Crop Adaptation to Climate Change (Yadav/Crop Adaptation to Climate Change) || Adaptation of the Potato Crop to Changing Climates

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Chapter 11

Adaptation of the Potato Cropto Changing ClimatesRoland Schafleitner, Julian Ramirez, Andy Jarvis, Daniele Evers, Raymundo Gutierrez,and Mariah Scurrah

Introduction

Potato (Solanum tuberosum L.) is the third mostimportant food crop after rice and wheat with aworldwide production of 314 million tons (FAO-STAT 2008). During the last decades, demand forpotato increased strongly, particularly in devel-oping countries, where this crop is ever more rec-ognized as key for food security, since it yieldsmore nutritious food under harsher conditionsthan any other major crop. Actually, potato isgrown in 149 countries from latitudes 65◦N to50◦S and at altitudes from sea level to 4000 m(Hijmans 2001). In spite of its broad geograph-ical distribution, modern cultivars require mod-erate climates and appropriate water supply forhigh yields.

Climate change scenarios project a significantrise of average temperatures in the twenty-firstcentury, accompanied by more frequent extremeweather events such as droughts and heat waves.Higher temperatures also increase the crop waterrequirement, which together with changing rain-fall patterns could bring about greater droughtstress risks for potato. Crop growth models sug-gest potato yield decreases in consequence to

climate change of up to 32% on a global scaleuntil 2050.

Expected climate change effectsfor potato areas

Climate changes projections for potato grow-ing regions around the globe for 2020 were es-tablished as described in Beebe et al. (Chap-ter 16, this book). Changes of precipitation arepredicted to remain relatively modest, while tem-perature changes are likely to be more criticalfor potatoes, as their predominant cultivationareas overlap with regions with high projectedtemperature raises. Changes in suitability ofpotato cultivation areas were determined usingthe niche-based Ecocrop model (Hijmans 2001)calibrated with potato production data fromHijmans (2001). According to this model,present suitable potato cultivation areas wereidentified, which greatly overlap with the ac-tual production areas (Fig. 11.1a). However, asthis model includes precipitation data, it doesnot pinpoint areas, where potato is grown un-der irrigated conditions (many of these regionsare marked in gray in Fig. 11.1a). The current

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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288 CROP ADAPTATION TO CLIMATE CHANGE

(a)

(b)

(c)

Fig. 11.1. (a) Area harvested from FAOSTAT (2009), (b) predicted suitability for current conditions, (c) current climaticconstraint for potato cultivation, (d) predicted future suitability (2020) using an ensemble (average) of four GCM patterns, and(e) predicted suitability change (future to current) by 2020.



ADAPTATION OF THE POTATO CROP TO CHANGING CLIMATES 289

(d)

(e)

Fig. 11.1. (Continued)

abiotic constraints to present production showminimum temperatures to be the limiting fac-tor in high latitude areas, while most subtropicalregions are more limited by maximum tempera-tures than by other climatic factors (Fig. 11.1b).According to this model, drought is a limitingfactor in a few cultivation areas only; however,the present model does not take in account theoccurrence of drought spells and thus probablysubestimates actual and future drought risks de-rived from more extended dry seasons through-out the year.

Future climate suitability for potato cultiva-tion is likely to be severely affected by climatechange by 2020 (Fig. 11.1c,d). The most signifi-cant losses in potato suitability occur in tropical

highlands and in southern Africa as well as inpotato-growing areas at high latitudes (supportedby Tubiello et al. 2001; Hijmans 2003). However,the average expected change in suitability is ac-tually positive (+1.3%); but these gains, whichare considerable in area and amount (Fig. 11.1d),occur in very high latitudes or high tropical al-titudes, where the current climatic constraint areminimum temperatures.

We evaluated the potential benefits fromimproved abiotic stress tolerance in potatoes(Fig. 11.2) and found that from the total currentpotato harvested area, some 7.7 million hectares(65.3% of current potato cropping area) wouldbe less impacted by climate change and croppingarea could expand to 15.5 million new hectares,




Degree of improvement

Cha

nge

in s

ize

of s

uita

ble

area

s (%

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Fig. 11.2. Expected potential benefits from different breeding technologies for pota-toes. Degree of improvement indicates the amount at which growing parameters weremodified (each 0.5◦C for cold and heat tolerance and each 5% for drought tolerance).

if improvements in heat tolerance are made. Coldtolerance would bring opportunities for expand-ing the crop’s agricultural frontier toward morenorthern or southern latitudes, with 8.7 millionhectares of new land benefitting. Drought toler-ance appears as the trait with the least impactwith 2.8 million hectares benefitting from tol-erance, most of which overlap with areas thatwould benefit from heat tolerance improvement.As already mentioned above, this assumptiondoes not include drought risks due to higher wa-ter requirement under heat stress.

Projected rises of the CO2 concentrationcould bring about additional benefits for potatoproduction in form of a substantial increase ofnet photosynthesis, resulting in yield increasesin some regions (Miglietta et al. 2000).

Beside increased abiotic stress, climatechange will also bring about changes in pestand disease incidence for potato. Both pathogensand beneficial organisms can be affected directlyor indirectly by changing environmental condi-tions. Warmer temperatures increase the num-ber of insect and nematode generations makingthese pests probably more abundant (Porter et al.1991). Potato tuber moth (Phthorimaea oper-culella Zeller) develops faster under higher tem-

peratures and is projected to spread northwardand upward (Sporleder et al. 2008). Also, fungaland bacterial disease incidence depends on theweather, and climate change is likely to promotenew emerging diseases for potato. However, drierand warmer conditions during the summer couldreduce the incidence of the most severe potatodisease, late blight in some regions. Droughtevents also can favor disease outbreaks, suchas aphid-borne virus diseases (Bagnall 1988).Shifts in pest and disease incidence in potatowill require changes in crop protection strategies,and according to data summarized by Koleva andSchneider (2009), hotter temperatures will bringabout higher pesticide costs for potato.

Potato responses to climatechange effects—heat, drought,and cold stress

Heat stress is a serious threat for potato pro-duction. Elevated temperatures accelerate leafsenescence and double respiration rates for each10◦C increase, resulting in a negative carbon bal-ance on hot days (Winkler 1971). Tuber initia-tion is restricted in most genotypes when nighttemperatures exceed 17◦C (Menzel 1985; Prange




et al. 1990) and the optimal temperature for tuberbulking ranges between 14◦C and 22◦C. Highertemperatures decrease tuber growth (Basu andMinhas 1991) and can cause misshapen or chaintubers, high sugar levels, low starch contents,and glycoalkaloid accumulation (Kleinkopf et al.1988). Under hot and dry conditions, high va-por pressure deficit increases transpiration andinsufficient water supply causes drought stress,resulting in further yield losses.

From available studies, we can conclude thatphotosynthesis, respiration, carbon partition, andhormonal effects affect yield in potato under heatstress, but we know very little about the geneticor physiological basis of tolerance. In other plantsystems, it has been shown that photosynthesislimitation under heat is conferred by a mixture ofribulose bisphosphate carboxylase activity, elec-tron transport, and pyrophosphate regenerationeffects (Sage and Kubien 2007). In potato underheat stress, both increase and decrease of photo-synthetic rate was observed (Lafta and Lorenzen1995; Timlin et al. 2006), but increased respi-ration rate appears to have a greater effect onnet carbon accumulation under heat than drop ofphotosynthetic rate (Sarquis et al. 1996). Moremoderate increases of respiration rates have beenrecognized as a heat-tolerance trait in Solanumchacoense and Solanum acaule (Wivutvongvana1979).

Disturbance of carbon partition is one of themain implications of heat stress on potato pro-ductivity. It was shown that under heat stress,sucrose is still transported to tubers, but conver-sion to starch is inhibited (Wolf et al. 1991). Inconsequence, sucrose concentrations rise at thephloem unloading site, resulting in a reductionof overall sucrose transport through the phloemto the tubers, as this transport relies on a su-crose gradient between source and sink tissues.This suggests that reduction in tuber growth andstarch deposition under heat at the first placeis not a result of decreased assimilate availabil-ity, but is due to declined sink strength of thetubers. Heat tolerance candidate genes that in-fluence sink strength might be involved with

the regulation of phloem unloading, sucrose andstarch metabolism in tubers, such as sugar trans-porters, invertase, sucrose synthase, and ADP-glucose pyrophosphorylase (Viola et al. 2001).ADP-glucose pyrophosphorylase is regulated al-losterically, on transcriptional level and by theredox status of the cell (Tiessen et al. 2002).Abiotic stress such as heat alters the cellularredox status through induction of reactive oxy-gen species production (reviewed by Suzuki andMittler 2006), so accumulation of reactive oxy-gen species under heat could be involved withstarch synthesis and sink strength regulation anddetoxification of reactive oxygen species couldcontribute to heat tolerance. The heat stress re-sponse on gene expression level in potato tu-bers and tuber periderm, respectively, was tar-geted by microarray studies (Rensink et al. 2005;Ginzberg et al. 2009) that could be mined to iden-tify further target genes for heat stress effects andtolerance.

Tuber initiation is even more susceptible toheat stress than carbon partition, although a fewaccessions of different potato species show con-sistent tuberization at 30–40◦C (Reynolds andEwing 1989). Tuberization in potato consists ofa complex process involving various hormonaland environmental signals (reviewed by Sarkar2008). The heat-triggered inhibitory effect is me-diated through increased gibberellic acid levels,but the temperature-responsive component re-mains to be elucidated.

Beside heat stress, drought stress risk mightincrease under climate change conditions. Tu-ber production generally correlates with plant-available water; however, some genotypes pro-duce higher yields than others under limitedwater supply. Considerable variation in droughttolerance has been found in potato varieties,breeding clones, landraces, and wild species(Levy 1986; Ochoa 1998; Schafleitner et al.2007a; Coleman 2008). Screening tetraploidbreeding clones and landraces with differentploidy levels from the potato collection heldin trust by the International Potato Center re-vealed up to now 192 accessions with yield drops




below 25% when exposed to drought from day40 after planting in the winter season of the Pe-ruvian coastal desert, while susceptible cloneslost up to 100% of their yield under the sameconditions (Enrique Chujoy, Rolando Cabello,unpublished). A trait related to drought tol-erance is high water-use efficiency. Water-useefficiency depends on stomatal density and be-havior, mesophyll conductance, and photosyn-thetic efficiency. Large variation of water-useefficiency, i.e., the amount of tuber dry mass pro-duced per liter of water, has been found in ad-vanced breeding clones (Raymundo Gutierrez,unpublished).

Drought tolerance may be based on reducedtranspiration, improved water uptake, toleranceto a certain degree of dessication, or a combi-nation of these mechanisms. Reduced stomatalbehavior minimizes water loss and thus con-tributes to turgor maintenance, but also limitsCO2 access to the photosynthetic apparatus forcarbon assimilation. Enhanced water uptake isanother way for plants to remain turgescent un-der water stress conditions. Long roots increasethe water uptake capacity of plants and sustainleaf growth and plant productivity under droughtin environments, where water remains availablein deeper soil layers. Variation in root length hasbeen found in potato, and tuber yield was signifi-cantly correlated to root dry mass under stressedconditions (Iwama 2008).

Molecular and biochemical investigations in-dicated several tolerance mechanisms present inpotato landraces. Results of gene expression pro-filing experiments with microarrays pointed to-ward detoxification of reactive oxygen species asone of the key traits endorsing survival and tuberproduction under water stress. Genes encodingenzymes such as superoxide dismutase, ascor-bate peroxidase, catalase, glutathione peroxi-dase, and peroxiredoxin were strongly inducedunder drought in tolerant accessions (Watkinsonet al. 2006; Schafleitner et al. 2007b; Mane et al.2008; Vasquez-Robinet et al. 2008). Addition-ally, compounds with active oxygen scavengeractivity, such as sugar alcohols, amino acids, and

an array of secondary metabolites, accumulatedunder water stress in drought tolerant potato andpresumably mitigate the deleterious effects ofactive oxygen species in the cells (Mane et al.2008). Osmotic adjustment, which contributesto turgor maintenance under drought stress andprotects membranes and proteins from harmfuldesiccation effects, could also be a factor indrought tolerance, although its usefulness as atolerance trait is controversially discussed. Atleast it could support root development in orderto reach water that may be available deeper in thesoil profile (Serraj and Sinclair 2002). Expres-sion profiling data further suggest a role of cellrescue mechanisms including membrane modi-fication and protein stabilization by chaperones,late embryo abundant, and heat shock proteinsto drought tolerance of potato (Schafleitner et al.2007b).

Potato cold tolerance is also an important traitfor adaptation to climate change scenarios. Whenplanting dates are anticipated or potato cultiva-tion is shifted to higher altitudes and latitudesin the course of adaptation strategies to avoidheat stress, the crop might be exposed occa-sionally to lower temperatures than in the cur-rent potato cultivation areas and thus suffer fromcold stress or frost. According to Oufir et al.(2008), cold and drought stress share many com-mon features. Among the molecular responses tocold exposure in potato, genes involved in sev-eral metabolisms such as amino acid, carbohy-drates, energy, detoxification, and photosynthe-sis, as well as dehydrins and antifreeze proteins,are differentially regulated (Oufir et al. 2008;Renaut et al. 2009). Regulatory factors influ-encing the expression of cold-responsive genesare the cold-induced CBF (C-repeat binding fac-tor) transcription factors. The Arabidopsis CBF1gene has been shown to increase freezing toler-ance in several species, including potato (Pinoet al. 2008). Changes at the biochemical levelsuggest, among others, a role for proline and forcarbohydrates, both increasing internal osmoticpressure and preventing cellular water loss dur-ing dehydration (Pino et al. 2008).




Cultivated potato is generally frost-sensitiveand unable to cold acclimate, i.e., to evolve andadapt to cold by increasing its tolerance (Oufiret al. 2008). Some wild potato species are morefrost hardy and capable of cold acclimation. Theythus are an important resource for breeding freez-ing tolerance traits into S. tuberosum. S. com-mersomnii, a diploid tuber-bearing potato, cancold acclimate and survives temperatures downto −12◦C (Pennycoke et al. 2008).

Potato biodiversity—sourcesfor abiotic stress tolerance

The region from the southwestern United Statesto central Argentina and Chile represents the cen-ter of diversity for Solanum species. Potato wasdomesticated over 7000 years before present inthe Andes of South America. During its world-wide adaptation, potato passed through a geneticbottleneck resulting in a relatively narrow ge-netic base of the present varieties. In spite of theefforts to broaden the gene pool that have startedafter the late blight epidemic in the nineteenthcentury and continue up to now, the current va-rieties are still strongly interrelated and a fewclones appear repeatedly in their pedigrees. Inconsequence, for many weaknesses, such as sus-ceptibility to abiotic stresses, there is no effectivegenetic solution available in the breeding pool.Nevertheless, some variation in drought and heattolerance in modern potato cultivars has beenreported (Levy 1986, 1991; Stark et al. 1988;Midmore and Prange 1991; Ekanayake andMidmore 1992; Ranalli et al. 1997; Minie andDe Ronde 2008). In addition to the toler-ance found in varieties and breeding clones,in contrast to most other crops, many potatolandraces and wild relatives are available forbreeding.

Solanum species evolved under a range ofdifferent conditions; they can be found betweensea level and 4500 m altitude, in temperate en-vironments, humid tropical climates, and evenin deserts. Consequently, Solanum species of-fer a vast diversity of tolerance traits for breed-

ing (Mendoza and Estrada 1979; Smillie et al.1983; Reynolds and Ewing 1986; Ochoa 1998;Schafleitner et al. 2007a, 2007b). Up to now,only a small sample of this biodiversity has beenused in breeding programs mainly as a source ofdisease resistance genes.

The potato germplasm pool can be di-vided into the primary gene pool consistingof tetraploid bred potato, the secondary genepool comprising diploid to pentaploid landracesof cultivated relatives (S. tuberosum cultivar-groups), and the tertiary gene pool, the wildSolanum species, comprising accessions withploidy levels from diploid (2n = 24) to hexaploid(6n = 72) (Hijmans et al. 2007). Accord-ing to Huaman and Spooner (2002), all lan-drace populations of cultivated potato, previ-ously treated as distinct species, are, like themodern potato cultivars, members of S. tubero-sum, but can be divided into eight CultivarGroups: Ajanhuiri Group, Andigenum Group,Chaucha Group, Chilotanum Group, CurtilobumGroup, Juzepczukii Group, Phureja Group, andStenotomum Group. The diploid landrace culti-var groups Phureja and Stenotomum groups havebeen used widely in breeding programs. Wildtuber-bearing Solanum L. species are relatives ofthe cultivated potato and comprise 196 species inSolanum sect. Petota, and three in Solanum sect.Etuberosum (Spooner and Hijmans 2001). Muchof the potato biodiversity including wild species,cultivated landraces, as well as thousands of cul-tivars, breeding, and genetic stocks from nearly200 years of modern breeding is maintained ingene banks. The world’s largest potato collectioncomprising more than 11,000 accessions of 8 cul-tivated and 142 wild species is held in trust by theInternational Potato Center (CIP, Lima, Peru).Other important potato gene banks are the Com-monwealth Potato Collection (CPC, Dundee,Scotland), the Dutch-German Potato Collec-tion (CGN, Wageningen, The Netherlands),the Groß Lusewitz Potato Collection (JuliusKuhn Institut, Groß Lusewitz, Germany), thePotato Collection of the Vavilov Institute (VIR,St Petersburg, Russia), the US Potato Genebank




(NRSP-6, Sturgeon Bay, USA), and Potato Col-lections in Argentina, Bolivia, and Peru.

Breeding for abiotic stresstolerance in potato

Past potato breeding mainly focused on highyield, yield stability, product uniformity, tuberquality, and resistance to pests and pathogensand was less oriented toward abiotic stress tol-erance, as the latter has not been recognizedas primary breeding aim. Nevertheless, severalbreeding programs yielded cultivars with in-creased tolerance to high temperatures (reviewedby Levy and Veilleux 2007). Present variation inabiotic stress tolerance in the potato primary, sec-ondary, and tertiary gene pool could be used tobreed varieties that are adapted to higher tem-peratures and can support drought periods betterthan present cultivars.

Intercrossing different Solanum species maybe complicated due to differences in ploidy lev-els, but several natural and artificial mechanismsare available to circumvent this crossing barrier(Hougas and Peloquin 1960; Ortiz 1998). A ma-jor obstacle in using wild species for breedingis the number of backcrosses that are requiredto reestablish the phenotype of a cultivar. It tookfrom three to seven backcrosses to transfer a ma-jor dominant resistance gene from a wild speciesinto a successful cultivar (Bradshaw et al. 2006)and it probably would take many generationsmore to reconstitute the phenotype of a com-mercial cultivar after introducing a highly multi-genic and epistatic trait from wild into cultivatedpotato, except selective markers for the traits ofinterest would be available. Finally, partial orcomplete pollen sterility remains a problem forbreeding programs.

In general, breeding for abiotic stress is dif-ficult for two reasons: the erratic nature of cli-mate stress effects and the complexity of plantabiotic stress tolerance. Nature and strength ofabiotic stress events are highly variable and theirduration may change from year to year makingselection for stress tolerance under natural envi-

ronments difficult, as different stress levels overseasons might activate different stress responsemechanisms. Accordingly, in each selection cy-cle, different genes would be selected for, result-ing in no breeding progress over time.

Most tolerance traits are encoded by a multi-tude of interacting genes. Tolerance traits fur-thermore are subject of strong gene by envi-ronment interaction and thus might not func-tion in the expected way in a broad range ofenvironments. Therefore, heritability of abioticstress tolerance traits is often low and a spe-cific trait might only function in a specific stressenvironment.

For the efficient use of the Solanum biodi-versity to improve abiotic stress tolerance ofmodern cultivars, more information on tolerancetraits present in the wild species and landracesis required, implicating systematic screening ofSolanum germplasm and identification of poten-tially useful traits. Presence or absence of toler-ance traits has been predicted based on taxon-omy, where traits have been linked to particularspecies that are adapted to particular ecologicalconditions (Hawkes 1990; Ochoa 2004). For ex-ample, tolerance to drought, cold, or heat mightbe likely in species or populations growing inhot and dry or cold areas (Rick 1973; Nevoet al. 1982). Hijmans et al. (2007) describeda Geographical Information System-based ap-proach to analyze the extent to which taxo-nomic, geographic, and ecological factors canpredict the presence of frost tolerance in wildpotatoes. Extending this research to heat anddrought tolerance of Solanum could help to focusscreening and trait capture efforts on a reason-able number of accessions instead of characteriz-ing whole collections. Careful analysis of envi-ronmental constrains together with scrupulousassessment of the potential of different stresstolerance traits and trait combinations for yieldmaintenance under stress in the scope of pre-breeding trials and simulation modeling is prob-ably the best avenue to identify those traits thatfinally can improve crop performance in stressfulenvironments.




Genomics has emerged as a powerful toolfor understanding genome structure and genefunctions. It provides new means to unveil ge-netic variation of traits of interest for breeders.Extensive expressed sequence tags (EST) col-lections have put the basis for the developmentof microarrays to study gene expression changesduring developmental processes or responses tovarious stresses in potato (Rensink et al. 2005;Kloosterman et al. 2008). A big step ahead forpotato genomics will be the availability of thewhole genome sequence, which will greatly im-prove the understanding of the gene content ofpotato and give way to analyze linkage phase andepistatic interactions in this heterozygous poly-ploid crop. Now, next-generation sequencing andhigh-throughput genotyping can be applied tounderstand the diversity of Solanum species andto assess links between genotypes and pheno-types on many loci at the same time to identifyand track favorable alleles and the interactionbetween them for potato improvement.

Gene technologies can offer alternative andcomplementary paths for adaptation of potatoto changing environments. Resistance genes notpresent in the potato gene pool could be in-troduced by transgenic technologies into breed-ing populations. Cis-genic approaches using ex-clusively potato sequences could be applied tobring in genes from exotic potato gene pools intobreeding populations without the need of labo-rious backcrossing (Conner et al. 2007). In spiteof the in-general polygenic inheritance of abioticstress tolerance, tolerance traits not necessarilyare encoded by a multitude of genes. Reactiveoxygen species detoxification under stress maybe enhanced by the transgenic expression of oneor a few genes in potato (Van der Mescht et al.2007; Tang et al. 2008). Furthermore, transgenicintroduction of a regulative gene such as a tran-scription factor or signaling gene might influencethe expression of dozens of downstream actinggenes. Thus, transgenic expression of one or afew genes might regulate a complex polygenictrait and could serve to improve abiotic stresstolerance in potato.

Conclusions

The potato crop is sensitive to heat, cold, anddrought; consequently, climate change is pro-jected to have a large, and in many cases, negativeimpact on future yields in many potato grow-ing regions. Breeding for increased stress tol-erance is needed to maintain potato productionlevels face to rising temperatures and increaseddrought stress risk. Potato breeders have accessto a wealth of genetic resources for potato im-provement, such as a vast diversity of landracesand wild relatives that could be used for abi-otic stress tolerance improvement. The wholepotato genome sequence, genetic and physicalmaps, and high throughput resequencing havethe potential to enhance molecular breeding ef-forts for potato. However, increased investmentin the characterization of tolerance traits, par-ticularly those functioning under combined tem-perature and drought stress and under increasedCO2 levels, will be required to adapt potato tochanging climates.

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Crop Adaptation to Climate Change (Yadav/Crop Adaptation to Climate Change) || Adaptation of the Potato Crop to Changing Climates