Abstract
Somatic hybridization has been used in potato to overcome the sexual barriers between the cultivated (Solanum tuberosum L.) and wild species. To date hundreds of inter/intra-specific somatic hybrids have been produced via protoplast fusions using 23 Solanum species and characterized for multiple traits such as agronomic, disease/pest resistance, salinity, frost and others. With increasing success in recovery of fusion products, somatic hybrids have been exploited in potato genetics, breeding and genomics studies. Here, we report on progress in somatic hybridization research in potato during the past 40 years.
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Introduction
A huge genetic diversity is available in Solanum species for various desirable traits. Wild Solanum species have been used in potato breeding but they represent only a small fraction to the total Solanum diversity (Bradshaw et al. 2006). Huge efforts are involved in successful utilization of wild species to widen the narrow genetic base of the cultivated potato. Many useful genes of wild sources cannot be transferred to common potato through conventional breeding because of sexual barriers caused by the differences in ploidy level and endosperm balance number (EBN) (Spooner and Salas 2006). However, it is now possible to overcome the sexual barriers using methods such as manipulation of ploidy and EBN, bridge crosses, mentor pollination, embryo rescue, hormone treatment, reciprocal crosses, and somatic hybridization (reviewed by Jansky 2006).
Somatic hybridization aims to strengthen the potato gene pool by introducing genes from wild species (Helgeson et al. 1993). This technique allows several advantages over conventional breeding and transgenic methods, such as: (i) produces fertile somatic hybrids with target traits of wild Solanum species, (ii) provides access to basic pre-breeding material for effective utilization in breeding, (iii) enables easy transfer of monogenic and polygenic traits in one step, (iv) results recombination of nuclear and cytoplasmic genomes, and (v) avoids biosafety regulatory issues associated with transgenics. In the past, a few researchers discussed protoplast fusion in Solanum species (Orczyk et al. 2003), plant species (Waara and Glimelius 1995; Davey et al. 2005; Eeckhaut et al. 2013), citrus (Guo et al. 2013), asymmetric fusion (Lakshmanan et al. 2013), asymmetric fusion in wheat (Liu and Xia 2014), integeneric fusion (Liu et al. 2005) and crop improvement in China (Wang et al. 2013).
During the past 40 years, hundreds of symmetric somatic hybrids were produced, while limited number of asymmetric somatic hybrids. More than a decade back, Orczyk et al. (2003) reviewed application of potato somatic hybrids in genetics and breeding, and since then several progresses have been achieved in this area. Hence, this review aims to progress in somatic hybridization research in potato and their application to genetics, breeding and genomics studies.
Production of potato somatic hybrids
Protoplast fusion and culture
Protoplast isolation was first reported by Cocking (1960) and after a decade successful regeneration of plants from tobacco protoplasts was obtained (Takebe et al. 1971). With the development of electrofusion method, reports on recovery of somatic hybrid regenerants were more prevalent (Fish et al. 1988; Bates et al. 1987) might be due to easy control of fusion parameters than the chemical method using polyethylene glycol (Wallin et al. 1974). In food crops, electrofusion was first reported in potato between Solanum tuberosum and semi-cultivated (S. phureja) (Puite et al. 1986). Leaf-mesophyll cells of in vitro-grown plants were used largely to isolate protoplasts in protoplast-digestion-medium (Binding et al. 1978), then culture of fused products in the VKM medium (Binding and Nehls 1977) followed by shoot development on the MS13K medium (Behnke 1975). Recently, selection of somatic hybrids (S. tuberosum + S. chacoense) based on callus growth tagged with green fluorescent protein has been observed (Rakosy-Tican and Aurori 2015). Over the 40 years, many fusion experiments were executed and hundreds of potato somatic hybrids were produced and characterized, as summarized in Table 1.
Confirmation of somatic hybrids
A large number of potato somatic hybrids have been produced via protoplast fusion between the common and the wild species S. acaule, S. berthaultii, S. brevidens, S. bulbocastanum, S. cardiophyllum, S. chacoense, S. circaefolium, S. commersonii, S. etuberosum, S. × michoacanum, S. melongena, S. nigrum, S. phureja, S. pinnatisectum, S. tuberosum, S. sanctae-rosae, S. spegazzinii, S. stenotomum, S. tarnii, S. torvum, S. vernei, S. verrucosum, and S. villosum (Table 1). Following regeneration, somatic hybrids are characterized through various methods such as cytological (flow cytometry, chromosome count, guard cell count, FISH-fluorescence in situ hybridization and GISH-genomic in situ hybridization), isozyme, molecular markers (e.g. RAPD-random amplified polymorphic DNA, RFLP-restriction fragment length polymorphism, ISSR-inter simple sequence repeat, SSR-simple sequence repeat, AFLP-amplified fragment length polymorphism, and DArT-diversity array technology), phenotypes (e.g. foliage, stem, leaf, flower and tuber traits) and pollen fertility. Somatic hybrids are also analyzed for cytoplasm types (W/α, T/β, W/γ, W/δ and S/ε: Lössl et al. 1999) based on organelle (chloroplast and mitochondria) genomes-specific markers as described by Lössl et al. (2000). Finally, somatic hybrids are examined for the presence of target traits under field or controlled conditions and for many other traits (Table 1).
Application of potato somatic hybrids
Genetics
Genetics of potato somatic hybrids and their segregating progenies has been studied in nuclear genome to dissect the recombination patterns. Chromosomal segregation pattern was analyzed in hexaploid somatic hybrid (S. brevidens + S. tuberosum) and their progenies with S. tuberosum using RFLP (Williams et al. 1993) and RAPD markers (McGrath et al. 1996). Study suggests that S. brevidens ribosomal (r)DNA loci are primarily contributed to isochromosome formation in the hybrids and progenies (McGrath and Helgeson 1998). Further, a chromosome substitution line was developed in S. breviden-somatic hybrids and progenies (BC1, BC2 and BC3) with S. tuberosum. Study demonstrates that a single copy of chromosome 8 from S. brevidens replaced the same in the BC3 clone and has significant impact on transferring resistance to tuber soft rot and early blight (Tek et al. 2004). The importance of combining GISH and DNA markers was also suggested to study chromosomal behaviours in potato (Dong et al. 1999). Poor chromosomal pairing was observed in somatic hybrid (S. etuberosum + S. tuberosum) and progenies (BC1 and BC2), and further suggests that genome dosage affects tuber formation but has less effect on potato virus Y (PVY) resistance (Gavrilenko et al. 2003). Evidence of tetrasomic inheritance was investigated in a tetraploid somatic hybrid (S. commersonii + S. tuberosum) and F2 progeny (90 individuals) using RAPD and AFLP markers. Segregation pattern was investigated by RFLP markers in a progeny of hexaploid somatic hybrids (S. acaule + S. tuberosum) crossed with S. tuberosum. Study concludes that somatic hybridization allows the effective use of S. acaule genes into cultivated potato (Yamada et al. 1998). Recently, somatic hybrid (S. chacoense + S. tuberosum) exhibited tetrasomic or disomic segregation ratio using SSR markers and suggested that pentaploid hybrid exhibits tetraploid inheritance pattern (Chen et al. 2016).
Interaction between nuclear and cytoplasmic genes can affects fertility and agronomic traits of somatic hybrids and progenies (Lössl et al. 1994). Segregation and recombination patterns of organelle genomes were investigated in potato (Frei et al. 1998) and analyzed variations (Tiwari et al. 2014, 2016). Majority of somatic hybrids follow recombination of mitochondrial genome from both parents, and chloroplast pattern from only one parent (e.g. S. bulbocastanum: Iovene et al. 2007; S. pinnaticetum: Sarkar et al. 2011; S. chacoense: Chen et al. 2013) except recombination of chloroplast genome was observed only once in S. vernei-somatic hybrid (Trabelsi et al. 2005). Loss of male sterility was observed in nuclear-mitochondrial genomes re-arrangement in S. commersonii-somatic hybrids and BC1 progeny (Cardi et al. 1999). Study suggests possibility of exploitation of novel cytoplasm in potato breeding (Scotti et al. 2003), especially variation in a hot spot mitochondrial region (rpl5–rps14) (Scotti et al. 2004). Recently, random and non-random segregations of organelle genomes were observed in somatic hybrid (S. × michoacanum + S. tuberosum) using DArT markers (Smyda-Dajmund et al. 2016).
In search of new genes, a few potato somatic hybrids and their progenies were exploited in linkage mapping studies. The RB gene (Rpi-blb1) originates from diploid wild species S. bulbocastanum of somatic hybrid (S. bulbocastanum + S. tuberosum) and confers durable resistance to late blight (Helgeson et al. 1998; Song et al. 2003). The RB gene was mapped to potato chromosome 8 through analysis of somatic hybrid progenies (BC1 and BC2) (Naess et al. 2000, 2001). Besides, genetic stability of in vitro plants of somatic hybrids (S. tuberosum dihaploid ‘C-13’ + S. pinnatisectum, and ‘C-13’ + S. etuberosum) was confirmed using methylation-sensitive amplified polymorphism (MSAP) and AFLP markers (Tiwari et al. 2013b, 2015d, e).
Breeding
Improvement of somatic hybrids is essential for desirable agronomic traits through breeding methods to decrease the undesirable effects of wild species. In addition, transfer of disease/pest resistance traits from somatic hybrids to progenies is also important. Assessment of genetic and phenotypic variation among somatic hybrids (Gavrilenko et al. 1999) and development of their advanced progenies such as F2, BC1, BC2 and BC3 led to the effective utilization somatic hybrids in potato breeding (Table 1). Somatic hybrids performed better in field trials in terms of tuber traits and phenotypes (Carrasco et al. 2000), and a few hybrid produced higher yield, tuber number and weight than parents (Möllers et al. 1994). Tuber yield per plant of backcrossed progenies of somatic hybrid improved considerably to that of parents (Carputo et al. 2000). Recently, potential S. pinnatisectum-somatic hybrids (P4, P8 and P10) were selected for adaptability, tuber traits, late blight resistance and keeping quality traits in the sub-tropical plains of India, where nearly 90% of potato is grown (Luthra et al. 2016). Further, hybrid progeny (BC1) was also generated that can be utilized in potato breeding (Luthra et al. 2016). Good tuber yield and quality were observed in S. tarnii-somatic hybrids derived BC1 progenies in the field trials (Thieme et al. 2008). Further, study confirmed that somatic hybrid had resistance to both PVY and late blight, of which only PVY resistance was transferred to BC1 progeny. Multiple years of field evaluations of S. etuberosum-somatic hybrids and progenies showed stable transmission and expression of PLRV and PVY resistances in three (BC1, BC2 and BC3) and two (BC1 and BC2) generations, respectively (Novy et al. 2007). Besides, resistances to PVY, potato leaf roll virus (PLRV), and Green peach aphid in BC2 progeny were also observed (Novy et al. 2002). Another study demonstrates that late blight resistance can be transferred successfully through breeding from tetraploid somatic hybrids (S. × michoacanum + S. tuberosum and autofused S. × michoacanum) to common varieties (Smyda-Dajmund et al. 2017). Bacterial wilt resistance was transferred to advanced progenies of S. commersonii-somatic hybrids, and three highly resistant clones (BC1 and BC2) were selected as breeding materials (Kim-Lee et al. 2005). The effect of genetic constitution of S. tuberosum was investigated in S. bulbocastanum-somatic hybrids and progenies (BC1 and BC2) (Rakosy-Tican et al. 2015). Field performance for foliage maturity and tuber traits (tuber yield, tuber number, tuber weight and specific gravity) was observed in somatic hybrids (S. brevidens/S. commersonii + S. tuberosum) progenies and implicated to breeding for freezing tolerance (Chen et al. 1999a, b, c). Somatic hybrids (S. commersonii + S. tuberosum) were observed more similar to cultivated potato for phenotypes in field evaluations and developed F2 progeny, and suggested to fast transfer of useful traits from S. commersonii into cultivated background (Cardi 1998; Cardi et al. 2002). Recently, SSR alleles were identified for bacterial wilt resistance breeding in S. chacoense-somatic hybrids and backcross progenies (Chen et al. 2016). To our knowledge, there are investigations up to BC3 generation and all attempts to exploit somatic hybrid material so far have not resulted in registration of a potato cultivar.
Genomics
Somatic hybrids and their parents have been exploited to a limited extent in genomics studies. Whole genome sequences of chloroplast genome of wild potato species (S. commersonii), a commonly used fusion parent, was deciphered and identified two Indel markers for application in chloroplast genotyping (Cho et al. 2016). High-throughput genotyping of somatic hybrids (S. × michoacanum + S. tuberosum) showed presence of both parental chromosomes and loss of some markers (13.9–29.6%) in the hybrids using 5358 DArT markers analysis (Smyda-Dajmund et al. 2016). In functional genomics, genes controlling potato tuberization in tuber-bearing somatic hybrids (S. tuberosum + S. etuberosum) vs. control parent (S. etuberosum-non tuberous) were identified using microarray. Findings suggest candidate genes expression in leaf tissue of somatic hybrid are implicated to tuber growth and development process such as transport, carbohydrate metabolism, phytohormones and transcription/translation/binding functions (Tiwari et al. 2015a, b). In another study, late light resistance genes were identified in somatic hybrid (S. tuberosum + S. pinnatisectum) by microarrays and study suggests a broad spectrum of candidate genes involved in late blight resistance in the hybrid (Singh et al. 2016a). Further, recent identification of eight miRNAs (miR395, 821, 1030, 1510, 2673, 3979, 5021 and 5213) in S. pinnatisectum-somatic hybrid for late blight resistance genes and their targets has led a new insight in potato biology (Singh et al. 2016b). Most of the predicted target genes are associated with different biological processes such as disease resistance proteins and transcription factors families. These miRNAs could be manipulated through RNAi technique for transgenic development (Singh et al. 2016b). Thus, with the advancement in next-generation sequencing technologies, there is a huge scope to deploy these novel tools in somatic hybrids to study potato biology and improvement.
Conclusions
Limited utilization of Solanum species and therefore narrow genetic base of the cultivated potato is the cause of concern of yield stagnation. Development of potato somatic hybrids using wild Solanum species with desirable attributes could be helpful to address this issue. Analyzing breeding potential of somatic hybrids, development of advanced progenies by hybridization with common varieties and identification of linked molecular markers are important for their successful applications in potato breeding. Further, information on nuclear-cytoplasmic interaction between cultivated and wild species could also be useful while exploitation of somatic hybrids in breeding. Novel genomics tools like whole genome sequencing, DArT markers based genotyping, microRNAs, microarrays and many others could strengthen the somatic hybridization research in potato. Moreover, access to the next-generation sequencing technologies and the potato genome sequences are essential for application of genome-enabled breeding strategies and critical investigation of complex biological processes like heat tolerance, tuberization, yield, disease-pest resistance and other traits. The sequence data could be extensively deployed for gene discovery, marker development, phylogeny, comparative genomics, breeding, and engineering of new genotypes/phenotypes. Taken together, above studies would provide a myriad of useful information available in somatic hybrids for potato genetics, breeding and genomics improvement.
References
Ahn YK, Park T-H (2013) Resistance to common scab developed by somatic hybrids between Solanum brevidens and Solanum tuberosum. Acta Agri Scand B 63:595–603
Austin S, Baer M, Ehlenfeldt M, Kazmierczak PJ, Helgeson JP (1985a) Intra-specific fusions in Solanum tuberosum. Theor Appl Genet 71:172–175
Austin S, Baer MA, Helgeson JP (1985b) Transfer of resistance to potato leaf roll virus from Solanum brevidens into Solanum tuberosum by somatic fusion. Plant Sci 39:75–82
Austin S, Ehlenfeldt MK, Baer MA, Helgeson JP (1986) Somatic hybrids produced by protoplast fusion between S. tuberosum and S. brevidens: phenotypic variation under field conditions. Theor Appl Genet 71:682–690
Austin S, Lojkowska E, Ehlenfeldt MK, Kelman A, Helgeson JP (1988) Fertile interspecific hybrids of Solanum: a novel source of resistance to Erwinia soft rot. Phytopathology 78:1216–1220
Austin S, Pohlman JD, Brown CR, Mojtahedi H, Santo GS, Douches DS, Helgeson JP (1993) Interspecific somatic hybridization between Solanum tuberosum L. and S. bulbocastanum Dun. as a means of transferring nematode resistance. Am Potato J 70:485–495
Aversano R, Savarese S, Nova JMD, Frusciante L, Punzo M, Carputo D (2009) Genetic stability at nuclear and plastid DNA level in regenerated plants of Solanum species and hybrids. Euphytica 165:353–361
Barone A, Li J, Sebastiano A, Cardi T, Frusciante L (2002) Evidence for tetrasomic inheritance in a tetraploid Solanum commersonii (+) S. tuberosum somatic hybrid through the use of molecular markers. Theor Appl Genet 104:539–546
Bastia T, Carotenuto N, Basile B, Zoina A, Cardi T (2000) Induction of novel organelle DNA variation and transfer of resistance to frost and Verticillium wilt in Solanum tuberosum through somatic hybridization with 1EBN S. commersonii. Euphytica 116:1–10
Bastia T, Scotti N, Cardi T (2001) Organelle DNA analysis of Solanum and Brassica somatic hybrids by PCR with universal primers. Theor Appl Genet 102:1265–1272
Bates GW, Nea LJ, Hasenkampf CA (1987) Electrofusion and plant somatic hybridization. In: Sowers AE (ed) Cell fusion. Plenum Press, New York, pp 479–496
Behnke M (1975) Regeneration in Gewebekulturen einiger dihaplider Solanum tuberosum-Klone. Z Pflanzenzücht 75:262–265
Bidani A, Nouri-Ellouz O, Lakhoua L, Sihachakr D, Cheniclet C, Mahjoub A, Drira N, Gargouri-Bouzid R (2007) Interspecific potato somatic hybrids between Solanum berthaultii and Solanum tuberosum L. showed recombinant plastome and improved tolerance to salinity. Plant Cell Tissue Organ Cult 91:179–189
Binding H, Nehls R (1977) Regeneration of isolated protoplasts to plant Solanum dulcamara L. Z. Pflanzenphysiology 85:279–280
Binding H, Nehls R, Schieder O, Sopory SK, Wenzel G (1978) Regeneration of mesophyll protoplasts isolated from dihaploid clones of Solanum tuberosum L. Plant Physiol 43:52–54
Binding H, Jain SM, Finger J, Mordhorst G, Nehls R, Gressel J (1982) Somatic hybridization of an atrazine resistant biotype of Solanum nigrum with Solanum tuberosum. Part 1. clonal variation of morphology and in atrazine sensitivity. Theor Appl Genet 63:273–277
Bołtowicz B, Szczerbakowa A, Wielgat B (2005) RAPD analysis of the interspecific somatic hybrids Solanum bulbocastanum (+) S. tuberosum. Cell. Mol Biol Lett 10:51–162
Bradshaw JE, Bryan GJ, Ramsay G (2006) Genetic resources (including wild and cultivated Solanum species) and progress in their utilisation in potato breeding. Potato Res 49:49–65
Cai XK, Liu J, Xie CH (2004) Mesophyll protoplast fusion of Solanum tuberosum and Solanum chacoense and their somatic hybrid analysis. Acta Hortic Sin 31:623–626
Cardi T (1998) Multivariate analysis of variation among Solanum commersonii (+) S. tuberosum somatic hybrids with different ploidy levels. Euphytica 99:35–41
Cardi T, Ambrosio FD, Consoli D, Puite KJ, Ramulu KS (1993) Production of somatic hybrids between frost-tolerant Solanum commersonii and S. tuberosum: characterization of hybrid plants. Theor Appl Genet 87:193–200
Cardi T, Bastia T, Monti L, Earle ED (1999) Organelle DNA and male fertility variation in Solanum spp. and interspecific somatic hybrids. Theor Appl Genet 99:819–828
Cardi T, Mazzei M, Frusciante L (2002) Field variation in a tetraploid progeny derived by selfing a Solanum commersonii (+) S. tuberosum somatic hybrid: a multivariate analysis. Euphytica 124:111–119
Carputo D, Basile B, Cardi T, Frusciante L (2000) Erwinia resistance in backcross progenies of Solanum tuberosum × S. tarijense and S. tuberosum (+) S. commersonii hybrids. Potato Res 43:135–142
Carrasco A, De Galarreta JIR, Rico A, Ritter E (2000) Transfer of PLRV resistance from Solanum verrucosum Schlechdt to potato (S. tuberosum L.) by protoplast electrofusion. Potato Res 43:31–42
Chandel P, Tiwari JK, Ali N, Devi S, Sharma SH, Sharma SA, Luthra SK, Singh BP (2015) Interspecific potato somatic hybrids between Solanum tuberosum and S. cardiophyllum, potential sources of late blight resistance breeding. Plant Cell Tissue Organ Cult 123:579–589
Chen Y-KH, Bamberg JB, Palta JP (1999a) Expression of freezing tolerance in the interspecific F1 and somatic hybrids of potatoes. Theor Appl Genet 98:995–1004
Chen Y-KH, Palta JP, Bamberg JB, Kim H, Haberlach GT, Helgeson JP (1999b) Expression of non-acclimated freezing tolerance and cold acclimation capacity in somatic hybrids between hardy wild Solanum species and cultivated potatoes. Euphytica 107:1–8
Chen Y-KH, Palta JP, Bamberg JB (1999c) Freezing tolerance and tuber production in selfed and backcross progenies derived from somatic hybrids between Solanum tuberosum L. and S. commersonii Dun. Theor Appl Genet 99:100–107
Chen L, Guo X, Xie C, He L, Cai X, Tian L, Song B, Liu J (2013) Nuclear and cytoplasmic genome components of Solanum tuberosum + S. chacoense somatic hybrids and three SSR alleles related to bacterial wilt resistance. Theor Appl Genet 126:1861–1872
Chen L, Guo X, Wang H, Xie C, Cai X, He L, Zhou J, Liu J (2016) Tetrasomic inheritance pattern of the pentaploid Solanum chacoense (+) S. tuberosum somatic hybrid (resistant to bacterial wilt) revealed by SSR detected alleles. Plant Cell Tissue Organ Cult 127:315–323
Cheng J, Saunders JA, Sinden SL (1995) Colorado potato beetle resistant somatic hybrid potato plants produced via protoplast electrofusion. In Vitro Cell Dev Biol 31:90–95
Cho K-S, Cheon K-S, Hong S-Y, Cho J-H, Im J-S, Mekapogu M, Yu Y-S, Park T-H (2016) Complete chloroplast genome sequences of Solanum commersonii and its application to chloroplast genotype in somatic hybrids with Solanum tuberosum. Plant Cell Rep 35:2113–2123
Cocking EC (1960) A method for the isolation of plant protoplasts and vacuoles. Nature 187:962–963
Cooper-Bland S, Main DMJ, Fleming MH, Phillips MS, Powell W, Kumar A (1994) Synthesis of intraspecific somatic hybrids of Solanum tuberosum: assessments of morphological, biochemical and nematode (Globodera pallida) resistance characteristics. J Exp Bot 45:1319–1325
Davey MR, Anthony P, Power JB, Lowe KC (2005) Plant protoplast technology: current status. Acta Physiol Plant 27:117–129
Dong F, Novy RG, Helgeson JP, Jiang J (1999) Cytological characterization of potato-Solanum etuberosum somatic hybrids and their backcross progenies by genomic in situ hybridization. Genome 42:987–992
Eeckhaut T, Lakshmanan PS, Deryckere D, Von Bockstaele E, Van Huylenbroeck J (2013) Progress in plant protoplast research. Planta 238:991–1003
Fehér A, Preizner J, Litkey Z, Casanádi G, Dudits D (1992) Characterization of chromosome instability in interspecific somatic hybrids obtained by X-ray fusion between potato (Solanum tuberosum L.) and S. brevidens Phil. Theor Appl Genet 84:880–890
Fish N, Karp A, Jones MGK (1988) Production of somatic hybrids by electrofusion in Solanum. Theor Appl Genet 76:260–266
Fock I, Collonier C, Purwito A, Luisetti J, Sonvannavong V, Vedel F, Servaes A, Ambroise A, Kodja H, Ducreux G, Sihachakr (2000) Resistance to bacterial wilt in somatic hybrids between Solanum tuberosum and S. phureja. Plant Sci 160:165–176
Fock I, Collonier C, Purwito A, Luisetti J, Sonvannavong V, Vedel F, Servaes A, Ambroise A, Kodja H, Ducreux G, Sihachakr (2001) Use of S. stenotomum for introduction of resistance to bacterial wilt in somatic hybrids of potato. Plant Physiol Biohem 39:899–908
Frei U. Stattmann M, Lössl A, Wenzel G (1998) Aspects of fusion combining ability of dihaploid Solanum tuberosum L.: influence of the cytoplasm. Potato Res. 41:155–162
Gavrilenko T, Thieme R, Tiemann H (1999) Assessment of genetic and phenotypic variation among interspecific somatic hybrids of potato, Solanum tuberosum L. Plant Breed 118:205–213
Gavrilenko T, Thieme R, Heimbach U, Thieme T (2003) Fertile somatic hybrids of Solanum etuberosum + dihaploid Solanum tuberosum and their backcrossing progenies: relationships of genome dosage with tuber development and resistance to potato virus Y. Euphytica 131:323–332
Gibson RW, Jones MGK, Fish N (1988) Resistance to potato leaf roll virus and potato virus Y in somatic hybrids between dihaploid Solanum tuberosum and S. brevidens. Theor Appl Genet 76:113–117
Gillen AM, Novy RG (2007) Molecular characterization of the progeny of Solanum etuberosum identifies a genomic region associated with resistance to potato leafroll virus. Euphytica 155:403–415
Greplová M, Polzerová H, Vlastníková H (2008) Electrofusion of protoplasts from Solanum tuberosum, S. bulbocastanum and S. pinnatisectum. Acta Physiol Plant 30:787–796
Guo X, Xie C, Cai X, Song B, He L, Liu J (2010) Meiotic behavior of pollen mother cells in relation to ploidy level of somatic hybrids between Solanum tuberosum and S. chacoense. Plant Cell Rep 29:1277–1285
Guo W-W, Xiao S-X, Deng X-X (2013) Somatic cybrid production via protoplast fusion for citrus improvement. Scientia Hort 163:20–26
Harding K, Millam S (2000) Analysis of chromatin, nuclear DNA and organelle composition in somatic hybrids between Solanum tuberosum and Solanum sanctae-rosae. Theor Appl Genet 101:939–947
Helgeson JP, Haberlach GT, Ehlenfeldt MK, Hunt G, Pohlman JD, Austin S (1993) Fertile somatic hybrids of potato and wild Solanum species: potential for use in breeding programs. Am Potato J 70:437–452
Helgeson JP, Pohlman JD, Austin S, Haberlach GT, Wielgus SM, Ronis D, Zambolim L, Tooley P, McGrath JM, James RV, Stevenson WR (1998) Somatic hybrids between Solanum bulbocastanum and potato: a new source of resistance to late blight. Theor Appl Genet 96:738–742
Horsman K, Bergervoet JEM, Jacobsen E (1997) Somatic hybridization between Solanum tuberosum and species of the S. nigrum complex: Selection of vigorously growing and flowering plants. Euphytica 96:345–352
Iovene M, Savarese M, Cardi T, Frusciante L, Scott N, Simon PW, Carputo D (2007) Nuclear and cytoplasmic genome composition of Solanum bulbocastanum (+) S. tuberosum somatic hybrids. Genome 50:443–450
Iovene M, Aversano R, Savarese S, Caruso I, Dimatteo A, Cardi T, Frusciante L, Carputo D (2012) Interspecific somatic hybrids between Solanum bulbocastanum and S. tuberosum and their haploidization for potato breeding. Biol Plant 56:1–8
Jadari R, Sihachakr D, Rossignol L, Ducreux G (1992) Transfer of resistance to Verticillium dahliae Kleb. from Solanum torvum S.W. into potato (Solanum tuberosum L.) by protoplast electrofusion. Euphytica 64:39–47
Jansky S (2006) Overcoming hybridization barriers in potato. Plant Breed 125:1–12
Kim H, Coi SU, Chae MS, Weilgus SM, Helgeson JP (1993) Identification of somatic hybrids produced by protoplast fusion between Solanum commersonii and S. tuberosum haploid. Korean J Plant Tissue Cult 20:337–344
Kim-Lee H, Moon JS, Hong YJ, Kim MS, Cho HM (2005) Bacterial wilt resistance in the progenies of the fusion hybrids between haploid of potato and Solanum commersonii. Am J Potato Res 82:129–137
Kozukue N, Misoo S, Yamada T, Kamijima O, Friedman M (1999) Inheritance of morphological characters and glycoalkaloids in potatoes of somatic hybrids between dihaploid Solanum acaule and tetraploid Solanum tuberosum. J Agric Food Chem 47:4478–4483
Laferriere LT, Helgeson JP, Allen C (1999) Fertile Solanum tuberosum + S. commersonii somatic hybrids as sources of resistance to bacterial wilt caused by Ralstonia solanacearum. Theor Appl Genet 98:1272–1278
Lakshmanan PS, Tom E, Dieter D, Erik VB, Johan VH (2013) Asymmetric somatic plant hybridization: status and applications. Am J Plant Sci 4:1–10
Laurila J, Laasko I, Larkka J, Gavrilenko T, Rokka V-M, Pehu E (2001) The proportions of glycoalkaloid aglycones are dependent on the genome constitutions of interspecific hybrids between two Solanum species (S. brevidens and S. tuberosum). Plant Sci 161:677–683
Lightbourn GJ, Veilleux RE (2007) Production and evaluation of somatic hybrids derived from monoploid potato. Am J Potato Res 84:425–435
Lightbourn GJ, Jelesko JG, Veilleux RE (2007) Retrotransposon-based markers from monoploids used in somatic hybridization. Genome 50:492–501
Liu S, Xia G (2014) The place of asymmetric somatic hybridization in wheat breeding. Plant Cell Rep 33:595–603
Liu J, Xu X, Deng X (2005) Intergeneric somatic hybridization and its application to crop genetic improvement. Plant Cell Tissue Org Cult 82:19–44
Lössl A, Frei U, Wenzel G (1994) Interaction between cytoplasmic composition and yield parameters in somatic hybrid of S. tuberosum L. Theor Appl Genet 89:873–878
Lössl A, Adler N, Horn R, Frei U, Wenzel G (1999) Chondriome-type characterization of potato: mt α, β, γ, δ, ε and novel plastid-mitochondrial configurations in somatic hybrids. Theor Appl Genet 98:1–10
Lössl A, Götz M, Braun A, Wenzel G (2000) Molecular markers for cytoplasm in potato: male sterility and contribution of different plastid-mitochondrial configurations to starch production. Euphytica 116:221–230
Luthra SK, Tiwari JK, Lal M, Chandel P, Kumar V (2016) Breeding potential of potato somatic hybrids: evaluations for adaptability, tuber traits, late blight resistance, keeping quality and backcross (BC1) progenies. Potato Res 59:375–391
Mattheij WM, Puite KJ (1992) Tetraploid potato hybrids through protoplast fusions and analysis of their performance in the field. Theor Appl Genet 83:807–812
McGrath JM, Helgeson JP (1998) Differential behavior of Solanum brevidens ribosomal DNA loci in a somatic hybrid and its progeny with potato. Genome 41:435–439
McGrath JM, Wielgus SM, Helgeson JP (1996) Segregation and recombination of Solanum brevidens synteny groups in progeny of somatic hybrids with S. tuberosum: intra-genomic equals or exceeds inter-genomic recombination. Genetics 142:1335–1348
McGrath JM, Williams CE, Haberlach GT, Wielgus SM, Uchytil TF, Helgeson JP (2002) Introgression and stabilization of Erwinia tuber soft rot resistance into potato after somatic hybridization of Solanum tuberosum and S. brevidens. Am J Potato Res 79:19–24
Menke U, Schilde-Rentschler L, Ruoss B, Zanke C, Hemleben V, Ninnemann H (1996) Somatic hybrids between the cultivated potato Solanum tuberosum L. and the 1EBN wild species Solanum pinnatisectum Dun.: morphological and molecular characterization. Theor Appl Genet 92:617–626
Mojtahedi H, Brown CR, Santo GS (1995) Characterization of resistance in a somatic hybrid of Solanum bulbocastanum and S. tuberosum to Meloidogyne chitwoodi. J Nematology 27:86–93
Möllers C, Wenzel G (1992) Somatic hybridization of dihaploid potato protoplasts as a tool for potato breeding. Botanica Acta 105:133–139
Möllers C, Frei U, Wenzel G (1994) Field evaluation of tetraploid somatic potato hybrids. Theor Appl Genet 88:147–152
Naess SK, Bradeen JM, Wielgus SM, Haberlach GT, McGrath JM, Helgeson JP (2000) Resistance to late blight in Solanum bulbocastanum is mapped to chromosome 8. Theor Appl Genet 101:697–704
Naess SK, Bradeen JM, Wielgus SM, Haberlach GT, McGrath JM, Helgeson JP (2001) Analysis of the introgression of Solanum bulbocastaum DNA into potato breeding lines. Mol Genet Genomics 265:694–704
Nouri-Ellouz O, Gargouri-Bouzid R, Sihachakr D, Triki MA, Ducreux G, Drira N, Lakhoua L (2006) Production of potato intraspecific somatic hybrids with improved tolerance to PVY and Pythium aphanidermatum. J. Plant Physiol 163:1321–1332
Nouri-Ellouz O, Triki MA, Jbir-Koubaa R, Louhichi A, Charfeddine S, Drira N, Gargouri-Bouzid R (2016) Somatic hybrids between potato and S. berthaultii show partial resistance to soil-borne fungi and potato virus Y. J Phytopathol 164:485–496
Novy RG, Helgeson JE (1994a) Somatic hybrids between Solanum etuberosum and diploid, tuber bearing Solanum clones. Theor Appl Genet 89:775–782
Novy RG, Helgeson JP (1994b) Resistance to potato virus Y in somatic hybrids between Solanum etuberosum and S. tuberosum × S. berthaultii hybrid. Theor Appl Genet 89:783–786
Novy RG, Nasruddin A, Ragsdale DW, Radcliffe EB (2002) Genetic resistances to potato leafroll virus, potato virus Y, and green peach aphid in progeny of Solanum etuberosum. Am. J Potato Res 79:9–18
Novy RG, Gillen AM, Whitworth JL (2007) Characterization of the expression and inheritance of potato leafroll virus (PLRV) and potato virus Y (PVY) resistance in three generations of germplasm derived from Solanum etuberosum. Theor Appl Genet 114:1161–1172
Nyman M, Waara S (1997) Characterisation of somatic hybrids between Solanum tuberosum and its frost-tolerant relative Solanum commersonii. Theor Appl Genet 95:1127–1132
Oberwalder B, Ruoâ B, Schilde-Rentschler L, Hemleben V, Ninnemann H (1997) Asymmetric protoplast fusion between wild and cultivated species of potato (Solanum ssp.): detection of asymmetric hybrids and genome elimination. Theor Appl Genet 94:1104–1112
Oberwalder B, Schilde-Rentschler L, Ruoâ B, Wittemann S, Ninnemann H (1998) Asymmetric protoplast fusion between wild species and breeding lines of potato-effect of recipient and genome stability. Theor Appl Genet 97:1347–1354
Oberwalder B, Schilde-Rentschler L, Loffelhardt-Ruob B, Ninnemann H (2000) Differences between hybrids of Solanum tuberosum L. and Solanum circaeifolium Bitt. obtained from symmetric and asymmetric fusion experiments. Potato Res 43:71–82
Orczyk W, Przetakiewicz J, Nadoloska-Orczyk A (2003) Somatic hybrids of Solanum tuberosum-application to genetics and breeding. Plant Cell Tissue Organ Cult 74:1–13
Perl A, Aviv D, Galun E (1990) Potato fusion derived CMS potato cybrids: potential seed parents for hybrid, true potato seeds. J Heredity 81:438–442
Polgar Z, Wielgus SM, Horvath S, Helgeson JP (1999) DNA analysis of potato + Solanum brevidens somatic hybrid lines. Euphytica 105:103–107
Polzerová H, Patzak J, Greplová M (2011) Early characterization of somatic hybrids from symmetric protoplast electrofusion of Solanum pinnatisectum Dun. and Solanum tuberosum L. Plant Cell Tissue Organ Cult 104:163–170
Preiszner J, Feher A, Veisz O, Sutka J, Dudits D (1991) Characterization of morphological variation and cold resistance in interspecific somatic hybrids between potato (Solanum tuberosum L.) and S. brevidens Phil. Euphytica 57:37–49
Przetakiewicz J, Nadolska-Orczyk A, Kuc D, Orczyk W (2007) Tetraploid somatic hybrids of potato (Solanum tuberosum L.) obtained from diploid breeding lines. Cell Mol Biol Lett 12:253–267
Puite KJ, Roest S, Pijnacker LP (1986) Somatic hybrid potato plants after electrofusion of diploid Solanum tuberosum and Solanum phureja. Plant Cell Rep 5:262–265
Rakosy-Tican E, Aurori A (2015) Green fluorescent protein (GFP) supports the selection based on callus vigorous growth in the somatic hybrids Solanum tuberosum L. + S. chacoense Bitt. Acta Physiol Plant 37:201
Rakosy-Tican E, Thieme R, Nachtigall M, Molnar I, Denes T-E (2015) The recipient potato cultivar influences the genetic makeup of the somatic hybrids between five potato cultivars and one cloned accession of sexually incompatible species Solanum bulbocastanum Dun. Plant Cell Tissue Organ Cult 122:395–407
Rasmussen JO, Nepper JP, Rasmussen OS (1996) Analysis of somatic hybrids between two sterile dihaploid Solanum tuberosum L. breeding lines. Restoration of fertility and complementation of G. pallida Pa2 and Pa3 resistance. Theor Appl Genet 92:403–410
Rasmussen JO, Waara S, Rasmussen OS (1997) Regeneration and analysis of interspecific asymmetric potato-Solanum ssp hybrid plants selected by micromanipulation or fluorescence activated cell sorting (FACS). Theor Appl Genet 95:41–49
Rasmussen JO, Nepper JP, Kirk H-G, Tolstrup K, Rasmussen OS (1998) Combination of resistance to potato late blight in foliage and tubers by interspecific dihaploid protoplast fusion. Euphytica 102:363–370
Rasmussen JO, Lössl A, Rasmussen OS (2000) Analysis of the plastome and chondriome origin in plants regenerated after asymmetric Solanum ssp. protoplast fusions. Theor Appl Genet 101:336–343
Rokka V-M, Xu Y-S, Kankila J, Kuusela A, Pulli S, Pehu E (1994) Identification of somatic hybrids of dihaploid Solanum tuberosum lines and S. brevidens by species specific RAPD patterns and assessment of disease resistance of the hybrids. Euphytica 80:207–217
Rokka V-M, Tauriainen A, Pietilä L, Pehu E (1998) Interspecific somatic hybrids between wild potato Solanum acaule Bitt. and anther-derived dihaploid potato (Solanum tuberosum L.). Plant Cell Rep 18:82–88
Rokka V-M, Valkonen JPT, Tauriainen A, Pietila L, Lubecka R, Zimnoch-Guzowska E, Pehu E (2000) Production and characterization of ‘second generation’ of somatic hybrids derived from protoplast fusion between interspecific somatohaploid and dihaploid Solanum tuberosum L. Am J Potato Res 77:149–159
Rokka V-M, Laurila J, Tauriainen A, Laakso I, Larkka J, Metzler M, Pietiä L (2005) Glycoalkaloid aglycone accumulations associated with infection by Clavibacter michiganensis ssp. sepedonicus in potato species Solanum acaule and Solanum tuberosum and their interspecific somatic hybrids. Plant Cell Rep 23:683–691
Sarkar D, Tiwari JK, Sharma SH, Poonam, Sharma SA, Gopal J, Singh BP, Luthra SK, Pandey SK, Pattanayak D (2011) Production and characterization of somatic hybrids between Solanum tuberosum L. and S. pinnatisectum Dun. Plant Cell Tissue Organ Cult 107:427–440
Scotti N, Monti L, Cardi T (2003) Organelle DNA variation in parental Solanum spp. genotypes and nuclear-cytoplasmic interactions in Solanum tuberosum (+) S. commersonii somatic hybrid-backcross progeny. Theor Appl Genet 108:87–94
Scotti N, Marechal-Drouard L, Cardi T (2004) The rpl5–rps14 mitochondrial region: a hot spot for DNA rearrangements in Solanum spp. somatic hybrids. Curr Genet 45:378–382
Serraf I, Sihachakr D, Ducreux G, Brown SC, Allot M, Barghi N, Rossignol L(1991) Interspecific somatic hybridization in potato by protoplast electrofusion. Plant Sci 76: 115–126
Sharma S, Sarkar D, Pandey SK, Chandel P, Tiwari JK (2011) Stoloniferous shoot protoplast, an efficient cell system in potato for somatic cell genetic manipulations. Sci Hortic 128:84–91
Shi YZ, Chen Q, Li HY, Beasley D, Lynch DR (2006) Somatic hybridization between Solanum tuberosum and S. cardiophyllum. Can J Plant Sci 86:539–545
Sidorov VA, Zubko MK, Kuchko AA, Komarnitsky IK, Gleba YY (1987) Somatic hybridization in potato: use of γ-irradiated protoplasts of Solanum pinnatisectum in genetic reconstruction. Theor Appl Genet 74:364–368
Singh R, Tiwari JK, Rawat S, Sharma V, Singh BP (2016a) Monitoring gene expression pattern in somatic hybrid of Solanum tuberosum and S. pinnatisectum for late blight resistance using microarray analysis. Plant Omics J 9:99–105
Singh R, Tiwari JK, Rawat S, Sharma V, Singh BP (2016b) In silico identification of candidate microRNAs and their targets in potato somatic hybrid Solanum tuberosum (+) S. pinnatisectum for late blight resistance. Plant Omics J 9:159–164
Smyda P, Jakuczun H, Debski K, Śliwka J, Thieme R, Nachtigall M, Wasilewicz-Flis I, Zimnoch-Guzowska E (2013) Development of somatic hybrids Solanum × michoacanum Bitter. (Rydb.) (+) S. tuberosum L. and autofused 4x S. × michoacanum plants as potential sources of late blight resistance for potato breeding. Plant Cell Rep 32:1231–1241
Smyda-Dajmund P, Sliwka J, Wasilewicz-Flis I, Jakuczun H, Zimnoch-Guzowska E (2016) Genetic composition of interspecific potato somatic hybrids and autofused 4x plants evaluated by DArT and cytoplasmic DNA markers. Plant Cell Rep 35:1345–1358
Smyda-Dajmund P, Śliwka J, Wasilewicz-Flis I, Jakuczun H, Zimnoch-Guzowska E (2017) BC1 and F1 progeny from Solanum × michoacanum (+) S. tuberosum somatic hybrids, autofused 4 × S. michoacanum and cultivated potato. Am J Potato Res. https://doi.org/10.1007/s12230-017-9568-2
Song J, Bradeen JM, Naess SK, Raasch JA, Wielgus SM, Haberlach GT, Liu J, Kuang H, Austin-Phillips S, Buell CR, Helgeson JP, Jiang J (2003) Gene RB cloned from Solanum bulbocastanum confers broad spectrum resistance to potato late blight. Proc Natl Acad Sci USA 100:9128–9133
Spooner DM, Salas A (2006) Structure, biosystematics, and genetic resources. In: Gopal J, Khurana SM, Paul (eds) Handbook of potato production, improvement and postharvest management. Food Product Press, New York, pp 1–40
Szczerbakowa A, Maciejewska U, Wielgat B (2000) Plant regeneration from the protoplasts of Solanum tuberosum, S. nigrum and S. bulbocastanum. Acta Physiol Plant 22:3–10
Szczerbakowa A, Maciejewska U, Pawlowski P, Skierski JS, Wielgat B (2001) Electrofusion of protoplasts from Solanum tuberosum, S. nigrum and S. bulbocastanum. Acta Physiol Plant 23:169–179
Szczerbakowa A, Boltowicz D, Wielgat B (2003a) Interspecific somatic hybrids Solanum bulbocastanum (+) S. tuberosum H-8105. Acta Physiol Plant 25:365–373
Szczerbakowa A, Maciejewska U, Zimnoch-Guzowska E, Wielgat B (2003b) Somatic hybrids Solanum nigrum (+) S. tuberosum: morphological assessment and verification of hybridity. Plant Cell Rep 21:577–584
Szczerbakowa A, Boltowicz D, Lebecka R, Radomski P, Wielgat B (2005) Characteristics of the interspecific somatic hybrids Solanum pinnatisectum (+) S. tuberosum H-8105. Acta Physiol Plant 27:265–273
Szczerbakowa A, Tarwacka J, Oskiera M, Jakuczun H, Wielgat B (2010) Somatic hybridization between the diploids of S. × michoacanum and S. tuberosum. Acta Physiol Plant 32:867–873
Szczerbakowa A, Tarwacka J, Sliwinska E, Wielgat B (2011) Nuclear DNA content and chromosome number in somatic hybrid allopolyploids of Solanum. Plant Cell Tissue Organ Cult 106:373–380
Takebe I, Labib G, Melchers G (1971) Regeneration of whole plants from isolated mesophyll protoplasts of tobacco. Naturwissenschaften 58:318–320
Tarwacka J, Polkowska-Kowalczyk L, Kolano B, Śliwka J, Wielgat B (2013) Interspecific somatic hybrids Solanum villosum (+) S. tuberosum, resistant to Phytophthora infestans. J Plant Physiol 170:1541–1548
Tek AL, Stevenson WR, Helgeson JP, Jiang J (2004) Transfer of tuber soft rot and early blight resistances from Solanum brevidens into cultivated potato. Theor Appl Genet 109:249–254
Thach NQ, Frei U, Wenzel G (1993) Somatic fusion for combining virus resistances in Solanum tuberosum L. Theor Appl Genet 85:863–867
Thieme R, Darsow U, Gavrilenko T, Dorokhov D, Tiemann H (1997) Production of somatic hybrids between S. tuberosum L. and late blight resistant Mexican wild potato species. Euphytica 97:189–200
Thieme R, Gavrilenko T, Thieme T, Heimbach U (1999) Production of potato genotypes with resistance to potato virus Y by biotechnological methods. In: Altmann A et al (eds), Plant biotechnology and in vitro biology in the 21st century. Kluwer Academic Publishers, Dordrecht, pp 557–560
Thieme R, Darsow U, Rakosy-Tican L, Kang Z, Gavrilenko T, Antonova O, Heimbach U, Thieme T (2004) Use of somatic hybridization to transfer resistance to late blight and potato virus Y (PVY) into cultivated potato. Plant Breed Seed Sci 50:113–118
Thieme R, Rakosy-Tican E, Gavrilenko T, Antonova O, Schubert J, Nachtigall M, Heimbach U, Thieme T (2008) Novel somatic hybrids (Solanum tuberosum L. + Solanum tarnii) and their fertile BC1 progenies express extreme resistance to potato virus Y and late blight. Theor Appl Genet 116:691–700
Thieme R, Rakosy-Tican E, Nachtigall M, Schubert J, Hammann T, Antonova O, Gavrilenko T, Heimbach U, Thieme T (2010) Characterization of the multiple resistance traits of somatic hybrids between Solanum cardiophyllum Lindl. and two commercial potato cultivars. Plant Cell Rep 29:1187–1201
Tiwari JK, Poonam, Sarkar D, Pandey SK, Gopal J, Kumar SR (2010) Molecular and morphological characterization of somatic hybrids between Solanum tuberosum L. and S. etuberosum Lindl. Plant Cell Tissue Organ Cult 103:175–187
Tiwari JK, Poonam, Kumar V, Singh BP, Sharma S, Luthra SK, Bhardwaj V (2013a) Evaluation of potato somatic hybrids of dihaploid S. tuberosum (+) S. pinnatisectum for late blight resistance. Potato J 40:176–179
Tiwari JK, Saurabh S, Chandel P, Singh BP, Bhardwaj V (2013b) Analysis of genetic and epigenetic variation in potato somatic hybrid by AFLP and MASP markers. Electron J Biotechnol. https://doi.org/10.2225/vol16-issue6-fulltext-9
Tiwari JK, Chandel P, Singh BP, Bhardwaj V (2014) Analysis of plastome and chondriome genome types in potato somatic hybrids from Solanum tuberosum x Solanum etuberosum. Genome 57:29–35
Tiwari JK, Devi S, Sharma S, Chandel P, Rawat S, Singh BP (2015a) Allele mining in Solanum germplasm: cloning and characterization of RB-homologous gene fragments from late blight resistant wild potato species. Plant Mol Biol Rep 33:1584–1598
Tiwari JK, Devi S, Sundaresha S, Chandel P, Ali N, Singh B, Bhardwaj V, Singh BP (2015b) Microarray analysis of gene expression patterns in the leaf during potato tuberization in the potato somatic hybrid Solanum tuberosum and Solanum etuberosum. Genome 58:305–313
Tiwari JK, Poonam, Saurabh S, Devi S, Ali N, Bhardwaj V, Singh BP (2015c) Molecular characterization of potato somatic hybrids by inter simple sequence repeat (ISSR) markers. Potato J 42:1–7
Tiwari JK, Saurabh S, Chandel P, Singh BP, Bhardwaj V (2015d) Assessment of genetic and epigenetic variations in potato somatic hybrids by methylation-sensitive ISSR and RAPD markers. Bangladesh J Bot 44:45–50
Tiwari JK, Saurabh S, Chandel P, Devi S, Ali N, Bist CM, Singh BP (2015e) Analysis of genetic and epigenetic changes in potato somatic hybrids between Solanum tuberosum and S. etuberosum by AFLP and MSAP markers. Agric Res 4:339–346
Tiwari JK, Devi S, Chandel P, Ali N, Bhardwaj V, Singh BP (2016) Organelle genome analysis in somatic hybrids between Solanum tuberosum and S. pinnatisectum revealed diverse cytoplasm type in potato. Agric Res 5:22–28
Trabelsi S, Gargouri-Bouzid R, Vedel F, Nato A, Lakhoua L, Drira N (2005) Somatic hybrids between potato Solanum tuberosum and wild species Solanum verneï exhibit a recombination in the plastome. Plant Cell Tissue Organ Cult 83:1–11
Valkonen JPT, Rokka V-M (1998) Combination and expression of two virus resistance mechanisms in interspecific somatic hybrids of potato. Plant Sci 131:85–94
Waara S, Glimelius K (1995) The potential of somatic hybridization in crop breeding. Euphytica 85:217–233
Waara S, Nyman M, Johannisson A (1998) Efficient selection of potato heterokaryons by flow cytometric sorting and the regeneration of hybrid plants. Euphytica 101:293–299
Wallin A, Glimelius K, Eriksson T (1974) The induction of aggregation and fusion of Daucus carota protoplasts by polyethylene glycol. Z Pflanzenphysiol 74:64–80
Wang J, Jiang J, Wang Y (2013) Protoplast fusion for crop improvement and breeding in China. Plant Cell Tissue Organ Cult 112:131–142
Williams CE, Wielgus SM, Haberlach GT, Guenther C, Kim-Lee H, Helgeson JP (1993) RFLP analysis of chromosomal segregation in progeny from an interspecific hexaploid somatic hybrid between Solanum brevidens and Solanum tuberosum. Genetics 135:1167–1173
Xu YS, Murto M, Dunckley R, Jones MGK, Pehu E (1993) Production of asymmetric hybrids between Solanum tuberosum and irradiated S. brevidens. Theor Appl Genet 85:729–734
Yamada T, Misoo S, Ishii T, Ito Y, Takaoka K, Kamijima O (1997) Characterization of somatic hybrids between tetraploid Solanum tuberosum L. and dihaploid S. acaule. Breed Sci 47:229–236
Yamada T, Hosaka K, Kaide N, Nakagawa K, Misoo S, Kamijima O (1998) Cytological and molecular characterization of BC1 progeny from two somatic hybrids between dihaploid Solanum acaule and tetraploid S. tuberosum. Genome 41:743–750
Yu Y, Ye W, He L, Cai X, Liu T, Liu J (2013) Introgression of bacterial wilt resistance from eggplant to potato via protoplast fusion and genome components of the hybrids. Plant Cell Rep 32:1687–1701
Zimnoch-Guzowska E, Lebecka R, Kryszczuk A, Maciejewska U, Szczerbakowa A, Wielgat B (2003) Resistance to Phytophthora infestans in somatic hybrids of Solanum nigrum L. and diploid potato. Theor Appl Genet 107:43–48
Acknowledgements
JKT thanks to the DBT project (BT/PR4291/AGR/2/838/2011), the CABin project and Biotechnology program (HORTCPRICIL201500300131), ICAR-CPRI, Shimla for necessary supports. We are grateful to the Editors and anonymous reviewers for valuable inputs to improve the manuscript.
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JKT conceived the idea, collected literature and wrote the manuscript; JKT, SD, NA, SKL, VK contributed in development of interspecific somatic hybrids and literature collection; VB, RKS, SKC corrected the manuscript; and SR contributed in genomics work. All authors read and approved the manuscript.
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Communicated by Sergio J. Ochatt.
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Tiwari, J.K., Devi, S., Ali, N. et al. Progress in somatic hybridization research in potato during the past 40 years. Plant Cell Tiss Organ Cult 132, 225–238 (2018). https://doi.org/10.1007/s11240-017-1327-z
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DOI: https://doi.org/10.1007/s11240-017-1327-z