacil Alves de Souza, Adérico Júnior Badaró Pimentel and Guilherme Ribeiro 10 Breeding Perennial Species for Abiotic Stress . . . . . . . . . . . . . . . 157 Rinaldo Cesar de Paula, Nadia Figueiredo de Paula and Celso Luis Marino Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 Chapter 1 Abiotic Stresses: Challenges for Plant Breeding in the Coming Decades Aluízio Borém, Magno Antonio Patto Ramalho and Roberto Fritsche-Neto Abstract Modern agriculture has been providing food, feed, fiber and more recently biofuel to meet the World´s demand. One of the bases of this modern agriculture is the improved cultivars, which are much higher yielder than ancient ones. The scientific literature documents the significant contribution of plant breeding to agriculture and finally food production worldwide. Now-a-days, agriculturehasnewandhugechallenges,duetopopulationgrowth,thepressureon agriculture liability on the environmental conservation, and climate change. To cope with these new challenges, many plant breeding programs have reoriented their breeding scope to stress tolerance in the last years. So, in this book, experts on plant physiology and on plant breeding presents the most recent advances and discoveries applied toabiotic stresses, discussing the new physiological concepts, breeding methods, and modern molecular biological approaches to develop improved cultivars tolerant to most sorts of abiotic stress. Keywords Climatechange(cid:2)Globalwarming(cid:2)Improvedcultivars(cid:2)Breedingfor draught (cid:2) Biotechnology A.Borém(&)(cid:2)R.Fritsche-Neto UniversidadeFederaldeViçosa,Viçosa,Brazil e-mail:[email protected] R.Fritsche-Neto e-mail:[email protected] M.A.P.Ramalho UniversidadeFederaldeLavras,Lavras,Brazil e-mail:[email protected]fla.br R.Fritsche-NetoandA.Borém(eds.),PlantBreedingforAbioticStressTolerance, 1 DOI:10.1007/978-3-642-30553-5_1,(cid:2)Springer-VerlagBerlinHeidelberg2012 2 A.Borémetal. 1.1 Introduction Agriculture was the most important event in the man’s history, changing his nomadic lifestyle to start to settle in communities. Archeological evidences sug- gestthatagriculturebeganabout10,000–12,000 yearsago.Withagriculture,man accelerated his influence in the environment, especially by substituting the native and diverse flora by the species of their preference or needs. Atthebeginningofthenineteenthcentury,therewasanincreasingconcernthat therapidgrowthofthepopulationcouldresultfoodshortageandthepoor-resource population would go famine. The most famous writing about this was by the economist Thomas Malthus, who said that the population increase according to a geometric progression while the food supply would increase according to an arithmetic progression. While Malthus model was correct, he did consider the agricultural technology development such as use of fertilizers and especially the adoption of genetically improved cultivars. Therefore, even with the population increasing to 4.68 billion people during the twentieth century, food was available to meet the demand, except for short periods of time. It is important to say that today hunger is mainly due to wealth distribution and not due to food scarcity. Withmodernagricultureusingimprovedcultivarsandgoodcropmanagement, crop yield has been raised in just all agronomic species. This progress was more evidentaftertheSecondWorldWar,withthewideadoptionofimprovedcultivars, fertilization,insecticides,andfungicides.Withthesuccessofmodernagricultureto producefoodinquantityandquality,thepricesoffooddroppedduringthetwentieth century.Thefoodabundance,asaresultofagriculturaltechnology,haditsowntoll; society lost the sense that agriculture had importance. With plenty offood at low prices,manyotherareasofthehumanknowledgestarttodrainasignificantpartof theinvestmentsinR&D,andtodaytheagriculturalresearchlivesahugedilemma; how to develop the technology for the coming decades, especially with the new challengeswithclimatechange.Oneofthesignsoftheshortageofinvestmentsfor agriculturalresearchisthesmallnumberofnewlytrainedplantbreedersinthelast decade, in spite of plant breeding’s contribution to increase food production worldwide(VencovskyandRamalho2006;Duvicketal.2004). Althoughthepopulationthatgoeshungryhasbeenreducedinthelast50 years, there are evidences that it could spike in the medium and long run. Currently, about925millionpeopledonothaveaccesstotheminimumdailyamountoffood recommended by United Nations WHO considering the macronutrients (carbo- hydrates,proteins,andlipids).Itisestimatedthatanother1billionpeoplehavethe occulthunger,thatis,havebadnutritionwithdeficiencyofvitaminsandminerals (Evans, 1998; Borém and Rios 2011). On the other hand, nearly another 1 billion people is consuming food in excess with risk of obesity and diabetes. Nowadays,agriculturehasnewandhugechallenges,whichtendtobeevenbigger incomingyears.Thisisduetothefactthatthepopulationissupposedtoincreaseup to2050,whenaccordingtotheUnitedNationFAOtheworldpopulationwillsta- bilizereaching9.1billioninhabitants.Thepopulationincreasewillhaveprofound 1 AbioticStresses:ChallengesforPlant 3 Table1.1 Worldfoodandfiberproductiondemandin2005and2025(inmillionsoftons) Product Production Demand Additional in2005 in2025 production Grains 2.219,40 3.140,40 921 Oilygrains 595,05 750,97 155,96 Perennialcrops 242,81 321,99 70,18 Annualcrops 352,2 437,98 85,78 Coffee 7,72 9,4 1,68 Fibers 28,5 36,37 7,87 Timber 3.401,90 4.148,40 746,5 SourceAdaptedfromAgroanalysis(2007) Fig.1.1 Percapita consumptionoffivefood productsinemerging countriesfrom1996to2001. SourceAgroanalysis(2011) outcomestoourplanet.Itissurprisingthat,evenbeforethissentenceiscomplete, therewillbeanotherthreeinhabitantsontheEarth,thatis,anewpersonisbornevery three seconds, 259,000 new inhabitants per day, 7 million per month. It is worth notingthatittookseveralmillionyearsfortheEarthtoreachitsfirst2billionpeople, and that another 2 billion will be brought just in the next 25 years. Additionally, peoplearelivinglongerandalsomovingfromthecountrysidetourbanareas.Halfof theworldpopulationcurrentlyliveincitiesandsmalltowns,butthisnumbershould increaseupto60 %by2030,whenitissupposedtoexist26citieswithpopulation over10millionpeople(Beddington2010). Thechallengestobefacedbymanduetothepopulationincreaseinthecoming decades are various and often debated (UK Government 2011). One of them is how to provide food to meet the demand. On Table 1.1 is listed the world food productionanddemandin2005andin2025.Theincreaseddemandforfoodisnot onlyduetopopulationincrease,butalsoduetomanyotherfactors,amongthemis the increase in population wealth, especially in the emerging countries called BRICS(Brazil,Russia,India,China,andSouthAfrica),resultingindemandfora more diverse and rich diet (Fao 2009, 2010; Fig. 1.1a). It is well known that an increase in the per capita income results in an increase in food demand, and the BRICS countries represent 40 % of the world population and nearly 18 % world economy. 4 A.Borémetal. Anotherfactorpressingthefoodproductionisthecompetitionbetweentheareas designedforbioenergyproduction.Thedemandforbioenergyshouldincrease45 % inthenext25 years,andgrowingareasusedforgrainwillbeallocatedtosugarcane, jatropha,andotherenergycrops(Mitchel2008;Beddington2010). The pressure on agriculture will be even greater due to its environmental lia- bility. Among the environmental aspects used as input in agricultural water deserves a special attention. It is estimated that 70 % of the drinkable water is currently used for irrigation (Chris and Briscoe 2001). Therefore, water must be used more efficiently. Additionally, it is estimated that many areas around the world will have less soil water, creating the need for irrigation to be more wide- spread and intense in many growing areas. Therenewableagricultureinputswillalsobecomescarcerandmorecostly.For example, Brazil uses 2.5 millions of nitrogen, i.e., 2.5 % of the world consump- tion, and 3.7 million of K O, making up to 13.9 % of world consumption. 2 To complicate the scenario even more, there is an increasing pressure for loweringtheCO emission(Wright2010)anditisregardedagriculturewillsuffer 2 most (Lobell et al. 2003; Assad and Pelegrino 2007). To cope with this scenario many marginal areas will be added to the agricultural system. Many of those marginal areas have aluminum toxicity, salinity, or other abiotic stresses, demanding for cultivars adapted to such constrains. Mostoftheagriculturalsystemsarenotsustainable,mainlyduetotheintenseuse ofnon-renewableresourcesreinforcingthefeelingformorestress-tolerantcultivars. Thelackofsustainabilitywillmaketheinsects,diseases,weeds,andbioticstresses more prevalent and severe. To deal with those challenges, more planning and new technology is necessary. Besides the traditional techniques and breeding methods, Biotechnology(Quam2009;BorémandAlmeida2011)andotheremergingtechnol- ogies,suchasNanotechnology(Alexandratos2006;Bruinsma2009)mustbeadopted. The contribution of plant breeding to increase food, feed, fiber, and biofuel production(VencovskyandRamalho2006;Duvicketal.2004)iswelldocumented. Nevertheless,whateverhappensinthecomingdecade,plantbreederswillhaveto devote profound dedication and deep scientific knowledge to keep pace with the increasing food demand in a hungry world. The main goal of this chapter is to address the scope of abiotic stresses for agriculture and how to cope with them, helpingplantbreederstoestablishrightprioritiesanddevelopadequatecultivars. 1.2 Crop Yield Potential Beforediscussingwhatpotentialyieldis,itisneededtodefinebiologicefficiency. Theconceptofbiologicefficiency varieswiththecropspeciesandsituations.For example, for food, feed, and fiber biologic efficiency depends on the solar energy conversion. Yield should reflect the unit of the product per unit of absorbed solar energy.Cropscientistsandeconomists,amongothers,prefertoassociatebiologic efficiency to unit of area, such as amount of the product (kg, tons) per area unit