Visiting Potato from a Breeding Perspective: Accomplishments and Prospects

*Navjot Singh Brar, Sat Pal Sharma and Prashant Kaushik*

#### **Abstract**

Several enhancements to the conventional potato breeding are possible though they have encouragement as well as limitations. I n this direction, the markerassisted selection may be utilized to stack major genes as well as QTLs. Whereas the genetic transformation and genome editing methods accelerate the process of ricking of genes/transgenes. Moreover, these methodologies supplemented with the next-generation sequencing (NGS) platforms and pipelines further aid in reaching the potato ideotype. Here, we overviewed the critical topics that are related to potatoes, from general background, breeding behavior, breeding approaches employed to the potato improvement. Overall, this information complied might serve as background information that is important for potato breeders.

**Keywords:** Potato*,* varieties, heterosis, heterosis, polyploidy, wild relatives

#### **1. Introduction**

Potato is among the most important food staples that rank overall fourth after cereals (maize, wheat and rice), belonging to one of the largest genus Solanum (over 1500 species) of family Solanaceae [1, 2]. Solanaceae family comprises of about 90 genera consisting of 3000–4000 species. Potato offers a considerable component of the world's food source. From unknown until the sixteenth century in the six following centuries, potato cultivation had spread from its centre of origin, in South America into the rest of the world [3, 4]. The genetic diversity is harboured in wild relatives and landraces considered to be valuable sources of deviation for genetic enhancement and crop improvement because the genetic foundation of the modern cultivated potato is quite narrow [5]. At present, the collected developed to guarantee the long-term upkeep of potato hereditary resources and reaffirms the benefits of potato genetic resources [3]. Collaboration between potato researchers and gene bank curators promotes the utilization of the genetic resources [6].

Moreover, there are over 5000 cultivar varieties of potato-based on its size, color, shape, texture, flavor, taste, storage quality and cooking quality [7]. These varieties are differing in physiochemical properties (carotenoids and ascorbic acid content) because of the location, agronomy practices, climatic and degree of stress conditions of that area [8, 9]. The potato crop is affordable (the poorest and most undernourished households can afford), high in quality nutrients (potential high food security crop), matures rapidly (4–6 months), need moderate care (irrigation at an interval of 6–7 days), easy to cook, protect itself against microbes (impermeable to gases,

water and chemicals), feed entire populations from hunger (high food security crop), easy to digest (quick breakdown with high glycemic index), used for some byproducts production (starch and alcohol) and also consumed by the animal as fodder [10]. Potato has high dietary fibers, magnesium, manganese, potassium, phosphorus, amino acids, proteins, carbohydrates, minerals, moisture, starch content, vitamins (Vit-C, B6) as well as other antioxidants like polyphenols and carotenoids and low in fats [11, 12].

Worldwide, economic losses occur in potato because of diseases like late blight although these diseases are controlled by regular application of fungicides [13, 14]. Recent improvement in next-generation sequencing (NGS) technologies has resulted in a major reduction in the sequencing costs that makes genotyping with NGS systems cheaper and achievable [15]. Massive genotyping of the gene bank collections as well as posting the info will be a strategy to show the prospective utilization of germplasm collections in gene banks. Some gene banks have started distribution of germplasm collections together with all the genotyping information by NGS datasets [16]. NGS technologies are particularly helpful in the taxonomy that depends considerably on the herbarium specimen conceived from wild plants from the wild [17]. In this review, we have gathered the information from the general background, breeding behaviour, conventional breeding, genetic engineering to NGS methodologies employed to the potato improvement. This information is going to be a useful resource for potato breeders, offering information about the development made and prospects of reading a potato ideotype.

#### **2. Taxonomy**


The section Petota is splits in to 8 cultivated and 228 wild species of potato, which are further grouped into 21 taxonomic series (19 tuber bearing+2 nontuberous) [22]. Out of the cultivated species, only *S. tuberosum* ssp. *tuberosum* is extensively cultivated all around the globe, while others are cultivated especially in the Andean nations*.*


#### **3. Origin and evolution**

Series *tuberosa* (containing *S. tuberosum*) and other series of subsection *potatoe* have two centres of diversity. One is a long-stretching Andean terrain in Argentina, Bolivia, Colombia, Ecuador, Peru, and Venezuela, while the other is in central Mexico [18, 23–31]. This theory is based on the fact that the plants originally introduced into Europe were late flowering and tuberising, and the morphological description [32]. Such transition can take place in a fairly short period of approximately ten years of selection [33]. An alternative school of thought is that, after the potato blight epidemic in Europe, new germplasm of *S. tuberosum subsp. tuberosum, which* originated from Chile was introduced into Europe [18].

Hawkes [18] and Grun [34] opined that the cultigenic species *S. stenotomum* is the most primitive and progenitor of all other cultivated material. *S. leptophyes* Bitt. has been theorized as the probable progenitor of *S. stenotomum* based on morphological similarity [18]. The first cultivated material of *S. stenotomum*, has also been considered to be domesticated from *S. brevicaule* complex genepool [25, 34–37]. With advent of molecular techniques seven different chloroplast haplotypes were distinguished in a selection of wild and cultivated species [38]. Kardolus et al. [39] revealed that *S. tuberosum* subsp. *tuberosum* forms a cluster with *S. multidissectum* and *S. canasense* in the Brevicaule complex. *S. tuberosum* is believed to be a straight tetraploid of *S. stenotomum* by some workers but some evidence strongly support the allotetraploid origin of *S. tuberosum* [40]. As per another report the cultivated species are in the same clade as the northern Brevicaule clade that consists of *S. bukasovii*, *S. ambosinum*, *S. canasense*, *S. leptophyes*, *S. achacachense* Card. and *S. multidissectum* [41]. Multiple origin from *S. stenotomum* is believed to be cause of rising of initial populations of *S. tuberosum* subsp. *andigena* [42]*.*

To summarize, first diploid cultivated material (*S. stenotomum*) has probably descended from one of the species in the Brevicaule complex. Sexual polyploidization, accompanied by hybridization and human selection led to the development of tetraploid landraces (*S. tuberosum* subsp. *andigenum*). However, there is an absence of sufficient molecular data to point out a particular wild ancestral species.

#### **4. Domestication of potato**

Spanish conquerors introduced potato into the European countries by the 16th century [18, 43–46]. There are two competing theories about the nature of the first material to be introduced into Europe. Grun [34] and Hawkes [18] suggested the very first potato material brought to Europe consisted of *S. tuberosum* subsp. *andigena* from the Andes, quite probably from Colombia. The late blight epidemic in Europe during the 1840s led to the destruction of most of the original stock of potato. In the post epidemic period, new introductions consisted mainly of *S. tuberosum* subsp. *tuberosum.* Whereas Juzepczuk and Bukasov [47] were of the opinion that the subsp. *tuberosum* germplasm from Chile was already a part of early introductions in Europe, as morphology and growing conditions of early European plants and Chilean material bore similarities. Chilean potatoes were suitable for growing in Europe as they were adapted for tuberization under longday conditions. DNA analysis of the historical herbarium specimens suggested that although Andean potato arrived first but Chilean potato was present long before late blight epidemics in Europe [44].

Introduction of potato to the Bengal floodplains, Nile delta, Morocco and Nigeria was made by European colonizers, colonial governors, missionaries [48, 49].

Emigrant farmers carried the potato to Australia and South America that led to the establishment of the potato in Argentina and Brazil. The tuber spread was along the old Asian routes through the Caucasus to Turkey, and from Russian federation to western China [31].

During the 20th century, potato emerged as a truly global food. After the Second World War, the potato was grown on a huge span of arable land in Germany and Britain, and potato has surpassed cereal production in Belarus and Poland. Since 1960s, cultivation of potato has been expanding in the ever-developing world [50, 51], it is grown as a cash crop in Bangladesh.

#### **5. Floral biology**

Potato inflorescence is terminal comprising 1–30 (but usually 7–15) flowers, depending on the type of cultivar [52–55]. The inflorescence is cymose, and flowers are actinomorphic and hypogynous. Arrangement of floral parts is regular. Five petal arrangement of the flower gives it a star shape [56]. Depending upon the cultivar, shape and size of lobes of sepals vary. The androecium comprises of five stamens alternating with the petals. The anthers collectively form a cone shaped structure to conceal the ovary [55]. Anthers are bright yellow or orange coloured except in case of male sterile plants in which the colour of anthers is light yellow or yellow green [57]. The ovary is superior and bilocular with ovules arranged at the periphery of the placenta.


Details of the *S. tuberosum* inflorescence are given below:

Colour of the corolla varies from white to complex range of blue, red, and purple [53]. Opening of flowers start near the base of the inflorescence and proceed upward at the rate of about 2–3 flowers each day [54]. Long day length accompanied by high humidity and low temperature are conducive for potato flowering [57, 58]. Flower production and berry setting is favoured by 12–14 hour photoperiod and night temperature of 12-15°C [59, 60]. Short day duration at the time of flowering may result in abscission of floral bud [58]. Flower and fruit production in potato is influenced by several factors such as genotype, temperature, photoperiod, inflorescence position, plant/stem density, competition between flower and tuber, precipitation, date of planting and nutrient level [61–65]. Flowers remain open for 2–4 days, and out of this duration, pollen production and stigma remains receptive for about 2 days [57]. The fruit type is a berry, and are spherical to ovoid in shape, about 14 cm in diameter. Berries are green in colour or green-tinged, and upon ripening bear white or purple spots or bands [53, 66].

*Visiting Potato from a Breeding Perspective: Accomplishments and Prospects DOI: http://dx.doi.org/10.5772/intechopen.98519*

Floral bud abscission occurs in case of short days at the time of flowering, hence giving the impression of poor flowering of a cultivar [58]. Thus, conditions favourable for flowering and fruiting in tropics and subtropics can be found at higher altitudes (1500 m above sea level) [67]. Characteristics like days to flowering, flowering duration, the intensity of flowering and fruit set have wide genetic diversity [60]. A survey on flowering behaviour, male sterility and berry set was conducted across 25 countries by Gopal [67]. Flowering initiated after 6–15 weeks of planting and duration of flowering ranged from 1 to 10 weeks. The setting of berries ranged from 0 to more than 5 berries/plant, while there no setting in 31.8% of accessions in blooming. Production of flowers and fruits is influenced by several factors like temperature, photoperiod, genotype, inflorescence position, plant/stem density, flower and tuber competition, precipitation, date of planting and nutrient level [61–65]. The number of primary flowers increased with increase in plant density while the proportion of flowers on lateral stems reduced [62].

#### **6. Pollination**

Potato is predominantly a self-pollinated plant and is occasionally crosspollinated [54, 56]. Generally, diploid wild species are insect-pollinated and crossbreeding in nature. Presence of insects is imperative in facilitating cross-breeding and selfing in potato. Bumblebee species like *Bombus terricola* and *B. impatiens* are particularly good pollinators for potatoes [68, 69]. European honey bee (*Apis mellifera*) and *B. fervidus* do not contribute to the pollination of potato, as the flowers are devoid of nectar [70]. Despite the lack of pollinator resources provided by the crop, a great diversity of bees was recorded in a potato-dominated agroecosystem [71]. Wind does not play any role in the pollination of potato, and no seed set was observed [68]. There are no detailed studies of pollination behavior of potato in India. Controlled pollination can be achieved under field or greenhouse. However, crosses made under the field conditions are prone to losses from the environmental factors like wind, rain, heat and drought. Therefore, breeders prefer crossing in the greenhouse. The crossing should preferably be done during the early morning hours when the temperature is moderate [54].

#### **7. Wild relatives of potato**

Comprehensive taxonomic treatment by Hawkes [18] found there are 235 potato species in total, 228 outdoors and 7 cultivated potato species. Various studies, implementing advanced molecular resources with a considerable amount of samples covering a broad range of species have advised that a reconsideration of the taxonomic classification is necessary [72]. As previously, potato species are hugely sophisticated in taxonomic classification. A broad area of distribution, together with an extensive selection of altitudinal division, from sea level up to 4500 MSL, indicates a broad range of adaptation this has resulted in the huge diversity and adaptations in the potatoes [73].

Genetic diversity of the germplasm and usefulness has been the drive to incorporate wild genes into cultivated types. The achievements of the application of wild relatives for genetic improvement relies a great deal on crossability with developed species. The gene pool is essentially the most often used concept determining the level of relatedness between species [74]. Though the genepool concept has been generally accepted, efforts to utilize the genepool concept to the potato was also

presented [75]. Manipulation methods to alter the ploidy level in potato have been discovered. Even important genes from the tertiary genepool could be unveiled using bridge species in the crossing, embryo rescue, and somatic hybridization [76]. Currently, potato genetic materials are preserved in gene banks around the planet and therefore, are offered for potato breeders as well as researchers [77]. Cultivated potatoes are conserved primarily as clonal collections, like a tuber, *in vitro* and cryopreservation; on the flip side, wild potato species are primarily gathered up and also retained in the type of botanical seeds [78, 79].

#### **8. Fertility issues in potato breeding**

Potato is propagated sexually by seeds and asexually by tubers [80]. Most of *Solanum* species are diploid in nature with obligate allogamy (cross-pollination) which is result of multi-allelic gametophytic self-incompatibility (S) locus, thus preventing self-fertilization among Solanum species. In contrast to this, tetraploid cultivated potato (*Solanum tuberosum* ssp. *tuberosum* L.) [81]. However, their highly heterozygous nature (interlocus and intralocus) with tetrasomic inheritance pose difficulty in genetic complexity and challenge in potato breeding, and this is further aggravated by, high genetic load due to accumulation of deleterious alleles as a result of its vegetative propagation. Severe inbreeding depression is anticipated upon selfing, which results in the reduction of seedling germination and many reproductive complexities like reduction in flowering [57].

Conventionally potato varieties are developed through hybridization and selection, with a huge investment of time and resources because of its complex multi-locus inheritance and tetraploid genome. Successful hybridization programme between different potato populations have to deal with many barriers like pre-zygotic barriers including pollen and pistil incompatibility and post-zygotic barriers like embryo and endosperm abortion, sterility and hybrid breakdown in segregating generations [82], that leads to the hindering of the breeding programmes [60]. In male-sterile plants, flowers do not produce functional anthers or viable pollen, but the ovaries usually function [57]. The failure to produce pollen may be an inherent characteristic with sterility being dominant over fertility [83]. Even after successful fertilization by overcoming these issues, development of seeds requires proper endosperm development.

Male sterility is the result of nuclear cytoplasm interactions; the predominant Ms. gene interacts with the cytoplasm, for instance, the diploid hybrids between *S. tuberosum* Group Tuberosum haploids × Group Phureja yield all or perhaps nearly all-male sterile progeny [84]. The occurrence of male sterility typically leads to issues for potato breeders, as the option of parental lines can limit the introgression of characteristics [85]. The frequency of male fertile offspring in a hybrid between the group Tuberosum and Phureja are different because of their different ploidy levels [86–88].

In the last couple of years, a pattern emerged in a group of potato breeders to reconsider the pick as a diploid species constructed from a compilation of inbred lines that capture the favourable genetic diversity accessible in cultivated and wild potatoes [89]. Inbreeding due to selfing might be useful for organizing the entire gene pool into different favourably interacting and healthy epistatic systems. Whatever the nature of its, self-compatible 2x cultivars will offer an even more appealing self-compatible source than *S. chacoense* since they will avoid the undesirable linkage drag regarding the usage of an untamed species within the development of 2x inbred lines. Loss of S-RNase functionality is a standard route to self-compatibility [90].

### **9. Unilateral compatibility**

The endosperm balance number (EBN) seems to be very likely that a mechanism related to a loss of protein functionality results in the formation of 2n gametes. Although it is not complete, the consistency of the self-incompatible self-compatible rule indicates a link between inter- and intraspecific pollen rejection [91, 92]. The EBN concept was helpful to elucidate the nature of the pollinator result in haploid removal. The triploid block is a reproductive screen resulting from endosperm malfunction due to the epigenetic event of genomic imprinting. Evidence implies that the endosperm dosage devices are imprinted within the gametes; therefore, the similar gene being functionally different in paternal and maternal chromosomes [93].

Spooner et al. [22] proposed a concept particularly for the Potato, implementing 5 crossability groups based on self-compatible/self-incompatible systems and endosperm balance number (EBN). The main genepool of potato contains *S. tuberosum* ssp. *tuberosum* with all cultivars and landraces. All the cultivated potatoes are tetraploid (2n = 4x = forty eight) with 4EBN. Potato has a vast secondary gene pool comprising of related wild species that gives a rich, distinctive, and different supply of hereditary variation. The EBN is a unit identifying the realizations of interspecific crosses [94]. Hybridization within every group is anticipated to achieve success rather than hybridization across groups, and therefore the executions of hybridization may be predicted. Whereas the genepool principle, as well as the EBN model, provides assistance in the utilization of wild genetic resources, additionally, they provide insight into phylogenetic connection and also taxonomy. Nevertheless, species crossability are always crucial to offer concrete evidence. Potato researchers have developed strategies to conquer the hybridization screen to transfer genes from wild species of the secondary and even tertiary genepool [95].

Haploids exhibited disomic inheritance, that implies that every chromosome combined with its homolog, thus giving means for simplifying genetic research in potato. They can furthermore be well utilized for research on natural mutation and chromosome pairing accumulated at the tetraploid fitness level. In this direction, the reason behind the generation of haploids was acquiring a genetic bridge between the different genomes of Solanum species [96]. Haploids from tetraploids usually don't flower and can also be male sterile because of inbreeding throughout the tasks of haploidization [97]. Selection of haploids can result in diploid breeding lines; additionally, a particular kind of haploids are accustomed for understanding the segregation of characteristics at the tetraploid level if numerous haploids are made of a single tetraploid genotype [98]. Whereas, tuberization in potato is controlled by day length [99, 100], and plant hormones, such as gibberellin and jasmonic acid also play a crucial role in defining tuberization. Although, specific potato genotype tuberizes under a particular day length condition along with specific physiological requirements that vary from genotype to genotype. In *in vitro* studies, no particular method of tuberization is found, and it's regarded as a complex trait. Utilizing the genome sequence [101] as well as info on Ft, it was determined that the potato genomic locus StSP6A, induces movable tuberization signal. The StSP6A signal led to the induction of tuber growth at the stolon termini. They've postulated that diverse allelic deviation of this gene is connected with the domestication of potatoes.

#### **10. Breeding behavior of potato: from conventional to new breeding technologies**

Potato breeding and improvement is an uphill task owing to its complex genetic structure and multi-allelic gene action arising due to its tetraploid genome [102].

Any breeding programme relies on the objective of the programme, germplasm availability and breeding method/technique. Genetic resources of potato are quite rich as compared to any other cultivated plant consisting of about 190 wild and primitive species [103], resulting in great amount of genetic diversity readily available for exploitation. Its rich variations are also attributed due to its reproductive biology which shows there can be 40% (range 21–74%) natural cross-pollination [104]. Besides this, its tetraploid nature (2n = 4x = 48) having four sets of chromosomes entirely homologous shows random pairing at meiosis [57] further adding to its diversity and genetic variations. This sexual reproduction generates ample amount of diversity by recombining the variants of genes that arose by mutation. As a consequence, potatoes are highly heterozygous individuals that display inbreeding depression on selfing and thus become the major impediment for the exploitation of its heterosis [105, 106].

Despite the broad genetic base, progress in efforts for potato breeding is quite slow, and its genetic gains are not fixable due to the obligatory out-breeding nature. Several conventional, as well as modern breeding techniques, have been utilized for improvement in yield, processing, storage-quality [107] and against biotic stresses [108, 109]. Although conventional breeding approaches like hybridization, clonal selection, irradiation/mutagens and introgression has been successfully employed [57]. But the progress is limited and slower due to demanding tasks of introgression and phenotypic characterization of better performing individuals in successive generations. Apart from this, intraspecific incompatibilities and inbreeding depression lead to failure of trait incorporations in the polyploid crop.

Although conventional breeding has played an important role in potato improvement by developing coloured potatoes and potatoes with improved nutrients [110], but the progress is very slow. In order to overcome these challenges, biotechnological, molecular breeding and genome editing tools, considered as new breeding techniques, have played an important role to facilitate interspecies crosses, and towards augmenting and broadening of the genetic base of gene pool of cultivated material. Biotechnological techniques like *in vitro* meristem shoot tips culture have been successfully eliminated potato virus Y [111]. This method was crucial and reliable for supplying pathogen-free seed potatoes to farm [112]. Embryo culture technique has been used successfully for improving resistance to potato leafroll virus so as to circumvent interspecific incompatibility [113]. Utilization of somaclonal variation resulting heritable phenotypic changes arise during the cell culture and regeneration of potato tissue culture was reported from leaf protoplasts of 'Russet Burbank' cultivar [114] along with improved resistance to pathogens like *Phytophthora infestans*; *Alternaria solani* [114, 115] and tuber morphology [116]. The somatic fusion of potato protoplasts with protoplasts of wild relatives has also been extensively exploited for introgression of novel sources of disease and pest resistance [105, 109, 117–119].

Potato is a model crop in which transgenic or genetic engineering technology has been exploited to the maximum extent, and it is one of the first crops for which transgenic plants were regenerated [120]. Genetic engineering is an important and highly effective tool for incorporating single gene or pyramiding gene into elite potato cultivars with minimal or no disturbances to their genetic background [121]. Numerous transgenic genotypes have been developed for a wide range of traits, including pest and disease resistances; abiotic stress resistance; quality attributes for improved processing, nutrition and appearance etc. Gene silencing is another novel technique which uses RNAi for traits like increased carotenoid content and reducing cold-induced sweetening [122–124].

Apart from transgenics/genetic engineering techniques, marker-assisted breeding (MAB) has been successfully demonstrated in tetraploid potato [125] *Visiting Potato from a Breeding Perspective: Accomplishments and Prospects DOI: http://dx.doi.org/10.5772/intechopen.98519*

for potato cyst nematode resistance trait. Several other examples like resistance to the nematode *Globodera rostochiensis*, resistance to potato virus X and resistance to potato wart [126] are the success stories of the application of MAB in potato. But the progress in MAB is negligible as compared to other crops due to its complex tetrasomic inheritance and high allelic variation [127]. However, in the current era of genomic breeding, prediction of genomic information is the best method to use for making breeding decisions [128]. Rather than using only significant marker-trait associations to build a prediction version, genomic prediction makes simultaneous usage of all markers [129]. In potato, genomic selection (GS) models are being utilized for predicting the accuracies of prediction models for various traits like for maturity [130], tuber starch content and chipping quality [131], *Phytophthora infestans* infection, plant maturity, tuber starch yield and tuber yield have been successfully predicted using GS models [132].

For the successful application of genome editing technologies, the first and foremost requirement is the availability of efficient transformation systems. Since potato has excellent availability of genomic resources as well as genome sequence and efficient transformation systems, several workers used various genome editing approaches *viz.* zinc-finger nucleases (ZFNs) [133–137] for improving traits like herbicide resistance, modification of starch, bio-fortification and reducing anti-nutritional factors to enhance overall increased quality of produce. Earlier for targeting traits like insect resistance, proteins, vitamins and carotenoids, transgenic technology was extensively used. Still, due to their off-target, copy number variations and other drawbacks, the trend has been shifted towards these new breeding technologies whereby TALENs and more recently CRISPR/CAS9 genome editing technologies were used for targeting traits like alteration of starch composition or hormonal expressions, reduction of anti-nutritional elements, imparting herbicide resistance, improving starch quality and overcoming self-incompatibility issues.

#### **11. Conclusions and future prospects**

The genetic improvement of potato depends on germplasm sources. In the genomics era, germplasm development can be easily performed by incorporating noval alleles from wild species, landraces, cultivated varieties, and even from distantly related species. In corporating the genomics equipment will substantially enhance the effectiveness of introgressing multi genic characteristics. Introgression may be possible through sexual hybridization, or molecular manipulations. In the context of molecular manipulations, different breeding technologies as TALEN and CRISPR/Cas9 are already used to improve the potato ideotype as per the market requirements. Moreover, the potato genome sequence, as well as useful potato hereditary transformation strategies, have hugely facilitated potato genetic engineering. The commercialization of these engineered goods is challenging because of regulatory/ethical restrictions and consumer preferences.

Breeding objectives like bio-fortification, as well as the removal of antinutritional factors like steroidal glycoalkaloids as already achieved. Additionally, incorporation of abiotic (environmental, salinity, drought, temperature) anxiety resistance that comes with improved nutrition can facilitate potato to acclimatize in varied agro-ecological zones, therefore impeding food shortage in less fertile/ water deficit farming lands. Further expansion of food studies can establish several preliminary values to rationalize the health advantages of potato derived foods. Indeed, the potato genome sequence has facilitated the relative genomic analyses to determine the genes helpful for improving several agronomically significant characteristics as tuberization, damage of bitterness, along with ailments opposition.

Whereas, the studies concentrating on food safety and protection can offer considerable means to meet up the soaring food demands, particularly in the food-deficit countries. The rapid advancement of growing genetic engineering has supplied brand new exciting resources to produce crops with nutritional traits and enhanced yield. In this particular context, potato harvest has potential that is enormous to help with food security as it can offer inexpensive, energy food that is high at a sustainable basis.

## **Conflicts of interest**

The authors declare no conflict of interest.

## **Funding**

This research received no external funding.

## **Author details**

Navjot Singh Brar1 , Sat Pal Sharma1 and Prashant Kaushik<sup>2</sup> \*

1 Department of Vegetable Science, Punjab Agricultural University, Ludhiana, India

2 Instituto de Conservación y Mejora de la Agrodiversidad Valenciana, Universitat Politècnica de València, Valencia, Spain

\*Address all correspondence to: prakau@doctor.upv.es

© 2021 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

*Visiting Potato from a Breeding Perspective: Accomplishments and Prospects DOI: http://dx.doi.org/10.5772/intechopen.98519*

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#### **Chapter 11**

## Impact and Management of Diseases of *Solanum tuberosum*

*Olusola L. Oyesola, Oluwadurotimi S. Aworunse, Margaret I. Oniha, Onyemaechi H. Obiazikwor, Oluwakemi Bello, Olubunmi M. Atolagbe, Ayodele A. Sobowale, Jacob O. Popoola and Olawole O. Obembe*

#### **Abstract**

*Solanum tuberosum* (Potato) is one of the essential economic crops with the potential to reduce hunger due to its high yield per unit area of land compared with many economic crops. However, its yield losses due to pest and disease attacks could be as high as 100%, depending on its tolerance level and pest and disease. Over the years, several disease management strategies have been researched, ranging from synthetic pesticides to the formulation of biopesticides as disease control measures. Moreso, recent breakthroughs in genetic engineering have simplified plant disease management strategies by developing techniques for conferring resistance on plants. Potato is a vital food crop worldwide, and with the struggle to suppress world food insecurity, effective disease management strategies must be employed for high production of quality and quantity potato, enough to feed the ever-increasing world population. Therefore, attention must be given to how disease-free potatoes can be produced to meet the unending demand for food by the continually increasing world population.

**Keywords:** Potato, Disease-free, Crops, Pathogens, Biocontrol, Resistance

#### **1. Introduction**

Potato (*Solanum tuberosum* L.) is the most popular vegetable crop of great importance worldwide and follows only wheat and rice as a food crop [1]. It is a source of carbohydrate being a starchy vegetable; it is, however, as a vegetable a very significant source of vitamin C, potassium, and dietary fibre as well as magnesium, vitamin B6, iron, carotenoids, and phenolic acids [2, 3]. It grows in a wide range of climates and is adopted by a broad range of cultures [4]. Potato is a critical alternative to the major cereal crops for feeding the world's population [5]. However, its production has two main challenges: disease and nutrient management [6]. Pathogens such as bacteria, fungi, viruses, nematodes, and phytoplasmas attack potato plants, causing diseases, which result in a significant loss of yield [7]. Naming a pathogen that negatively affects the host's health is the primary means to define any disease [8]. For instance, potato plays host to heterothallic species, *Phytophthora infestans* [9], which causes late blight disease. This single pathogen

caused severe devastation in the late 1840s in Europe [10] and cost Ireland 25% of its population in just four years [11]. Potatoes still have many diseases, but many other alternative crops make most countries not depend on potatoes like Ireland in the 1800s [12]. In recent time, potato crop loss due to late blight disease alone is estimated at \$6.7 billion annually worldwide [5]. A significant challenge to the management of *P. infestans* is the rate at which it adapts to control strategies [5]. More research on epidemiology and the host-pathogen interaction is needed to devise the most appropriate management strategy [7]. Also, insight into pathogen population dynamics offers an essential input for effective disease management [13].

Meanwhile, effective management of the disease requires implementing an integrated disease management approach [14]. Guchi [7] proposed investigating several control options and implementing an integrated management strategy based on local needs [7]. Therefore, this chapter aims to discuss the general/overall impact of the diseases of *Solanum tuberosum* as well as their management. This would increase awareness and awaken researchers' intervention to develop globally effective control or management strategies.

#### **2. Some host pathogens and diseases of potato**

Diverse host-pathogens are associated with the different diseases of potatoes, among which are bacteria and fungi. Plant pathogens responsible for diseases in potatoes include viruses, fungi, oomycetes, and bacteria [15]. A pathogenic bacterium known as *Ralstonia solanacearum* is responsible for the devastating bacterial wilt of potato and other solanaceous plants [16, 17]. The bacterium, *Ralstonia solanacearum,* is a gram-negative, non-spore-forming, aerobic, soil-borne motile pathogen that hinders tuber production resulting in economic losses [16–18]. It is distributed worldwide, affecting more than 200 economically essential crops, including potato [19]. The pathogen, usually disseminated by infected seed tuber, soil, water, and farm machinery [20], penetrates to infect the roots through wounds or natural openings and rapidly propagates within the host to attack the plant's vascular system. Consequently, it forestalls the translocation of nutrients and water, culminating in wilt, collapse and complete deadening of the plant and its decay [21, 22]. The ubiquitous plant pathogenic fungus of *Colletotrichum coccodes* is responsible for the blemish disease of potatoes called black dot [23].

The typical characteristic of black dot disease is the microsclerotia on infected tissue present in all potato parts. These microsclerotia, which usually survive in the soil for lengthy periods, lead to high disease incidence when soil inoculum levels increase [24]. Sequel to fungal colonisation of roots are colonisations in the stems, stolons, and tubers [25], and fungal contamination of tubers with *C. coccodes* leads to the development of lesions on the epicarp and loss of water during storage [26, 27]. The potato late blight disease is caused by *Phytophthora infestans* [28]. It affects the potato foliage and tubers. The foliage symptoms begin with brown to black, water-soaked lesions on leaves and stem that produce visible white spores at the lesion margins under humid conditions. This may result in the rapid collapse of the entire plants and orchards. Sporangia in the soil from the foliage initiate the tuber infection that starts from the wounds, eyes, or lenticels. The lesions appear as copper brown, red, or purplish, and white spores appear on tuber surfaces in storage.

*Streptomyces* spp. is the bacterial pathogen responsible for common scab in potato, and characteristic tan to dark brown, circular or irregular lesions rough in texture are produced. The scab may be superficial (russet scab), slightly raised (erumpent scab), or sunken (pitted scab). Its lesion type is determined by potato cultivar, maturity of tuber at infection, soil organic matter content, pathogen

#### *Impact and Management of Diseases of* Solanum tuberosum *DOI: http://dx.doi.org/10.5772/intechopen.98899*

strain, and the environment [29]. Another disease caused by the bacterium is soft rot, which is the most destructive of all storage diseases caused by *Erwinia carotovora*. The disease symptoms include tan- to brown-coloured water-soaked areas of granular, mushy tissue often outlined by brown to black margins. During storage periods, soft rot bacteria penetrate tubers already infected with other potato diseases. The rottening from bacterial penetration is accelerated by the heat generated from the intense respiration in the storage environment.

Early blight of potatoes, caused by *Alternaria solani*, usually affects its leaves, but tuber infections can also occur. The lesions found in the tubers are dark, sunken, and circular, usually surrounded by purple to grey raised tissue. Its underlying tissues are void of moisture, leathery and these brown lesions may have increased during storage with shrivelled tubers [29]. *Fusarium sambucinum* or *F. coeruleum* is responsible for dry rot that causes inner light to be dark brown or black dry crumbly rot of potato with collapsed tissue often lined with secondary white other-coloured fungal growth. This rot may commence at an injury site (bruise or cut), and the fungus penetrates the tuber to rot out its centre. In furtherance, the extensive rotting results in the shrinking and complete collapse of tissue and usually leaves a dark sunken area outside the tuber and internal cavities [29]. The silver scurf, caused by *Helminthosporium solani*, infects only the uber periderm (skin). The lesions appear first at the stolon end as small pale brown spots that may be difficult to detect at harvest but continues development during storage. While in storage, these lesions darken, sloughing off the skin occurs with many small circular lesions coalescing to form large lesions. The potato tubers tend to dry out and become wrinkled from excessive moisture loss during storage [29]. The fungus *Rhizoctonia solani* causes the black scurf disease, which does not reduce yield, even in storage. Fungal sclerotia develop in irregular, black hard masses on the tuber surface that harvesting tubers may reduce immediately after vine-kill and skin set. Sclerotia allow the pathogens to survive in the soil. Inside wet soils, *R. solani* may induce dark, sunken lesions on underground sprouts and stolons with consequent deprivation of nutrients, the complete killing of the potato tubers, reduction in transfer of starches (results to reduced sizes) [29].

Pink rot infections caused by *Phytophthora erythroseptica* commence at the stolon end and culminates in rotten, internal rubbery skin that turns pink after about 15 to 20 minutes of exposure to warm air (with a clear delineation between healthy and diseased tissue). On exposure to air, the tuber flesh turns pink and then brown-black. The fungal pathogen *Pythium* spp. is responsible for leak infections, penetrates tubers through harvest wounds, and continues to grow in transit and storage. Its infections develop into internal watery, grey, or brown rot, but the outer cortex remains intact, with well-defined red-brown lines demarcating healthy and infected tissue [29].

Viruses are among the predominant phytopathogens that cause approximately 50% of all emerging plant diseases [15]. Potato virus Y (PVY) is one of the most harmful viruses infecting potatoes across the globe since the 1980s [30].

#### **3. The impacts of diseases on the yield (quality and quantity) of potato**

In 2013, more than 368 million tonnes were produced from 19.4 million hectares [31]**.** Though hundreds of varieties of potato are grown in temperate and sub-tropical areas, its diversification in various agroclimatic conditions leads to a decrease in its production and productivity due to its low genetic base and various biotic factors, which makes it susceptible to many devastating diseases. The crop infection due to fungi, viruses, bacteria, and viroids alters its metabolism. These pathogens

affect the crop's morphological, physiological, and biochemical characteristics leading to altered distribution of photoassimilates, with resultant effects on its quality ad quantity.

Viral diseases of potatoes are devastating because they are tough to manage and transmitted via the tubers to subsequent generations. Viruses have the potential to alter the physiology of potato plants drastically, causing disorders. These disorders of growth processes cause stunting, leaf deformation, dwarfing, and reduction in the yield of potato tubers and product quality up to 88% [32–34]. Tens of potato viruses have been discovered and characterised, and the most cataclysmic are: Potato virus M (PVM): Potato virus S (PVS); Potato virus X (PVX); Potato virus Y (PVY); and Potato leaf roll virus (PLRV, virus L). PVX can debilitate 10–40% of potato in a single infections cycle and possess enormous devastating effects when combined with other potato viruses; due to its synergistic interaction with potyviruses, tuber losses yield close to 80% [33]. For example, the yield of potato simultaneously infected with PVM and PVX will decline to 60%, and when it is a complex infection of PVM + PVX + PVY, it will decline by 83.7%, i.e., total loss of yield [35]. In potato tubers infected with viral diseases, the content nutrients become reduced compared to healthy ones. Other biochemical and physiological changes also occur, resulting in a decrease in the quantity and quality of starch grains in the debilitated tissues, the acidity of starch, and amylase content [36]. There are varying losses in potato production from viruses; they are determined by the variety's resistance, the viral pathocomplexity, the level of spread of a specific virus, and their combinations with other viruses [37].

Bacterial diseases are one significant biotic constraint of potato production in the subtropical and tropical regions. Several bacterial diseases devastate potato, resulting in severe damages, especially on tubers, leading to economic losses. The most acute diseases are bacterial wilt caused by *Ralstonia solanacearum* [38] and the backleg caused by *Pectobacterium atrosepticum, P. carotovorum* subsp. *brasiliensis, P. wasabiae, Dickeya solani* and *D. dianthicola* [39, 40]. Loss of yield in potato crop is due to bacterial diseases that could be direct and indirect. There are specific facets: short-term impacts like yield loss and unvendability, and others with long-term impacts with environmental, economic, and social effects [39].

To date, potato late blight is still one of the most devastating diseases in potatoproducing regions worldwide and causes substantial economic losses of about 25–57%. Pathogenic fungus, *Phytophthora infestans,* are responsible for late blight disease in potato. Late blight disease is highly destructive and one of the diseases threatening global food security [41]. Its outbreak in Ireland resulted in famine, which led to millions of people's starvation and eventual death and subsequent continuous significant losses of potatoes worldwide. Therefore, it remains the most debilitating disease of the food crop, which causes annual potato losses sufficient enough to feed several millions of people [42]. Despite the apparent debilitating potential of late blight, it is tough to estimate losses because of other environmental factors that simultaneously affect potato yield.

Meanwhile, the economic impact of potato late blight in the USA was appraised to be around 210 million US dollars, while a worldwide assessment of potato loss by late blight in the second world countries based on an average production was about 15%. This represents approximately 2.75 billion US dollars loss in developing countries. However, a critical method of estimating the economic impact of potato late blight is by determining the usage of fungicide. With this method, the estimated fungicide currently used in developing countries stands at 750 million US Dollar. Therefore, about 1 billion US Dollar is spent on fungicides yearly to manage fungal disease worldwide [43].

#### **4. Management strategies of the diseases of potato**

Potato is among the high-income-yielding crops globally and can contribute to poverty reduction in developing regions [44]. However, Potato cultivation is beset with several diseases caused by diverse pathogens in the field and during storage, accounting for 50 to 60% of annual losses [45–47]. Control strategies that have been deployed to manage diseases in potato include the application of chemical fungicides, biological control agents, and cultural practices involving crop rotation.

#### **4.1 Chemical control**

Diseases caused by fungi are critical in potato production and require several synthetic fungicide options to reduce them to tolerable economic levels. Fungicides are preparations of different organic and inorganic compounds which can inhibit or destroy phytopathogenic fungi. These chemicals exert their effects by disrupting cell membranes of their targets or instigating catalytic enzymes in plant host tissue to suppress fungal growth and proliferation [48]. Practically, conventional management of potato diseases relies on the timely application of preventive fungicides [48, 49]. To control black rot disease, seed tubers are immersed in the fungicides thiabendazole, captofal, chloramizol sulphate, prochloraz, or a combination of each before field planting. Pencycuron and thiabendazole have also been documented to control black scurf and silver scurf effectively, respectively [26, 27]. Rahman et al. [50] demonstrated the effectiveness of Filthane M-45, Melody Duo, Secure, Metaril, and Ridomil gold to minimise Phytophthora infestans-induced late blight improve the yield of potato. More so, the application of dimethomorph, mancozeb, and fenamidone + mancozeb can significantly reduce the severity of late blight and increase potato yield [51]. The application of the antagonist *Trichoderma harzianum* combined with flutolanil seed dressing offers protection against *Rhizoctonia solani* damage throughout the growing season (Wilson et al., [52]. Although fungicides have been shown to manage potato diseases effectively, they are not without their attendant problems. It is now known that continuous application of fungicides results in resistance in many pathogenic fungi of potato. Whereas metalaxyl containing fungicides show good action against *Phytophthora infestans*, prolonged applications have resulted in resistant *P. infestans* [53]. Several metalaxyl-insensitive genotypes of *P. infestans* have been reported in different regions of the world. For example, in 1980, phenylamide resistant isolates of *P. infestans* were detected on field-grown potatoes in Netherlands, Switzerland, and Ireland [48, 49, 54]. In addition to fungicide resistance, the harmful consequences on non-target organisms, risk to soil environment, and carcinogenic potentials have discouraged the use of synthetic fungicides, thereby prompting the search for efficient, safe, and eco-friendly disease management options [55, 56].

#### **4.2 Biological control**

Disease management using biological control agents is touted as efficient alternatives to chemical fungicides as they are more eco-friendly and reduce the risk of the emergence of fungicide-resistant strains of plant pathogens [57, 58]. A biological control refers to the application of microbial antagonists or their by-products to inhibit plant diseases. Organisms that antagonise plant pathogens are known as biological control agents (BCAs). Such organisms are highly specific in their action against target pathogens, their products are biodegradable, and their mass production requires low cost [59, 60].

Here, we discuss the biological control strategies – microbial inoculants (beneficial, non-pathogenic single-strains of microorganisms that antagonise plant pathogens), microbial consortium (combination of different genera or species of symbiotically living microorganisms) isolated from the natural environment, and the application of phytoextracts [61–63].

#### **4.3 Microbial inoculants**

These are single strains of active beneficial microorganisms that offer protection against diverse pathogens or promote crop productivity and health when applied to crops or incorporated into the soil [63]. Microbial inoculants are an effective and cheap alternative strategy to reduce the severity of plant diseases [64–66]. *Agrobacterium*, *Pseudomonas*, *Bacillus*, *Alcaligenes*, *Streptomyces,* and others have been reported as effective bacterial control agents [16, 17, 60]. These organisms suppress bacterial and fungal pathogens by releasing active compounds, including siderophores, antibiotics, enzymes, and the plant hormone, indole-1,3-acetic acid. *Pseudomonas* strain has been widely investigated for their potential as BCAs because of their active nature and abundance in the rhizosphere [60]. Tariq et al. [67] demonstrated the antagonistic potential of *Pseudomonas* sp. StS3 against *Rhizoctonia solani,* which causes potato black scurf. *Streptomyces violaceusniger* AC12AB promoted growth by 26.8% and significantly reduced potato typical scab disease severity by up to 90% in field trials [66]. In addition to enhancing potato tuber biomass by 33% and 22% in two location field trials, *Bacillus amyloliquefaciens* strain BAC03 considerably reduced the severity of potato scab disease by 17–57% compared to control. BAC03 also enhanced potato tuber weight by 33% and 26% in the two locations [68].

#### **4.4 Microbial consortium**

This combination of BCAs consists of various microbial strains that synergistically confer enhanced plant growth activities and superior pathogen inhibition capabilities [69–71]. Compared to single-species microbial inoculants, the microbial consortium is more useful in field applications as it offers a wide range of biocontrol activities that promote inoculant efficiency and, in turn, improve plant growth and disease suppressability [56]. The application of a microbial product comprising a consortium of *Bacillus subtilis* and *Trichoderma harzianum* inhibited common scab disease in potato caused *Streptomyces* spp. by 30.6%–46.1%, and improved yield by 23.0%–32.2% [72]. Inoculation of *Fusaria* infested soil with a bacterial consortium of *Pseudomonas aeruginosa* (B4, B23, B25, and B35), *Alcaligenes feacalis* (B16), and *S. marcescens* (B8) was reported to not only suppress fusarium wilt of potato by 94% but also considerably improved plant biomass by 186.9% (Fresh weight) and 214. 75% (dry weight) [56]. Treatment with a consortium formulation comprising *Enterobacter amnigenus* strain A167, *Serratia plymuthica* strain A294, *Serratia rubidaea* strain H440, *S. rubidaea* strain H469 and *Rahnella aquatilis* strain H145 significantly reduced potato soft rot severity and incidence by 62–75% and 48–61%, respectively, when compared to a positive control with pathogens alone [73]. Also, a combination of rhizobacteria in combination with commercial arbuscular mycorrhiza fungi (AMF) have been reported to effective in abating bacterial wilt of potato [16, 17].

#### **4.5 Phytoextracts**

Green plants harbour a plethora of secondary metabolites that could serve as eco-friendly, natural alternatives to chemical fungicides [50, 74, 75]. Phytoextracts

#### *Impact and Management of Diseases of* Solanum tuberosum *DOI: http://dx.doi.org/10.5772/intechopen.98899*

are botanicals, natural oils, and plant volatiles that show pest/pathogen control activities. They are usually extracted from fresh or dried plant parts using alcohol, water, or other solvents. Phytoextracts can be fungicidal or fungistatic in action and exert their effects by inducing conditions unfavourable for pathogen growth and proliferation [44]. The application of botanicals can significantly reduce the cost of crop protection and the occurrence of pathogen resistance [44]. Several phytoextracts have been widely tested and reported as effective suppressors of plant pathogens [50, 75]. Dried cheerota plant (*Swertia chirata* Ham.) and jute leaf (*Corchorus capsularis* L.) have been reported to exhibit *in vitro* antibacterial activity against *Erwinia carotovora* subsp. *carotovora* (Ecc) P-138 s, the causative pathogen of soft rot in potato. Under storage conditions, the plant extracts also considerably attenuated bacterial soft rot disease of different potato varieties [50]. Regardless of the mode of application (seed coating or soil inclusion), Canada milkvetch extract (MVE) effectively abated *Verticillium dahlia-*induced wilt by 55–84% in two potato cultivars – Kennebec and Russet Burbank compared to the control under growth room conditions. MVE also significantly reduced vascular discolouration and infection by 55% and 45%, respectively, in two potato cultivars in the first year of the field trial. In the second year, MVE reduced all wilt parameters by 19–31% while increasing yield by 18% on the cultivar Kennebec [76]. Soil drenching with aqueous leaf extracts of *Hibiscus sabdariffa*, *Eucalyptus globulus,* and *Punica granatum* substantially reduced the severity of bacterial wilt disease of potato relative to inoculated control under greenhouse and field conditions. While the reduction in disease severity under field conditions was similar (up to 63.23 to 68.39%) for all the three plant extracts, *E. globulus* leaf extract showed maximum abatement (94% reduction) of disease symptom development under greenhouse condition compared to extracts of *H. sabdariffa* and *P. granatum* [77]. Fumigation of seed tubers of potato with *Allium sativum* – derived essential oils has been shown to manage stem cancer, silver scurf, dry rot, black scurf, and gangrene in small–scale farming systems [78, 79].

#### **4.6 Cultural control**

A well-known cultural method to manage the diseases of potato is crop rotation. This refers to cultivating economic plants in recurrent succession and a sequential fashion on the same piece of land [80]. Rotation using different cover crops and suitable fallow periods can contribute to the attenuation of multiple soil-borne pathogens and diseases and enhance the diversity of beneficial soil microflora [81]. Evidence is mounting to show the use of *Brassica* spp. like cabbage, broccoli, cabbage, kale, cauliflower, turnip, rapeseed, canola, radish, different mustards, and other related plants as rotation or green manure crops [82, 83]. These crops produce sulphur-containing glycosinolates degraded as part of a biofumigation process to generate isothiocyanates deleterious to several soil pathogens. *Brassica* spp. have been effectively used to abate populations of soil-borne fungal pathogens, nematodes, and weeds and promote crop yield and soil properties [82]. Other non-brassica crops like ryegrass have good suppression ability over soil-borne pathogens. In several rotation studies, rapeseed and canola crops prior to potato cultivation significantly attenuated (in the range of 25–75%) soil-borne disease due to common scab and *Rhizoctonia* over many seasons to less successful rotations or no rotation [84, 85]. A field trial at a highly infested site with a powdery scab, ryegrass, rapeseed, canola, and Indian mustard grown as rotation crops and green manure suppressed powdery scab in the subsequent potato crop 15–40%. Additionally, rapeseed and canola abated black scurf by 70–80% compared to a standard oats rotation (**Figure 1**) [82].


#### **Figure 1.**

*Management strategies for potato diseases.*

#### **5. Methods for raising disease-free potato**

Potato is affected by a wide range of fungal, viral, bacterial, and nematodal diseases [86]. These result in colossal yield loss annually. Therefore, it is imperative to exploit strategies for raising disease-free potato to reduce losses caused by pathogens, thus ensuring food security.

Some of the strategies for raising disease-free potato are:

#### **5.1 Conventional plant breeding**

The breeding of potato is a huge task due to inherent genetic and biological factors. Breeding for increased resistance to *Phytophthora infestans* (causal agent of late blight) is one of the most critical targets in potato breeding [87]. Plant breeders incorporated resistance against early and late blight disease by crossing hybrid lines with wild species (*S*. *brevidens* and *S. bulbocastanum*), which exhibited resistance against fungal pathogens [88, 89]. Potato plants resistant to diseases have been produced using conventional plant breeding. However, this process is tedious, and it takes time to achieve success.

#### **5.2 Induced resistance**

Resistance in plants can be induced by applying exogenous substances, or agents including living and non-living agents. Resistance to both fungal and viral diseases has been reported in potato. Quintanilla and Brishammar [90] reported systemic induced resistance to late blight in potato by treating with salicylic acid and *Phytophthora crptogea*. In their study, the non-pathogenic fungus *Phytophthora crptogea* and salicylic acid were used as inducer agents. Nadia *et al*., [91] showed that chemicals under greenhouse and field conditions induced resistance against early and late blight diseases. The inducers used in this study were ascorbic acid, dichloro-isonicotinic acid, ethylene diamine tetraacetic acid, and calcium chloride. Chemicals and fungicides (at low concentration) can induce resistance [92]; similar reports include

Andreu *et al*., [93]. Several studies have reported using biological agents as inducers of resistance in potato [94–98] reported mycorrhiza-induced resistance in potato. Induced resistance against potato virus Y (PVYNTN) has also been achieved [99].

#### **5.3 Genetic engineering approach**

Genetic engineering has been used to raise-disease free transgenic potato plants. However, this technique requires specialised skill, sophisticated equipment, and technical know-how. However, the problem of acceptance and ethical issues may also arise.

Extreme resistance to late blight disease by transferring 3 *R* genes from wild relatives into African farmer-preferred potato varieties was reported by [100]. Three late blight resistance genes from wild potato species were transferred as a stack into the farmer-preferred varieties, Tigoni and Shangi. *R* gene expression analysis in 18 transgenic events showed different transgenic events exhibiting different expression levels in the three genes. Engineering virus resistance using a modified potato gene has been reported by [101]. They reported that the transgenic expression of the *pvrl*<sup>2</sup> gene from pepper confers resistance to potato virus Y (PVY) in potato. The development of late blight-resistant potato by cisgene stacking was studied by Jo *et al*., [102].

RNA interference (RNAi) is an emerging post-transcriptional technique that has been used to produce crops resistant to diseases. Production of potato lines resistant to *P. infestans* through the RNAi technique has been reported [103]. RNAi technology can be directed to degrade the pathogen's mRNA that enters the host cell or silence endogenous genes of the host cell that aid pathogenicity. RNAi's mechanism of pathogen control is not dependent on producing a foreign protein that could be allergenic or toxic in the host plants. This makes this technology more acceptable than the typical transgenic approaches for disease control [104].

#### **5.4 Plant tissue culture techniques**

This technique can be used to produce disease-free pre-basic seeds. Disease-free pre-basic seed potato was produced through tissue culture in Nepal [105]. The use of disease-free seeds can help reduce the transmission of pathogens from propagating materials such as tuber to the field. It has been reported that quality seeds alone can increase yield by 15–20% in Bangladesh [106]. Therefore, micropropagation of potato can help reduce disease transmission through propagating materials; however, little has been achieved on the use of somatic embryos [107], and more researches are required for more remarkable breakthroughs in this regard.

#### **5.5 New/advanced breeding techniques**

Genome editing of potato using new technologies such as zinc-finger nucleases (ZFNs), transcription activator-like effector nucleases (TALENs), and clustered regularly interspaced palindromic repeats (CRISPR) associated nuclease 9 is currently been exploited. CRISPR/Cas9 has emerged as a breakthrough in gene editing; however, limited studies have been done on potatoes using this technique [108]. Genome editing using CRISPR/Cas9 has been used to engineer virus resistance in plants by targeting host genes directly involved in host-viral interactions [109–113]. This technique has been used to knock out potato genes/factors like eukaryotic translation initiation factors (*elf4E* and isoform *elf*(*iso*) *4E* that interact with viruses to assist viral infection [114]. Potato varieties resistant to viruses can be produced using this technique. Late blight resistance in potato has also been achieved using CRISPR/Cas9 genome editing. Functional knockouts of *stDND1*, *StCHL1,* and *DMG400000582* (*STDMR6*–1) genes generated increased resistance against late blight in potato [115].

Therefore, holistic and integrated approaches are required for raising diseasefree potato in order to overcome the ever-evolving phytopathogens and mitigate losses; including post-harvest losses caused by these pathogens, therefore ensuring food security.

## **6. Conclusions**

This chapter discusses the host-pathogens association of different diseases in potato and their impact on yield. The findings highlight management strategies of these diseases: chemical control, biological control, microbial inoculants, microbial consortium, phytoextracts, and cultural control. In addition, current methods for raising disease-free potatoes to reduce annual yield loss were reported in detail. Based on the presented findings, annual yield loss (pre-and post-harvest) is still high. Thus, the management strategies alone are promising but combining the different methods and exploiting disease-free potato can translate into an integrated management approach of potato diseases.

## **Acknowledgements**

We wish to appreciate Covenant University for sponsoring this publication.

## **Conflict of interest**

The authors declare no conflict of interests.

## **Author details**

Olusola L. Oyesola1 , Oluwadurotimi S. Aworunse1 , Margaret I. Oniha1 , Onyemaechi H. Obiazikwor2 , Oluwakemi Bello1 , Olubunmi M. Atolagbe1 , Ayodele A. Sobowale3 , Jacob O. Popoola1 and Olawole O. Obembe1,4\*

1 Department of Biological Sciences, Covenant University, Otta, Ogun State, Nigeria

2 Faculty of Life Sciences, Department of Plant Biology and Biotechnology, University of Benin, Benin City, Nigeria

3 Botany Department, University Of Ibadan, Nigeria

4 UNESCO Chair on Plant Biotechnology, Covenant University, Nigeria

\*Address all correspondence to: olawole.obembe@covenantuniversity.edu.ng

© 2021 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

*Impact and Management of Diseases of* Solanum tuberosum *DOI: http://dx.doi.org/10.5772/intechopen.98899*

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#### **Chapter 12**
