Somaclonal variation in plants: Causes and detection methods

Plant Growth Regulation

Volumn 63, Issue 2, 2011, Pages 147-173

  Bairu, Michael W ^a   Aremu, Adeyemi O ^a   van Staden, Johannes ^a  

^a UNIVERSITY OF KWAZULU NATAL   (South Africa)

Author keywords

Cell and tissue culture; Cell cycle; Chimeras; Chromosome aberration; Cryptic transposable elements; Induced variation

Indexed keywords

EID: 79952184595     PISSN: 01676903     EISSN: 15735087     Source Type: Journal    
DOI: 10.1007/s10725-010-9554-x     Document Type: Article

References (305)

1. Adir N, Zer H, Shochat S, Ohad I (2003) Photoinhibition-a historical perspective. Photosynth Res 76: 343-370.
2. Agarwal M, Shrivastava N, Padh H (2008) Advances in molecular marker techniques and their applications in plant sciences. Plant Cell Rep 27: 617-631.
3. Ahloowalia BS (1986) Limitations to the use of somaclonal variation in crop improvement. In: Semal J (ed) Somaclonal variations and crop improvement. Martinus Nijhoff Publishers, Dordrecht, pp 14-27.
4. Ahmed K, Abdelkareem A (2005) Somaclonal variation in bread wheat (Triticum aestivum L.). II. Field performance of somaclones. Cereal Res Commun 33: 485-492.
5. Ahmed EU, Hayashi T, Yazawa S (2004) Auxins increase the occurrence of leaf-colour variants in Caladium regenerated from leaf explants. Sci Hortic 100: 153-159.
6. Albani MC, Wilkinson MJ (1998) Inter simple sequence repeat polymerase chain reaction for the detection of somaclonal variation. Plant Breed 117: 573-575.
7. Althoff DM, Gitzendanner MA, Segraves KA (2007) The utility of amplified fragment length polymorphisms in phylogenetics: a comparison of homology within and between genomes. Syst Biol 56: 477-484.
8. Al-Zahim MA, Ford-Lloyd BV, Newbury HJ (1999) Detection of somaclonal variation in garlic (Allium sativum L.) using RAPD and cytological analysis. Plant Cell Rep 18: 473-477.
9. Anastassopoulos E, Keil M (1996) Assessment of natural and induced genetic variation in Alstroemeria using random amplified polymorphic DNA (RAPD) markers. Euphytica 90: 235-244.
10. Araújo LG, Prabhu AS, Filippi MC, Chaves LJ (2001) RAPD analysis of blast resistant somaclones from upland rice cultivar IAC 47 for genetic divergence. Plant Cell Tissue Organ Cult 67: 165-172.
11. Arihara A, Kita T, Igarashi S, Goto M, Irikura Y (1995) White Baron: a non-browning somaclonal variant of Danshakuimo (Irish cobbler). Am J Potato Res 72: 701-705.
12. Arun B, Joshi AK, Chand R, Singh BD (2003) Wheat somaclonal variants showing earliness, improved spot blotch resistance and higher yield. Euphytica 132: 235-241.
13. Arun B, Singh BD, Sharma S, Paliwal R, Joshi AK (2007) Development of somaclonal variants of wheat (Triticum aestivum L.) for yield traits and disease resistance suitable for heat stressed and zero-till conditions. Field Crops Res 103: 62-69.
14. Asif MJ, Othman RY (2005) Characterization of fusarium wilt-resistant and fusarium wilt-susceptible somaclones of banana cultivar rastali (Musa AAB) by random amplified polymorphic DNA and retrotransposon markers. Plant Mol Biol Rep 23: 241-249.
15. Aversano R, Savarese S, Maria De Nova J, 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.
16. Baer G, Yemets A, Stadnichuk N, Rakhmetov D, Blume Y (2007) Somaclonal variability as a source for creation of new varieties of finger millet (Eleusine coracana (L.) Gaertn.). Cytol Genet 41: 204-208.
17. Bairu MW, Fennell CW, van Staden J (2006) The effect of plant growth regulators on somaclonal variation in Cavendish banana (Musa AAA cv. 'Zelig'). Sci Hortic 108: 347-351.
18. Bajaj YPS (1990) Somaclonal variation-origin, induction, cryopreservation, and implications in plant breeding. In: Bajaj YPS (ed) Biotechnology in agriculture and forestry II. Somaclonal variation in crop improvement. Springer, Berlin, pp 3-35.
19. Baránek M, Křižan B, Ondrušíková E, Pidra M (2010) DNA-methylation changes in grapevine somaclones following in vitro culture and thermotherapy. Plant Cell Tissue Organ Cult 101: 11-22.
20. Barret P, Brinkman M, Beckert M (2006) A sequence related to rice Pong transposable element displays transcriptional activation by in vitro culture and reveals somaclonal variations in maize. Genome 49: 1399-1407.
21. Barwale UB, Widholm JM (1987) Somaclonal variation in plants regenerated from cultures of soybean. Plant Cell Rep 6: 365-368.
22. Bayliss MW (1980) Chromosomal variation in tissue culture. In: Vasil IK (ed) Perspectives in plant cell and tissue culture. Academic Press, New York, pp 113-144.
23. Bednarek P, Orlowska R, Koebner R, Zimny J (2007) Quantification of the tissue-culture induced variation in barley (Hordeum vulgare L.). BMC Plant Biol 7: 10. doi: 10. 1186/1471-2229-7-10.
24. Beemster GTS, Fiorani F, Inzé D (2003) Cell cycle: the key to plant growth control? Trends Plant Sci 8: 154-158.
25. Belaj A, Satovic Z, Cipriani G, Baldoni L, Testolin R, Rallo L, Trujillo I (2003) Comparative study of the discriminating capacity of RAPD, AFLP and SSR markers and of their effectiveness in establishing genetic relationships in olive. Theor Appl Genet 107: 736-744.
26. Bennici A, Anzidei M, Vendramin GG (2004) Genetic stability and uniformity of Foeniculum vulgare Mill. regenerated plants through organogenesis and somatic embryogenesis. Plant Sci 166: 221-227.
27. Bernardo Royo J, Itoiz R (2004) Evaluation of the discriminance capacity of RAPD, isoenzymes and morphologic markers in apple (Malus × domestica Borkh.) and the congruence among classifications. Genet Resour Crop Evol 51: 153-160.
28. Bhat KV, Bhat SR, Chandel KPS (1992) Survey of isozyme polymorphism for clonal identification in Musa. I. Esterase, acid phosphate and catalase. J Hortic Sci 67: 501-507.
29. Bhatia R, Singh KP, Jhang T, Sharma TR (2009) Assessment of clonal fidelity of micropropagated gerbera plants by ISSR markers. Sci Hortic 119: 208-211.
30. Bhattacharya S, Dey T, Bandopadhyay T, Ghosh P (2008) Genetic polymorphism analysis of somatic embryo-derived plantlets of Cymbopogon flexuosus through RAPD assay. Plant Biotechnol Rep 2: 245-252.
31. Biswas MK, Dutt M, Roy UK, Islam R, Hossain M (2009) Development and evaluation of in vitro somaclonal variation in strawberry for improved horticultural traits. Sci Hortic 122: 409-416.
32. Bogdanova ED (2003) Epigenetic variation induced in Triticum aestivum L. by nicotinic acid. Russ J Genet 39: 1029-1034.
33. Bonner JE, Warner RM, Brewbaker JL (1974) A chemosystematic study of Musa cultivars. HortScience 9: 325-328.
34. Bordallo PN, Silva DH, Maria J, Cruz CD, Fontes EP (2004) Somaclonal variation on in vitro callus culture potato cultivars. Hort Brasil 22: 300-304.
35. Botstein D, White RL, Skolnick M, Davis RW (1980) Construction of a genetic linkage map in man using restriction fragment length polymorphisms. Am J Hum Genet 32: 314-331.
36. Bouharmont J (1994) Application of somaclonal variation and in vitro selection to plant improvement. Acta Hortic 355: 213-218.
37. Bouman H, De Klerk GJ (2001) Measurement of the extent of somaclonal variation in begonia plants regenerated under various conditions. Comparison of three assays. Theor Appl Genet 102: 111-117.
38. Brar DS, Jain SM (1998) Somaclonal variation: mechanism and applications in crop improvement. In: Jain SM, Brar DS, Ahloowalia BS (eds) Somaclonal variation and induced mutations in crop improvement. Kluwer Academic Publishers, Dordrecht, pp 15-37.
39. Braun AC (1959) A demonstration of the recovery of the crown-gall tumor cell with the use of complex tumors of single-cell origin. Proc Natl Acad Sci USA 45: 932-938.
40. Bright S, Jarret V, Nelson R, Creissen G, Karp A, Franklin J, Norbury P, Kueh J, Rognes S, Miflin B (1983) Modification of agronomic traits using in vitro technology. In: Mantell SH, Smith H (eds) Plant biotechnology. Cambridge University Press, New York, pp 251-265.
41. Brutovská R, Čellárová E, Doležel J (1998) Cytogenetic variability of in vitro regenerated Hypericum perforatum L. plants and their seed progenies. Plant Sci 133: 221-229.
42. Bryant JA (1976) The cell cycle. In: Bryant JA (ed) Molecular aspect of gene expression in plants. Academic Press, New York, pp 117-216.
43. Caetano-Anollés G, Bassam B (1993) DNA amplification fingerprinting using arbitrary oligonucleotide primers. Appl Biochem Biotechnol 42: 189-200.
44. Campbell B, LeMare S, Piperidis G, Godwin I (2010) IRAP, a retrotransposon-based marker system for the detection of somaclonal variation in barley. Mol Breed. doi: 10. 1007/s11032-010-9422-4.
45. Carsono N, Yoshida T (2007) Variation in spikelet-related traits of rice plants regenerated from mature seed-derived callus culture. Plant Prod Sci 10: 86-90.
46. Cassells AC, Walsh M (1995) Screening for Sclerotinia resistance in Helianthus tuberosus L. (Jerusalem artichoke) varieties, lines and somaclones, in the field and in vitro. Plant Pathol 44: 428-437.
47. Cassells AC, Walsh C, Periappuram C (1993) Diplontic selection as a positive factor in determining the fitness of mutants of Dianthus 'Mystère' derived from x-irradiation of nodes in in vitro culture. Euphytica 70: 167-174.
48. Chandrika M, Thoyajaksha, Ravishankar Rai V, Ramachandra Kini K (2008) Assessment of genetic stability of in vitro grown Dictyospermum ovalifolium. Biol Plant 52: 735-739.
49. Chandrika M, Ravishankar Rai V, Thoyajaksha (2010) ISSR marker based analysis of micropropagated plantlets of Nothapodytes foetida. Biol Plant 54: 561-565.
50. Chen WH, Chen TM, Fu YM, Hsieh RM, Chen WS (1998) Studies on somaclonal variation in Phalaenopsis. Plant Cell Rep 18: 7-13.
51. Chen J, Henny R, Devanand P, Chao C (2006) AFLP analysis of nephthytis (Syngonium podophyllum Schott) selected from somaclonal variants. Plant Cell Rep 24: 743-749.
52. Chintapalli P, Moss J, Sharma K, Bhalla J (1997) In vitro culture provides additional variation for pigeonpea. In Vitro Cell Dev Biol Plant 33: 30-37.
53. Choi H-W, Lemaux PG, Cho M-J (2001) High frequency of cytogenetic aberration in transgenic oat (Avena sativa L.) plants. Plant Sci 160: 763-772.
54. Chomátová S, Turková V, Klozová E (1990) Protein complex and esterase isoenzyme patterns of Allium sativum L. cultivars and clones-regenerants. Biol Plant 32: 321-331.
55. Chuang SJ, Chen CL, Chen JJ, Chou WY, Sung JM (2009) Detection of somaclonal variation in micro-propagated Echinacea purpurea using AFLP marker. Sci Hortic 120: 121-126.
56. Claxton J, Arnold D, Clarkson J, Blakesley D (1998) The regeneration and screening of watercress somaclones for resistance to Spongospora subterranea f. sp. nasturtii and measurement of somaclonal variation. Plant Cell Tissue Organ Cult 52: 155-164.
57. Cloutier S, Landry B (1994) Molecular markers applied to plant tissue culture. In Vitro Cell Dev Biol Plant 30: 32-39.
58. Coggins LW, O'Prey M (1989) DNA tertiary structures formed in vitro by misaligned hybridization of multiple tandem repeat sequences. Nucleic Acids Res 17: 7417-7426.
59. Cooper C, Crowther T, Smith BM, Isaac S, Collin HA (2006) Assessment of the response of carrot somaclones to Pythium violae, causal agent of cavity spot. Plant Pathol 55: 427-432.
60. Côte F, Teisson C, Perrier X (2001) Somaclonal variation rate evolution in plant tissue culture: contribution to understanding through a statistical approach. In Vitro Cell Dev Biol Plant 37: 539-542.
61. D'Amato F (1975) The problem of genetic stability in plant tissue and cell cultures. In: Frankel OH, Hawkes JG (eds) Crop genetic resources for today and tomorrow. Cambridge University Press, New York, pp 333-348.
62. D'Amato F (1977) Cytogenetics of differentiation in tissue and cell culture. In: Reinert J, Bajaj YPS (eds) Applied and fundamental aspects of plant cell, tissue and organ culture. Springer, New York, pp 343-464.
63. Damasco OP, Godwin ID, Smith MK, Adkins SW (1996) Gibberellic acid detection of dwarf off-types in micropropagated Cavendish bananas. Aust J Exp Agric 36: 237-341.
64. Damasco OP, Smith MK, Godwin ID, Adkins SW, Smillie RM, Hetherington SE (1997) Micropropagated dwarf off-type Cavendish bananas (Musa spp., AAA) show improved tolerance to suboptimal temperatures. Aust J Agric Res 48: 377-384.
65. Damasco OP, Smith MK, Adkins SW, Hetherington SE, Godwin ID (1998) Identification and characterisation of dwarf off-types from micropropagated 'Cavendish' bananas. Acta Hortic 490: 79-84.
66. Daub ME (1986) Tissue culture and the selection of resistance to pathogens. Annu Rev Phytopathol 24: 159-186.
67. Daub ME, Jenns AE (1989) Field and greenhouse analysis of variation for disease resistance in tobacco somaclones. Phytopathology 79: 600-605.
68. De la Puente R, González A, Ruiz M, Polanco C (2008) Somaclonal variation in rye (Secale cereale L.) analyzed using polymorphic and sequenced AFLP markers. In Vitro Cell Dev Biol Plant 44: 419-426.
69. Dennis ES, Brettell RIS, Peacock WJ (1987) A tissue culture induced Adh1 null mutant of maize results from a single base change. Mol Gen Genet 210: 181-183.
70. Devarumath RM, Nandy S, Rani V, Marimuthu S, Muraleedharan N, Raina SN (2002) RAPD, ISSR and RFLP fingerprints as useful markers to evaluate genetic integrity of micropropagated plants of three diploid and triploid elite tea clones representing Camellia sinensis (China type) and C. assamica ssp. assamica (Assam-India type). Plant Cell Rep 21: 166-173.
71. Devarumath RM, Doule RB, Kawar PG, Naikebawane SB, Nerkar YS (2007) Field performance and RAPD analysis to evaluate genetic fidelity of tissue culture raised plants vis-à-vis conventional setts derived plants of sugarcane. Sugar Tech 9: 17-22.
72. Do G-S, Seo B-B, Ko J-M, Lee S-H, Pak J-H, Kim I-S, Song S-D (1999) Analysis of somaclonal variation through tissue culture and chromosomal localization of rDNA sites by fluorescent in situ hybridization in wild Allium tuberosum and a regenerated variant. Plant Cell Tissue Organ Cult 57: 113-119.
73. Doležel J (1997) Application of flow cytometry for the study of plant genomes. J Appl Genet 38: 285-302.
74. Doležel J, Bartoš JAN (2005) Plant DNA flow cytometry and estimation of nuclear genome size. Ann Bot 95: 99-110.
75. Doležel J, Valárik M, Vrána J, Lysák MA, Hřibová E, Bartoš J, Gasmanová N, Doleželová M, Šafář J, Šimková H (2004) Molecular cytogenetics and cytometry of bananas (Musa spp.). In: Jain SM, Swennen R (eds) Banana improvement: cellular, molecular biology, and induced mutations. Science Publishers, Inc., Enfield, pp 229-244.
76. Dugdale LJ, Mortimer AM, Isaac S, Collin HA (2000) Disease response of carrot and carrot somaclones to Alternaria dauci. Plant Pathol 49: 57-67.
77. Eastwood RF, Kollmorgen JF, Hannah M, Williams WM (1994) Reaction of somaclonal variants of wheat to the take-all fungus (Gaeumannomyces graminis var. tritici). Plant Pathol 43: 644-650.
78. Ehsanpour AA, Madani S, Hoseini M (2007) Detection of somaclonal variation in potato callus induced by UV-C radiation using RAPD-PCR. Gen Appl Plant Physiol 33: 3-11.
79. Eizenga GC, Cornelius PL (1991) Comparison of the isozyme variation in tall fescue parents and their somaclones. Euphytica 51: 249-256.
80. El-Sayed OE, Rizkalla AA, Sabri SRS (2007) In vitro mutagenesis for genetic improvement of salinity tolerance in wheat. Res J Agric Biol Sci 4: 377-383.
81. Etienne H, Bertrand B (2003) Somaclonal variation in Coffea arabica: effects of genotype and embryogenic cell suspension age on frequency and phenotype of variants. Tree Physiol 23: 419-426.
82. Evans DA, Sharp WR, Medina-Filho HP (1984) Somaclonal and gametoclonal variation. Am J Bot 71: 759-774.
83. Fiuk A, Bednarek P, Rybczyński J (2010) Flow cytometry, HPLC-RP, and metAFLP analyses to assess genetic variability in somatic embryo-derived plantlets of Gentiana pannonica Scop. Plant Mol Biol Rep 28: 413-420.
84. Fourré JL, Berger P, Niquet L, André P (1997) Somatic embryogenesis and somaclonal variation in Norway spruce: morphogenetic, cytogenetic and molecular approaches. Theor Appl Genet 94: 159-169.
85. Gao D-Y, Vallejo V, He B, Gai Y-C, Sun L-H (2009) Detection of DNA changes in somaclonal mutants of rice using SSR markers and transposon display. Plant Cell Tissue Organ Cult 98: 187-196.
86. Gao X, Yang D, Cao D, Ao M, Sui X, Wang Q, Kimatu J, Wang L (2010) In vitro micropropagation of Freesia hybrida and the assessment of genetic and epigenetic stability in regenerated plantlets. J Plant Growth Regul 29: 257-267.
87. Gengenbach BG, Umpeck P (1982) Characteristics of T-cytoplasm revertants from tissue culture. Maize Genet Coop News Lett 56: 140-142.
88. George EF (1993) Plant propagation by tissue culture, part 1: the technology. Exegetics Ltd, London.
89. Gesteira AS, Otoni WC, Barros EG, Moreira MA (2002) RAPD-based detection of genomic instability in soybean plants derived from somatic embryogenesis. Plant Breed 121: 269-271.
90. Giménez C, de García E, de Enrech NX, Blanca I (2001) Somaclonal variation in banana: cytogenetic and molecular characterization of the somaclonal variant CIEN BTA-03. In Vitro Cell Dev Biol Plant 37: 217-222.
91. Godwin ID, Sangduen N, Kunanuvatchaidach R, Piperidis G, Adkins SW (1997) RAPD polymorphisms among variant and phenotypically normal rice (Oryza sativa var. indica) somaclonal progenies. Plant Cell Rep 16: 320-324.
92. González A, De la Fuente M, De Ron A, Santalla M (2010) Protein markers and seed size variation in common bean segregating populations. Mol Breed 25: 723-740.
93. Gostimsky SA, Kokaeva ZG, Konovalov FA (2005) Studying plant genome variation using molecular markers. Russ J Genet 41: 378-388.
94. Graebe JE (2003) Gibberellin biosynthesis and control. Annu Rev Plant Physiol 38: 419-465.
95. Guo W, Gong L, Ding Z, Li Y, Li F, Zhao S, Liu B (2006a) Genomic instability in phenotypically normal regenerants of medicinal plant Codonopsis lanceolata Benth. et Hook. f., as revealed by ISSR and RAPD markers. Plant Cell Rep 25: 896-906.
96. Guo W, Li Y, Gong L, Li F, Dong Y, Liu B (2006b) Efficient micropropagation of Robinia ambigua var. idahoensis (Idaho Locust) and detection of genomic variation by ISSR markers. Plant Cell Tissue Organ Cult 84: 343-351.
97. Guo W, Wu R, Zhang Y, Liu X, Wang H, Gong L, Zhang Z, Liu B (2007) Tissue culture-induced locus-specific alteration in DNA methylation and its correlation with genetic variation in Codonopsis lanceolata Benth. et Hook. f. Plant Cell Rep 26: 1297-1307.
98. Hadi MZ, Bridgen MP (1996) Somaclonal variation as a tool to develop pest resistant plants of Torenia fournieri 'Compacta Blue'. Plant Cell Tissue Organ Cult 46: 43-50.
99. Hao Y-J, Deng X-X (2002) Occurrence of chromosomal variations and plant regeneration from long-term-cultured citrus callus. In Vitro Cell Dev Biol Plant 38: 472-476.
100. Hartmann C, Henry Y, Buyser J, Aubry C, Rode A (1989) Identification of new mitochondrial genome organizations in wheat plants regenerated from somatic tissue cultures. Theor Appl Genet 77: 169-175.
101. Hashim ZN, Campbell WF, Carman JG (1990) Morphological analyses of spring wheat (CIMMYT cv. PCYT-10) somaclones. Plant Cell Tissue Organ Cult 20: 95-99.
102. Hashmi G, Huettel R, Meyer R, Krusberg L, Hammerschlag F (1997) RAPD analysis of somaclonal variants derived from embryo callus cultures of peach. Plant Cell Rep 16: 624-627.
103. Hautea DM, Molina GC, Balatero CH, Coronado NB, Perez EB, Alvarez MTH, Canama AO, Akuba RH, Quilloy RB, Frankie RB, Caspillo CS (2004) Analysis of induced mutants of Philippine bananas with molecular markers. In: Jain SM, Swennen R (eds) Banana improvement: cellular, molecular biology, and induced mutations. Science Publishers, Inc., Enfield, pp 45-58.
104. Havey MJ, Muehlbauer FJ (1989) Linkages between restriction fragment length, isozyme, and morphological markers in lentil. Theor Appl Genet 77: 395-401.
105. Heinz DJ, Mee GWP (1971) Morphologic, cytogenetic, and enzymatic variation in Saccharum species hybrid clones derived from callus tissue. Am J Bot 58: 257-262.
106. Henry Y, Marcotte JL, de Buyser J (1996) The effect of aneuploidy on karyotype abnormalities in wheat plants regenerated from short- and long-term somatic embryogenesis. Plant Sci 114: 101-109.
107. Hirochika H (1993) Activation of tobacco transposons during tissue culture. EMBO J 12: 2521-2528.
108. Hornero J, Martínez I, Celestino C, Gallego FJ, Torres V, Toribio M (2001) Early checking of genetic stability of cork oak somatic embryos by AFLP analysis. Int J Plant Sci 162: 827-833.
109. Hossain AM, Konisho K, Minami M, Nemoto K (2003) Somaclonal variation of regenerated plants in chili pepper (Capsicum annuum L.). Euphytica 130: 233-239.
110. Hsu T-W, Tsai W-C, Wang D-P, Lin S, Hsiao Y-Y, Chen W-H, Chen H-H (2008) Differential gene expression analysis by cDNA-AFLP between flower buds of Phalaenopsis Hsiang Fei cv. H. F. and its somaclonal variant. Plant Sci 175: 415-422.
111. Hunter RL, Merkert CL (1957) Histochemical demonstration of enzymes separated by zone electrophoresis in starch gels. Science 125: 1294-1295.
112. Israeli Y, Reuveni O, Lahav E (1991) Qualitative aspects of somaclonal variations in banana propagated by in vitro techniques. Sci Hortic 48: 71-88.
113. Israeli Y, Lahav E, Reuveni O (1995) In vitro culture of bananas. In: Gowen S (ed) Bananas and plantians. Chapman and Hall, London, pp 147-178.
114. Israeli Y, Ben-Bassat D, Reuveni O (1996) Selection of stable banana clones which do not produce dwarf somaclonal variants during in vitro culture. Sci Hortic 67: 197-205.
115. Jain SM (1993) Somaclonal variation in Begonia × elatior and Saintpaulia ionantha L. Sci Hortic 54: 221-231.
116. Jain SM (1997) Micropropagation of selected somaclones of Begonia and Saintpaulia. J Biosci 22: 582-592.
117. Jain SM (2001) Tissue culture-derived variation in crop improvement. Euphytica 118: 153-166.
118. Jain SM (2006) Mutation-assisted breeding for improving ornamental plants. Acta Hortic 714: 85-98.
119. Jaligot E, Rival A, Beulé T, Dussert S, Verdeil JL (2000) Somaclonal variation in oil palm (Elaeis guineensis Jacq.): the DNA methylation hypothesis. Plant Cell Rep 19: 684-690.
120. Jaligot E, Beulé T, Rival A (2002) Methylation-sensitive RFLPs: characterisation of two oil palm markers showing somaclonal variation-associated polymorphism. Theor Appl Genet 104: 1263-1269.
121. James AC, Peraza-Echeverria S, Herrera-Valencia VA, Martinez O (2004) Application of the amplified fragment length polymorphism (AFLP) and the methylation-sensitive amplification polymorphism (MSAP) techniques for the detection of DNA polymorphism and changes in DNA methylation in micropropagated bananas. In: Jain SM, Swennen R (eds) Banana improvement: cellular, molecular biology, and induced mutations. Science Publishers, Inc., Enfield, pp 287-306.
122. Jarret RL (1986) Biochemical/genetic markers and their uses in the genus Musa. In: Persley GJ, Langhe EA (eds) Banana and plantian strategies, Cairns, Australia. Australia Centre for International Agricultural Research (ACIAR) proceeding no. 21, Canberra.
123. Jarret RL, Gawel N (1995) Molecular markers, genetic diversity and systematics in Musa. In: Gowen S (ed) Bananas and plantians. Chapman and Hall, London, pp 66-83.
124. Jarret RL, Litz RE (1986) Isozymes as genetic markers in bananas and plantains. Euphytica 35: 539-549.
125. Jin S, Mushke R, Zhu H, Tu L, Lin Z, Zhang Y, Zhang X (2008) Detection of somaclonal variation of cotton (Gossypium hirsutum) using cytogenetics, flow cytometry and molecular markers. Plant Cell Rep 27: 1303-1316.
126. Jones CJ, Edwards KJ, Castaglione S, Winfield MO, Sala F, van de Wiel C, Bredemeijer G, Vosman B, Matthes M, Daly A, Brettschneider R, Bettini P, Buiatti M, Maestri E, Malcevschi A, Marmiroli N, Aert R, Volckaert G, Rueda J, Linacero R, Vazquez A, Karp A (1997) Reproducibility testing of RAPD, AFLP and SSR markers in plants by a network of European laboratories. Mol Breeding 3: 381-390.
127. Joshi RK, Rao GJN (2009) Somaclonal variation in submergence tolerant rice cultivars and induced diversity evaluation by PCR markers. Int J Genet Mol Biol 1: 80-88.
128. Kaeppler S, Phillips R (1993) DNA methylation and tissue culture-induced variation in plants. In Vitro Cell Dev Biol Plant 29: 125-130.
129. Kaeppler SM, Kaeppler HF, Rhee Y (2000) Epigenetic aspects of somaclonal variation in plants. Plant Mol Biol 43: 179-188.
130. Kanwar K, Bindiya K (2003) Random amplified polymorphic DNA (RAPDs) markers for genetic analysis in micropropagated plants of Robinia pseudoacacia L. Euphytica 132: 41-47.
131. Kaplan DR (1992) The relationship of cells to organisms in plants: problem and implications of an organismal perspective. Int J Plant Sci 153: S28-S37.
132. Karp A (1994) Origins, causes and uses of variation in plant tissue cultures. In: Vasil IK, Thorpe TA (eds) Plant cell and tissue culture. Kluwer Academic Publishers, Dordrecht, pp 139-152.
133. Karp A (1995) Somaclonal variation as a tool for crop improvement. Euphytica 85: 295-302.
134. Karp A, Seberg OLE, Buiatti M (1996) Molecular techniques in the assessment of botanical diversity. Ann Bot 78: 143-149.
135. Kawiak A, Łojkowska E (2004) Application of RAPD in the determination of genetic fidelity in micropropagated Drosera plantlets. In Vitro Cell Dev Biol Plant 40: 592-595.
136. Khlestkina E, Röder M, Pshenichnikova T, Börner A (2010) Functional diversity at the Rc (red coleoptile) gene in bread wheat. Mol Breeding 25: 125-132.
137. Kole P, Chawla H (1993) Variation of Helminthosporium resistance and biochemical and cytological characteristics in somaclonal generations of barley. Biol Plant 35: 81-86.
138. Krikorian AD, Irizarry H, Cronauer-Mitra SS, Rivera E (1993) Clonal fidelity and variation in plantain (Musa AAB) regenerated from vegetative stem and floral axis tips in vitro. Ann Bot 71: 519-535.
139. Krikorian AD, Irizarry H, Goenaga R, Scott ME, Lockhart BEL (1999) Stability in plant and bunch traits of a 'French-type' dwarf plantain micropropagated from the floral axis tip and five lateral corm tips of a single mother plant: good news on the tissue culture and bad news on banana streak virus. Sci Hortic 81: 159-177.
140. Kumar M, Barker R, Reed B (1999) Morphological and molecular analysis of genetic stability in micropropagated Fragaria × ananassa cv. pocahontas. In Vitro Cell Dev Biol Plant 35: 254-258.
141. Kunitake H, Koreeda K, Mii M (1995) Morphological and cytological characteristics of protoplast-derived plants of statice (Limonium perezii Hubbard). Sci Hortic 60: 305-312.
142. Kuznetsova OI, Ash OA, Hartina GA, Gostimskij SA (2005) RAPD and ISSR analyses of regenerated pea Pisum sativum L. plants. Russ J Genet 41: 60-65.
143. Ładyżyński M, Burza W, Malepszy S (2002) Relationship between somaclonal variation and type of culture in cucumber. Euphytica 125: 349-356.
144. Lamseejan S, Jompuk P, Wongpiyasatid A, Deeseepan S, Kwanthammachart P (2000) Gamma-rays induced morphological changes in chrysanthemum (Chrysanthemum morifolium). Kasetsart J (Nat Sci) 34: 417-422.
145. Landry BS, Kesseli R, Leung H, Michelmore RW (1987) Comparison of restriction endonucleases and sources of probes for their efficiency in detecting restriction fragment length polymorphisms in lettuce (Lactuca sativa L.). Theor Appl Genet 74: 646-653.
146. Larkin PJ (1998) Introduction. In: Jain SM, Brar DS, Ahloowalia BS (eds) Somaclonal variation and induced mutations in crop improvement. Kluwer Academic Publishers, Dordrecht, pp 3-13.
147. Larkin P, Scowcroft W (1981) Somaclonal variation-a novel source of variability from cell cultures for plant improvement. Theor Appl Genet 60: 197-214.
148. Larkin PJ, Ryan SA, Brettell RIS, Scowcroft WR (1984) Heritable somaclonal variation in wheat. Theor Appl Genet 67: 443-455.
149. Lee M, Phillips RL (1988) The Chromosomal basis of somaclonal variation. Annu Rev Plant Physiol Plant Mol Biol 39: 413-437.
150. Lee S-H, Shon Y, Kim C, Chun H, Cheong Y, Kim Z, Choe Z, Choi Y, Cho M (1999) Variations in the morphology of rice plants regenerated from protoplasts using different culture procedures. Plant Cell Tissue Organ Cult 57: 179-187.
151. Lee YI, Lee IS, Lim YP (2002) Variation in sweetpotato regenerates from gamma-ray irridiated embryogenic callus. J Plant Biotechnol 4: 163-170.
152. Letham D, Gollnow B (1985) Regulators of cell division in plant tissues. XXX. Cytokinin metabolism in relation to radish cotyledon expansion and senescence. J Plant Growth Regul 4: 129-145.
153. Levinson G, Gutman GA (1987) Slipped-strand mispairing: a major mechanism for DNA sequence evolution. Mol Biol Evol 4: 203-221.
154. Li Y-C, Korol AB, Fahima T, Beiles A, Nevo E (2002) Microsatellites: genomic distribution, putative functions and mutational mechanisms: a review. Mol Ecol 11: 2453-2465.
155. Li X, Yu X, Wang N, Feng Q, Dong Z, Liu L, Shen J, Liu B (2007) Genetic and epigenetic instabilities induced by tissue culture in wild barley (Hordeum brevisubulatum (Trin.) Link). Plant Cell Tissue Organ Cult 90: 153-168.
156. Litt M, Luty JA (1989) A hypervariable microsatellite revealed by in vitro amplification of a dinucleotide repeat within the cardiac muscle actin gene. Am J Hum Genet 44: 397-401.
157. Long SP, Humphries S, Falkowski PG (1994) Photoinhibition of photosynthesis in nature. Annu Rev Plant Physiol Plant Mol Biol 45: 633-662.
158. LoSchiavo F, Pitto L, Giuliano G, Torti G, Nuti-Ronchi V, Marazziti D, Vergara R, Orselli S, Terzi M (1989) DNA methylation of embryogenic carrot cell cultures and its variations as caused by mutation, differentiation, hormones and hypomethylating drugs. Theor Appl Genet 77: 325-331.
159. Loureiro J, Pinto G, Lopes T, Doležel J, Santos C (2005) Assessment of ploidy stability of the somatic embryogenesis process in Quercus suber L. using flow cytometry. Planta 221: 815-822.
160. Loureiro J, Capelo A, Brito G, Rodriguez E, Silva S, Pinto G, Santos C (2007) Micropropagation of Juniperus phoenicea from adult plant explants and analysis of ploidy stability using flow cytometry. Biol Plant 51: 7-14.
161. Maddock SE, Semple JT (1986) Field assessment of somaclonal variation in wheat. J Exp Bot 37: 1065-1078.
162. Mandal A, Maiti A, Chowdhury B, Elanchezhian R (2001) Isoenzyme markers in varietal identification of banana. In Vitro Cell Dev Biol Plant 37: 599-604.
163. Martin K, Pachathundikandi S, Zhang C, Slater A, Madassery J (2006) RAPD analysis of a variant of banana (Musa sp.) cv. grande naine and its propagation via shoot tip culture. In Vitro Cell Dev Biol Plant 42: 188-192.
164. Marum L, Rocheta M, Maroco J, Oliveira M, Miguel C (2009) Analysis of genetic stability at SSR loci during somatic embryogenesis in maritime pine (Pinus pinaster). Plant Cell Rep 28: 673-682.
165. Matheka JM, Magiri E, Rasha AO, Machuka J (2008) In vitro selection and characterization of drought tolerant somaclones of tropical maize (Zea mays L.). Biotechnology 7: 641-650.
166. McClintock B (1950) The origin and behavior of mutable loci in maize. Proc Natl Acad Sci USA 36: 344-355.
167. McPheeters K, Skirvin RM (1983) Histogenic layer manipulation in chimeral 'Thornless Evergreen' trailing blackberry. Euphytica 32: 351-360.
168. Mehta YR, Angra DC (2000) Somaclonal variation for disease resistance in wheat and production of dihaploids through wheat × maize hybrids. Genet Mol Biol 23: 617-622.
169. Miñano HS, González-Benito ME, Martín C (2009) Molecular characterization and analysis of somaclonal variation in chrysanthemum cultivars using RAPD markers. Sci Hortic 122: 238-243.
170. Modgil M, Mahajan K, Chakrabarti SK, Sharma DR, Sobti RC (2005) Molecular analysis of genetic stability in micropropagated apple rootstock MM106. Sci Hortic 104: 151-160.
171. Mohanty S, Panda M, Subudhi E, Nayak S (2008) Plant regeneration from callus culture of Curcuma aromatica and in vitro detection of somaclonal variation through cytophotometric analysis. Biol Plant 52: 783-786.
172. Mohmand AS, Nabors MW (1990) Somaclonal variant plants of wheat derived from mature embryo explants of three genotypes. Plant Cell Rep 8: 558-560.
173. Mondal TK, Chand PK (2002) Detection of genetic variation among micropropagated tea [Camellia sinensis (L.) O. Kuntze] by RAPD analysis. In Vitro Cell Dev Biol Plant 38: 296-299.
174. Mujib A (2005) Colchicine induced morphological variants in pineapple. Plant Tissue Cult Biotechnol 15: 127-133.
175. Mujib A, Banerjee S, Dev Ghosh P (2007) Callus induction, somatic embryogenesis and chromosomal instability in tissue culture-raised hippeastrum (Hippeastrum hybridum cv. United Nations). Propag Ornam Plants 7: 169-174.
176. Mulcahy DL, Cresti M, Sansavini S, Douglas GC, Linskens HF, Mulcahy GB, Vignani R, Pancaldi M (1993) The use of random amplified polymorphic DNAs to fingerprint apple genotypes. Sci Hortic 54: 89-96.
177. Müller E, Brown PTH, Hartke S, Lörz H (1990) DNA variation in tissue-culture-derived rice plants. Theor Appl Genet 80: 673-679.
178. Munthali MT, Newbury HJ, Ford-Lloyd BV (1996) The detection of somaclonal variants of beet using RAPD. Plant Cell Rep 15: 474-478.
179. Nakano M, Nomizu T, Mizunashi K, Suzuki M, Mori S, Kuwayama S, Hayashi M, Umehara H, Oka E, Kobayashi H, Asano M, Sugawara S, Takagi H, Saito H, Nakata M, Godo T, Hara Y, Amano J (2006) Somaclonal variation in Tricyrtis hirta plants regenerated from 1-year-old embryogenic callus cultures. Sci Hortic 110: 366-371.
180. Nayak S, Debata BK, Srivastava VK, Sangwan NS (2003) Evaluation of agronomically useful somaclonal variants in Jamrosa (a hybrid Cymbopogon) and detection of genetic changes through RAPD. Plant Sci 164: 1029-1035.
181. Nehra NS, Kartha KK, Stushnott C, Giles KL (1992) The influence of plant growth regulator concentrations and callus age on somaclonal variation in callus culture regenerants of strawberry. Plant Cell Tissue Organ Cult 29: 257-268.
182. Ngezahayo F, Dong Y, Liu B (2007) Somaclonal variation at the nucleotide sequence level in rice (Oryza sativa L.) as revealed by RAPD and ISSR markers, and by pairwise sequence analysis. J Appl Genet 48: 329-336.
183. Niizeki M, Ishikawa R, Saito K (1990) Variation in a single protoplast- and seed-derived population of Lotus corniculatus L. Theor Appl Genet 80: 732-736.
184. Noro Y, Takano-Shimizu T, Syono K, Kishima Y, Sano Y (2007) Genetic variations in rice in vitro cultures at the EPSPs-RPS20 region. Theor Appl Genet 114: 705-711.
185. Nwauzoma AB, Tenkouano A, Crouch JH, Pillay M, Vuylsteke D, Daniel Kalio LA (2002) Yield and disease resistance of plantain (Musa spp., AAB group) somaclones in Nigeria. Euphytica 123: 323-331.
186. Olmos S, Lavia G, Di Renzo M, Mroginski L, Echenique V (2002) Genetic analysis of variation in micropropagated plants of Melia azedarach L. In Vitro Cell Dev Biol Plant 38: 617-622.
187. Oono K (1985) Putative homozygous mutations in regenerated plants of rice. Mol Gen Genet 198: 377-384.
188. Orbović V, Ćalović M, Viloria Z, Nielsen B, Gmitter F, Castle W, Grosser J (2008) Analysis of genetic variability in various tissue culture-derived lemon plant populations using RAPD and flow cytometry. Euphytica 161: 329-335.
189. Palombi MA, Lombardo B, Caboni E (2007) In vitro regeneration of wild pear (Pyrus pyraster Burgsd) clones tolerant to Fe-chlorosis and somaclonal variation analysis by RAPD markers. Plant Cell Rep 26: 489-496.
190. Pardha Saradhi P, Alia (1995) Production and selection of somaclonal variants of Leucaena leucocephala with high carbon dioxide assimilating potential. Energ Convers Manage 36: 759-762.
191. Peraza-Echeverria S, Herrera-Valencia VA, Kay A-J (2001) Detection of DNA methylation changes in micropropagated banana plants using methylation-sensitive amplification polymorphism (MSAP). Plant Sci 161: 359-367.
192. Peschke VM, Phillips RL (1992) Genetic implications of somaclonal variation in plants. Adv Genet 30: 41-75.
193. Peschke VM, Phillips RL, Gengenbach BG (1987) Discovery of transposable element activity among progeny of tissue culture-derived maize plants. Science 238: 804-807.
194. Petolino JF, Roberts JL, Jayakumar P (2003) Plant cell culture: a critical tool for agricultural biotechnology. In: Vinci VA, Parekh SR (eds) Handbook of industrial cell culture: mammalian, microbial and plant cells. Humana Press, New Jersey, pp 243-258.
195. Peyvandi M, Noormohammadi Z, Banihashemi O, Farahani F, Majd A, Hosseini-Mazinani M, Sheidai M (2009) Molecular analysis of genetic stability in long-term micropropagated shoots of Olea europaea L. (cv. Dezful). Asian J Plant Sci 8: 146-152.
196. Pfosser M, Heberle-Bors E, Amon A, Lelley T (1995) Evaluation of sensitivity of flow cytometry in detecting aneuploidy in wheat using disomic and ditelosomic wheat-rye addition lines. Cytometry 21: 387-393.
197. 1 dwarfism and shoot elongation in higher plants. Biol Plant 27: 172-179.
198. Pierik RLM (1987) In vitro culture of higher plants. Kluwer Academic Publishers, Dordrecht.
199. Pietsch G, Anderson N (2007) Epigenetic variation in tissue cultured Gaura lindheimeri. Plant Cell Tissue Organ Cult 89: 91-103.
200. Piola F, Rohr R, Heizmann P (1999) Rapid detection of genetic variation within and among in vitro propagated cedar (Cedrus libani Loudon) clones. Plant Sci 141: 159-163.
201. Plader W, Malepszy S, Burza W, Rusinowski Z (1998) The relationship between the regeneration system and genetic variability in the cucumber (Cucumis sativus L.). Euphytica 103: 9-15.
202. Podwyszyńska M (2005) Somaclonal variation in micropropagated tulips based on phenotype observation. J Fruit Ornam Plant Res 13: 109-122.
203. Polanco C, Ruiz ML (2002) AFLP analysis of somaclonal variation in Arabidopsis thaliana regenerated plants. Plant Sci 162: 817-824.
204. Pontaroli AC, Camadro EL (2005) Somaclonal variation in Asparagus officinalis plants regenerated by organogenesis from long-term callus cultures. Genet Mol Biol 28: 423-430.
205. Popescu AN, Isac VS, Coman MS MSR (1997) Somaclonal variation in plants regenerated by organogenesis from callus cultures of strawberry (Fragaria × ananassa). Acta Hortic 439: 89-96.
206. Prado M, Rodriguez E, Rey L, González M, Santos C, Rey M (2010) Detection of somaclonal variants in somatic embryogenesis-regenerated plants of Vitis vinifera by flow cytometry and microsatellite markers. Plant Cell Tissue Organ Cult 103: 49-59.
207. Predieri S (2001) Mutation induction and tissue culture in improving fruits. Plant Cell Tissue Organ Cult 64: 185-210.
208. Radhakrishnan R, Ranjitha Kumari B (2008) Morphological and agronomic evaluation of tissue culture derived Indian soybean plants. Acta Agric Slov 91: 391-396.
209. Rady M (2006) In vitro culture of Gypsophila paniculata L. and random amplified polymorphic DNA analysis of the propagated plants. Biol Plant 50: 507-513.
210. Rahman MH, Rajora OP (2001) Microsatellite DNA somaclonal variation in micropropagated trembling aspen (Populus tremuloides). Plant Cell Rep 20: 531-536.
211. Raimondi JP, Masuelli RW, Camadro EL (2001) Assessment of somaclonal variation in asparagus by RAPD fingerprinting and cytogenetic analyses. Sci Hortic 90: 19-29.
212. Rani V, Raina S (2000) Genetic fidelity of organized meristem-derived micropropagated plants: a critical reappraisal. In Vitro Cell Dev Biol Plant 36: 319-330.
213. Rani V, Parida A, Raina S (1995) Random amplified polymorphic DNA (RAPD) markers for genetic analysis in micropropagated plants of Populus deltoides Marsh. Plant Cell Rep 14: 459-462.
214. Rao IM, Roca WM, Ayarza MA, Tabares E, Garcia R (1992) Somaclonal variation in plant adaptation to acid soil in the tropical forage legume Stylosanthes guianensis. Plant Soil 146: 21-30.
215. Rasheed S, Tahira F, Khurram B, Tayyab H, Shiekh R (2003) Agronomical and physiochemical characterization of somaclonal variants in Indica basmati rice. Pak J Biol Sci 6: 844-848.
216. Ray T, Dutta I, Saha P, Das S, Roy SC (2006) Genetic stability of three economically important micropropagated banana (Musa spp.) cultivars of lower Indo-Gangetic plains, as assessed by RAPD and ISSR markers. Plant Cell Tissue Organ Cult 85: 11-21.
217. Reuveni O, Israeli Y (1990) Measures to reduce somaclonal variation in in vitro propagated bananas. Acta Hortic 275: 307-313.
218. Reuveni O, Israeli Y, Golubowicz S (1993) Factors influencing the occurrence of somaclonal variations in micropropagated bananas. Acta Hortic 336: 357-364.
219. Rietveld RC, Hasegawa PM, Bressan RA (1991) Somaclonal variation in tuber disc-derived populations of potato. Theor Appl Genet 82: 430-440.
220. Rietveld RC, Bressan RA, Hasegawa PM (1993) Somaclonal variation in tuber disc-derived populations of potato. II. Differential effect of genotype. Theor Appl Genet 87: 305-313.
221. Rivera FN (1983) Protein and isoenzyme banding patterns among Philippine cooking bananas and their wild parents (Musa species). Paradisiaca 6: 7-12.
222. Rodrigues PHV (2008) Somaclonal variation in micropropagated Heliconia bihai cv. Lobster Claw I plantlets (Heliconiaceae). Sci Agric 65: 681-684.
223. Rodrigues PHV, Tulmann Neto A, Cassieri Neto P, Mendes BMJ (1998) Influence of the number of subcultures on somoclonal variation in micropropagated Nanico (Musa spp., AAA group). Acta Hortic 490: 469-473.
224. Rodríguez López C, Bravo H, Wetten A, Wilkinson M (2010) Detection of somaclonal variation during cocoa somatic embryogenesis characterised using cleaved amplified polymorphic sequence and the new freeware Artbio. Mol Breeding 25: 501-516.
225. Rodríguez NV, Kowalski B, Rodríguez LG, Caraballoso IB, Suárez MA, Pérez PO, Quintana CR, González N, Ramos RQ (2007) In vitro and ex vitro selection of potato plantlets for resistance to early blight. J Phytopathol 155: 582-586.
226. Roels S, Escalona M, Cejas I, Noceda C, Rodriguez R, Canal MJ, Sandoval J, Debergh P (2005) Optimization of plantain (Musa AAB) micropropagation by temporary immersion system. Plant Cell Tissue Organ Cult 82: 57-66.
227. Roux NS (2004) Mutation induction in Musa-review. In: Jain SM, Swennen R (eds) Banana improvement: cellular, molecular biology, and induced mutation. Science Publishers, Inc., Enfield, pp 23-32.
228. Roy B, Mandal A (2005) Towards development of Al-toxicity tolerant lines in indica rice by exploiting somaclonal variation. Euphytica 145: 221-227.
229. Ryan SA, Larkin PJ, Ellison FW (1987) Somaclonal variation in some agronomic and quality characters in wheat. Theor Appl Genet 74: 77-82.
230. Sahijram L, Soneji J, Bollamma K (2003) Analyzing somaclonal variation in micropropagated bananas (Musa spp.). In Vitro Cell Dev Biol Plant 39: 551-556.
231. Sanchez-Teyer LF, Quiroz-Figueroa F, Loyola-Vargas V, Infante D (2003) Culture-induced variation in plants of Coffea arabica cv. Caturra rojo, regenerated by direct and indirect somatic embryogenesis. Mol Biotechnol 23: 107-115.
232. Sandoval J, Kerbellec F, Côte F, Doumas P (1995) Distribution of endogenous gibberellins in dwarf and giant off-types banana (Musa AAA cv. 'Grand nain') plants from in vitro propagation. Plant Growth Regul 17: 219-224.
233. Saxena G, Banerjee S, Rahman L, Mallavarapu GR, Sharma S, Kumar S (2000) An efficient in vitro procedure for micropropagation and generation of somaclones of rose scented Pelargonium. Plant Sci 155: 133-140.
234. Schlötterer C, Tautz D (1992) Slippage synthesis of simple sequence DNA. Nucleic Acids Res 20: 211-215.
235. Scowcroft WR (1984) Genetic variability in tissue culture: impact on germplasm conservation and utilization. International board for plant genetic resources (IBPGR) technical report AGPGIBPGR/84/152, Rome.
236. Senevirathna AMWK, Stirling CM, Rodrigo VHL (2008) Acclimation of photosynthesis and growth of banana (Musa sp.) to natural shade in the humid tropics. Exp Agric 44: 301-312.
237. Sengar A, Thind K, Kumar B, Pallavi M, Gosal S (2009) In vitro selection at cellular level for red rot resistance in sugarcane (Saccharum sp.). Plant Growth Regul 58: 201-209.
238. Shah SH, Wainwright SJ, Merret MJ (2003) Regeneration and somaclonal variation in Medicago sativa and Medicago media. Pak J Biol Sci 6: 816-820.
239. Sharma S, Bryan G, Winfield M, Millam S (2007) Stability of potato (Solanum tuberosum L.) plants regenerated via somatic embryos, axillary bud proliferated shoots, microtubers and true potato seeds: a comparative phenotypic, cytogenetic and molecular assessment. Planta 226: 1449-1458.
240. Shchukin A, Ben-Bassat D, Israeli Y (1997) Plant regenration via somatic embryogenesis in Grand Naine banana and its effect on somoclonal variation. Acta Hortic 447: 317-318.
241. Sheidai M, Aminpoor H, Noormohammadi Z, Farahani F (2008) RAPD analysis of somaclonal variation in banana (Musa acuminate L.) cultivar Valery. Acta Biol Szeged 52: 307-311.
242. Shen X, Chen J, Kane M, Henny R (2007) Assessment of somaclonal variation in Dieffenbachia plants regenerated through indirect shoot organogenesis. Plant Cell Tissue Organ Cult 91: 21-27.
243. Shepherd K, Dos Santos JA (1996) Mitotic instability in banana varieties. I. Plants from callus and shoot tip cultures. Fruits 51: 5-11.
244. Shushu DD, Comar JM, Abegaz BM (2009) Somaclonal variation in in vitro regenerated Ledebouria graminifolia (hyacinthaceae), an indigenous bulb in Botswana and its potential exploitation as an ornamental plant. J Biol Sci 9: 152-158.
245. Simmons MP, Zhang L-B, Webb CT, Müller K (2007) A penalty of using anonymous dominant markers (AFLPs, ISSRs, and RAPDs) for phylogenetic inference. Mol Phylogenet Evol 42: 528-542.
246. Singh A, Lai M, Singh M, Lai K, Singh S (2000) Variations for red rot resistance in somaclones of sugarcane. Sugar Tech 2: 56-58.
247. Singh M, Chaudhary K, Singal H, Magill C, Boora K (2006) Identification and characterization of RAPD and SCAR markers linked to anthracnose resistance gene in sorghum [Sorghum bicolor (L.) Moench]. Euphytica 149: 179-187.
248. Singh G, Sandhu S, Meeta M, Singh K, Gill R, Gosal S (2008a) In vitro induction and characterization of somaclonal variation for red rot and other agronomic traits in sugarcane. Euphytica 160: 35-47.
249. Singh R, Srivastava S, Singh S, Sharma M, Mohopatra T, Singh N (2008b) Identification of new microsatellite DNA markers for sugar and related traits in sugarcane. Sugar Tech 10: 327-333.
250. Siragusa M, Carra A, Salvia L, Puglia A, De Pasquale F, Carimi F (2007) Genetic instability in calamondin (Citrus madurensis Lour.) plants derived from somatic embryogenesis induced by diphenylurea derivatives. Plant Cell Rep 26: 1289-1296.
251. Sivanesan I (2007) Shoot regeneration and somaclonal variation from leaf callus cultures of Plumbago zeylanica Linn. Asian J Plant Sci 6: 83-86.
252. Skirvin RM (1978) Natural and induced variation in tissue culture. Euphytica 27: 241-266.
253. Skirvin RM, Norton M, McPheeters KD (1993) Somaclonal variation: has it proved useful for plant improvement? Acta Hortic 336: 333-340.
254. Skirvin RM, McPheeters KD, Norton M (1994) Sources and frequency of somaclonal variation. HortScience 29: 1232-1237.
255. Skoog F, Miller CO (1957) Chemical regulation of growth and organ formation in plant tissues cultured in vitro. Symp Soc Exp Biol 54: 118-130.
256. Smýkal P, Valledor L, Rodríguez R, Griga M (2007) Assessment of genetic and epigenetic stability in long-term in vitro shoot culture of pea (Pisum sativum L.). Plant Cell Rep 26: 1985-1998.
257. Squirrell J, Hollingsworth PM, Woodhead M, Russell J, Lowe AJ, Gibby M, Powell W (2003) How much effort is required to isolate nuclear microsatellites from plants? Mol Ecol 12: 1339-1348.
258. Srivastava S, Gupta PS, Srivastava BL (2005) Genetic relationship and clustering of some sugarcane genotypes based on esterase, peroxidase and amylase isozyme polymorphism. Cytologia 70: 355-363.
259. Stover RH (1987) Somaclonal variation in Grande Naine and Saba bananas in the nursery and in the field. In: Persley GJ, De Langhe E (eds) ACIAR proceeding no. 21, Canberra.
260. Sultana R, Tahira F, Tayyab H, Khurram B, Shiekh R (2005) RAPD characterization of somaclonal variation in indica basmati rice. Pak J Bot 37: 249-262.
261. Swartz HJ (1991) Post culture behaviour, genetic and epigenetic effects and related problems. In: Debergh PC, Zimmerman RH (eds) Micropropagation: technology and application. Kluwer Academic Publishers, Dodrecht, pp 95-122.
262. Symillides Y, Henry Y, Buyser J (1995) Analysis of Chinese Spring regenerants obtained from short- and long-term wheat somatic embryogenesis. Euphytica 82: 263-268.
263. Tangpong P, Taychasinpitak T, Jompuk C, Jompuk P (2009) Effects of acute and chronic gamma irradiations on in vitro culture of Anubias congensis N. E. Brown. Kasetsart J (Nat Sci) 43: 449-457.
264. Tautz D, Renz M (1984) Simple sequences are ubiquitous repetitive components of eukaryotic genomes. Nucleic Acids Res 12: 4127-4138.
265. Taylor PWJ, Geijskes JR, Ko HL, Fraser TA, Henry RJ, Birch RG (1995) Sensitivity of random amplified polymorphic DNA analysis to detect genetic change in sugarcane during tissue culture. Theor Appl Genet 90: 1169-1173.
266. Thieme R, Griess H (2005) Somaclonal variation in tuber traits of potato. Potato Res 48: 153-165.
267. Thomas J, Raj Kumar R, Mandal AKA (2006) Metabolite profiling and characterization of somaclonal variants in tea (Camellia spp.) for identifying productive and quality accession. Phytochemistry 67: 1136-1142.
268. Thomas JO, Margaret AC, Karl K, Iris E, Rony S, Christopher AC (2007) Genomic changes associated with somaclonal variation in banana (Musa spp.). Physiol Plant 129: 766-774.
269. Todorovska E, Trikonova A, Petrova M, Vitanova Z, Marinova E, Gramatikova M, Valcheva D, Zaprianov S, Mersinkov N, Atanassov A (1997) Agronomic performance and molecular assessment of tissue culture-derived barley lines. Plant Breed 116: 511-517.
270. Tremblay L, Levasseur C, Tremblay FM (1999) Frequency of somaclonal variation in plants of black spruce (Picea mariana, Pinaceae) and white spruce (P. glauca, Pinaceae) derived from somatic embryogenesis and identification of some factors involved in genetic instability. Am J Bot 86: 1373-1381.
271. Unai E, Iselen T, de Garcia E (2004) Comparison of characteristics of bananas (Musa sp.) from the somaclone CIEN BTA-03 and its parental clone Williams. Fruits 59: 257-263.
272. van Belkum A, Scherer S, van Alphen L, Verbrugh H (1998) Short-sequence DNA repeats in prokaryotic genomes. Microbiol Mol Biol Rev 62: 275-293.
273. van den Bulk RW, Löffler HJM, Lindhout WH, Koornneef M (1990) Somaclonal variation in tomato: effect of explant source and a comparison with chemical mutagenesis. Theor Appl Genet 80: 817-825.
274. van der Wurff AWG, Chan YL, van Straalen NM, Schouten J (2000) TE-AFLP: combining rapidity and robustness in DNA fingerprinting. Nucleic Acids Res 28: e105.
275. van Harten AM (1998) Mutation breeding: theory and practical applications. Cambridge University Press, Cambridge.
276. van Harten AM, Bouter H, Broertjes C (1981) In vitro adventitious bud techniques for vegetative propagation and mutation breeding of potato (Solanum tuberosum L.). II. Significance for mutation breeding. Euphytica 30: 1-8.
277. van Sint Jan V, Costa de Macedo C, Kinet J-M, Bouharmont J (1997) Selection of Al-resistant plants from a sensitive rice cultivar, using somaclonal variation, in vitro and hydroponic cultures. Euphytica 97: 303-310.
278. Vendrame WA, Kochert G, Wetzstein HY (1999) AFLP analysis of variation in pecan somatic embryos. Plant Cell Rep 18: 853-857.
279. Venkatachalam L, Sreedhar RV, Bhagyalakshmi N (2007a) Genetic analyses of micropropagated and regenerated plantlets of banana as assessed by RAPD and ISSR markers. In Vitro Cell Dev Biol Plant 43: 267-274.
280. Venkatachalam L, Sreedhar RV, Bhagyalakshmi N (2007b) Molecular analysis of genetic stability in long-term micropropagated shoots of bananas using RAPD and ISSR markers. Electron J Biotechnol 10: 1-8.
281. Venkatachalam L, Sreedhar RV, Bhagyalakshmi N (2008) The use of genetic markers for detecting DNA polymorphism, genotype identification and phylogenetic relationships among banana cultivars. Mol Phylogenet Evol 47: 974-985.
282. Vidal MDC, De García E (2000) Analysis of a Musa spp. somaclonal variant resistant to yellow Sigatoka. Plant Mol Biol Rep 18: 23-31.
283. Vos P, Hogers R, Bleeker M, Reijans M, Tvd Lee, Hornes M, Friters A, Pot J, Paleman J, Kuiper M, Zabeau M (1995) AFLP: a new technique for DNA fingerprinting. Nucleic Acids Res 23: 4407-4414.
284. Vuylsteke D, Swennen R, Wilson GF, De Langhe E (1988) Phenotypic variation among in vitro propagated plantain (Musa sp. cultivar 'AAB'). Sci Hortic 36: 79-88.
285. Vyskot B, Fajkus J, Kuglík P, Koukalová B, Kuhrová V (1991) Genome modifications in protoplast-derived tobacco plants: phenotypic evaluation and RFLP Analysis. Biol Plant 33: 455-460.
286. Wang Y, Wang F, Zhai H, Liu Q (2007) Production of a useful mutant by chronic irradiation in sweetpotato. Sci Hortic 111: 173-178.
287. Wattanasiri C, Walton PD (1993) Effects of growth regulators on callus cell growth, plant regeneration, and somaclonal variation of smooth bromegrass (Bromus inermis Leyss.). Euphytica 69: 77-82.
288. Weising K, Nybom H, Wolff K, Kahl G (2005) DNA fingerprinting in plants: principles, methods, and applications. CRC Press, New York.
289. Welsh J, McClelland M (1990) Fingerprinting genomes using PCR with arbitrary primers. Nucleic Acids Res 18: 7213-7218.
290. Welter L, Göktürk-Baydar N, Akkurt M, Maul E, Eibach R, Töpfer R, Zyprian E (2007) Genetic mapping and localization of quantitative trait loci affecting fungal disease resistance and leaf morphology in grapevine (Vitis vinifera L). Mol Breeding 20: 359-374.
291. Wenz H-M, Robertson JM, Menchen S, Oaks F, Demorest DM, Scheibler D, Rosenblum BB, Wike C, Gilbert DA, Efcavitch JW (1998) High-precision genotyping by denaturing capillary electrophoresis. Genome Res 8: 69-80.
292. Wilhelm E, Hristoforoglu K, Fluch S, Burg K (2005) Detection of microsatellite instability during somatic embryogenesis of oak (Quercus robur L.). Plant Cell Rep 23: 790-795.
293. Williams JGK, Kubelik AR, Livak KJ, Rafalski JA, Tingey SV (1990) DNA polymorphisms amplified by arbitrary primers are useful as genetic markers. Nucleic Acids Res 18: 6531-6535.
294. Xu Q-l, Hu Z, Li C-y, Wang X-y, Wang C-y (2009) Tissue culture of Sinningia speciosa and analysis of the in vitro-generated tricussate whorled phyllotaxis (twp) variant. In Vitro Cell Dev Biol Plant 45: 583-590.
295. Yadav P, Suprasanna P, Gopalrao K, Anant B (2006) Molecular profiling using RAPD technique of salt and drought tolerant regenerants of sugarcane. Sugar Tech 8: 63-68.
296. Yamagishi M, Koba T, Shimada T, Itoh K, Sukekiyo Y, Shimamoto K (1997) Characteristics of genetic variation in the progenies of protoplast-derived plants of rice, Oryza sativa cv Nipponbare. Theor Appl Genet 94: 1-7.
297. Yang H, Tabei Y, Kamada H, Kayano T, Takaiwa F (1999) Detection of somaclonal variation in cultured rice cells using digoxigenin-based random amplified polymorphic DNA. Plant Cell Rep 18: 520-526.
298. Yang W-R, Zhang Q-X, Pan H-T, Sun M (2010) In vitro regeneration of Lilium tsingtauense Gilg. and analysis of genetic variability in micropropagated plants using RAPD and ISSR techniques. Propag Ornam Plants 10: 59-66.
299. Zaffari GR, Peres LEP, Kerbauy GB (1998) Endogenous levels of cytokinins, indoleacetic acid, abscisic acid, and pigments in variegated somaclones of micropropagated banana leaves. J Plant Growth Regul 17: 59-61.
300. Zaid A, Al Kaabi H (2003) Plant-off types in tissue culture-derived date palm (Phoenix dactylifera L.). Emirates J Agric Sci 15: 17-35.
301. Zhang M, Wang H, Dong Z, Qi B, Xu K, Liu B (2010) Tissue culture-induced variation at simple sequence repeats in sorghum (Sorghum bicolor L.) is genotype-dependent and associated with down-regulated expression of a mismatch repair gene, MLH3. Plant Cell Rep 29: 51-59.
302. Zhao Y, Grout BWW, Crisp P (2005) Variations in morphology and disease susceptibility of micropropagated rhubarb (Rheum rhaponticum) PC49, compared to conventional plants. Plant Cell Tissue Organ Cult 82: 357-361.
303. Zheng K-L, Zhou Z-M, Wang G-L, Luo Y-K, Xiong Z-M (1989) Somatic cell culture of rice cultivars with different grain types: somaclonal variation in some grain and quality characters. Plant Cell Tissue Organ Cult 18: 201-208.
304. Zhong Z, Smith HG, Thomas TH (1993) In vitro culture of petioles and intact leaves of sugar beet (Beta vulgaris). Plant Growth Regul 12: 59-66.
305. Zietkiewicz E, Rafalski A, Labuda D (1994) Genome fingerprinting by simple sequence repeat (SSR)-anchored polymerase chain reaction amplification. Genomics 20: 176-183.