Due to mainly human population pressure and activities, global biodiversity is getting reduced and particularly plant biodiversity is becoming at high risk of extinction. Consequently, many efforts have been deployed to develop conservation methods. Because it does not involve cell dedifferentiation of differentiated cells but rather the development and growth of new shoots from preexisting meristems, the axillary bud proliferation approach is the method offering least risk of genetic instability. Indeed, meristems are more resistant to genetic changes than disorganized tissues. The present review explored through the scientific literature the axillary bud proliferation approach and the possible somaclonal variation that could arise from it. Almost genetic stability or low level of genetic variation is often reported. On the contrary, in a few cases studied to date, DNA methylation alterations often appeared in the progenies, showing epigenetic variations in the regenerated plants from axillary bud culture. Fortunately, epigenetic changes are often temporary and plants may revert to the normal phenotype. Thus, in the absence of genetic variations and the existence of reverting epigenetic changes over time, axillary bud culture can be adopted as an alternative nonconventional way of conserving and restoring of plant biodiversity. 1. Introduction Global biodiversity is defined as the variation of all life on earth and the ecological complexes in which it occurs [1]. Biodiversity refers to genetic diversity, species diversity, and ecosystem diversity [2, 3] and includes the forest and agricultural ecosystems and the wild animals [4]. Among the above components, plants represent a vital part of biodiversity and healthy ecosystems. They provide multiple ecosystem services including production of oxygen for the rest of living organisms [5, 6], removal of atmospheric carbon dioxide emissions in the photosynthesis process, creation and stabilization of soil, protection of watersheds, and provision of natural resources including food, fibre, fuel, shelter, and medicine [7]. They also play an important role in the water cycle and constitute habitat for a wide range of other living organisms. Thus, plants are the basis for life on earth and humans are quite dependent on them [8–10] given that they are fundamental structural and nutrient-sequestering components of most ecosystems. Due to dependency on biodiversity, the number of threatened plant species has gradually increased during the last decade, the maximum being observed in 2011 [11]. The key factor in
References
[1]
P. Leadley, H. M. Pereira, R. Alkemade et al., Biodiversity Scenarios: Projections of 21st Century Change in Biodiversity and Associated Ecosystem Services, Convention on Biological Diversity, Montreal, Canada, 2010.
[2]
S. I. Dodson, T. F. H. Allen, S. R. Carpenter et al., Ecology, Oxford University Press, New York, NY, USA, 1998.
[3]
K. J. Gaston and J. I. Spicer, Biodiversity: An Introduction, Blackwell Science, Oxford, UK, 1998.
[4]
World Bank, “Ensuring the Future: The World Bank and Biodiversity, 1998–2004,” Tech. Rep., World Bank, Environment Department, Washington, DC, USA.
[5]
M. A. Huston, Biological Diversity: The Coexistence of Species on Changing Landscapes, Cambridge University Press, Cambridge, Mass, USA, 1994.
[6]
R. B. Primack and R. T. Corlett, Tropical Rain Forests: An Ecological and Biogeographical Comparison, Blackwell Publishing, Oxford, UK, 2005.
[7]
E. O. Wilson, The Diversity of Life, Penguin, London, UK, 1992.
[8]
G. C. Daily, “Challenges in valuation,” in Nature's Services: Societal Dependence on Natural ecosystems, G. C. Daily, Ed., pp. 365–374, Island Press, Washington, DC, USA, 1997.
[9]
A. Balmford, A. Bruner, P. Cooper et al., “Ecology: economic reasons for conserving wild nature,” Science, vol. 297, no. 5583, pp. 950–953, 2002.
[10]
A. Hamilton and P. Hamilton, Plant Conservation: An Ecosystem Approach, Earthscan, London, UK, 2006.
[11]
IUCN Red List version 2011.2, https://www.bolgermany.de/dateien/How_many_species_IUCN_Data.pdf.
[12]
G. L. Kirkland Jr. and R. S. Ostfeld, “Factors influencing variation among states in the number of federally listed mammals in the United States,” Journal of Mammalogy, vol. 80, no. 3, pp. 711–719, 1999.
[13]
K. Thompson and A. Jones, “Human population density and prediction of local plant extinction in Britain,” Conservation Biology, vol. 13, no. 1, pp. 185–189, 1999.
[14]
M. Cardillo, A. Purvis, W. Sechrest, J. L. Gittleman, J. Bielby, and G. M. Mace, “Human population density and extinction risk in the world's carnivores,” PLoS Biology, vol. 2, no. 7, 2004.
[15]
E. C. Ellis, E. C. Antill, and H. Kreft, “All is not loss: plant biodiversity in the anthropocene,” PLoS ONE, vol. 7, no. 1, Article ID e30535, 2012.
[16]
United Nations Population Division, World Population Prospects, 2008.
[17]
United Nations Population Division, Briefing Packet, 1998 Revision of World Population Prospects, and World Population Prospects, the 2006 Revision.
[18]
A. K. Diraiappah and S. Naeem, Millennium Ecosystem Assessment: Ecosystems and Human Well-Being: Biodiversity Synthesis, World Resources Institute, Washington, DC, USA, 2005.
[19]
S. H. M. Butchart, M. Walpole, B. Collen et al., “Global biodiversity: indicators of recent declines,” Science, vol. 328, no. 5982, pp. 1164–1168, 2010.
[20]
F. S. Chapin III, E. S. Zavaleta, V. T. Eviner et al., “Consequences of changing biodiversity,” Nature, vol. 405, no. 6783, pp. 234–242, 2000.
[21]
M. Loreau, S. Naeem, P. Inchausti et al., “Ecology: biodiversity and ecosystem functioning: current knowledge and future challenges,” Science, vol. 294, no. 5543, pp. 804–808, 2001.
[22]
B. J. Cardinale, D. S. Srivastava, J. E. Duffy et al., “Effects of biodiversity on the functioning of trophic groups and ecosystems,” Nature, vol. 443, no. 7114, pp. 989–992, 2006.
[23]
D. U. Hooper, F. S. Chapin III, J. J. Ewel et al., “Effects of biodiversity on ecosystem functioning: a consensus of current knowledge,” Ecological Monographs, vol. 75, no. 1, pp. 3–35, 2005.
[24]
P. Balvanera, A. B. Pfisterer, N. Buchmann et al., “Quantifying the evidence for biodiversity effects on ecosystem functioning and services,” Ecology Letters, vol. 9, no. 10, pp. 1146–1156, 2006.
[25]
D. S. Srivastava and M. Vellend, “Biodiversity-ecosystem function research: is it relevant to conservation?” Annual Review of Ecology, Evolution, and Systematics, vol. 36, pp. 267–294, 2005.
[26]
H. Hillebrand and B. Matthiessen, “Biodiversity in a complex world: consolidation and progress in functional biodiversity research,” Ecology Letters, vol. 12, no. 12, pp. 1405–1419, 2009.
[27]
J. Emmett Duffy, “Why biodiversity is important to the functioning of real-world ecosystems,” Frontiers in Ecology and the Environment, vol. 7, no. 8, pp. 437–444, 2009.
[28]
J. T. Kerr and D. J. Currie, “Effects of human activity on global extinction risk,” Conservation Biology, vol. 9, no. 6, pp. 1528–1538, 1995.
[29]
D. J. Forester and G. E. Machlis, “Modeling human factors that affect the loss of biodiversity,” Conservation Biology, vol. 10, no. 4, pp. 1253–1263, 1996.
[30]
K. Thompson and A. Jones, “Human population density and prediction of local plant extinction in Britain,” Conservation Biology, vol. 13, no. 1, pp. 185–189, 1999.
[31]
R. P. Cincotta, J. Wisnewski, and R. Engelman, “Human population in the biodiversity hotspots,” Nature, vol. 404, no. 6781, pp. 990–992, 2000.
[32]
M. L. McKinney, “Role of human population size in raising bird and mammal threat among nations,” Animal Conservation, vol. 4, no. 1, pp. 45–57, 2001.
[33]
A. H. Harcourt and S. A. Parks, “Threatened primates experience high human densities: adding an index of threat to the IUCN Red List criteria,” Biological Conservation, vol. 109, no. 1, pp. 137–149, 2002.
[34]
A. H. Harcourt, S. A. Parks, and R. Woodroffe, “Human density as an influence on species/area relationships: double jeopardy for small African reserves?” Biodiversity and Conservation, vol. 10, no. 6, pp. 1011–1026, 2001.
[35]
G. Ceballos and P. R. Ehrlich, “Mammal population losses and the extinction crisis,” Science, vol. 296, no. 5569, pp. 904–907, 2002.
[36]
M. Parkes, “Personal commentaries on ‘ecosystems and human well-being: Health synthesis—a report of the millennium ecosystem assessment’,” EcoHealth, vol. 3, no. 3, pp. 136–140, 2006.
[37]
Y. A. Izrael, S. M. Semenov, O. A. Anisimov et al., “The fourth assessment report of the intergovernmental panel on climate change: working group II contribution,” Russian Meteorology and Hydrology, vol. 32, no. 9, pp. 551–556, 2007.
[38]
O. E. Sala, F. S. Chapin III, J. J. Armesto et al., “Global biodiversity scenarios for the year 2100,” Science, vol. 287, no. 5459, pp. 1770–1774, 2000.
[39]
R. E. Green, S. J. Cornell, J. P. W. Scharlemann, and A. Balmford, “Farming and the fate of wild nature,” Science, vol. 307, no. 5709, pp. 550–555, 2005.
[40]
J. A. Foley, N. Ramankutty, K. A. Brauman et al., “Solutions for a cultivated planet,” Nature, vol. 478, no. 7369, pp. 337–342, 2011.
[41]
http://www.fao.org/fishery/topic/3541/en.
[42]
B. Czech, P. R. Krausman, and P. K. Devers, “Economic associations among causes of species endangerment in the United States,” BioScience, vol. 50, no. 7, pp. 593–601, 2000.
[43]
M. L. McKinney, “Urbanization, biodiversity, and conservation,” BioScience, vol. 52, no. 10, pp. 883–890, 2002.
[44]
M. L. McKinney, “Measuring floristic homogenization by non-native plants in North America,” Global Ecology and Biogeography, vol. 13, no. 1, pp. 47–53, 2004.
[45]
P. M. Vitousek, H. A. Mooney, J. Lubchenco, and J. M. Melillo, “Human domination of Earth's ecosystems,” Science, vol. 277, no. 5325, pp. 494–499, 1997.
[46]
R. I. Mcdonald, P. Kareiva, and R. T. T. Forman, “The implications of current and future urbanization for global protected areas and biodiversity conservation,” Biological Conservation, vol. 141, no. 6, pp. 1695–1703, 2008.
[47]
G. Buczkowski and D. S. Richmond, “The effect of urbanization on ant abundance and diversity: a temporal examination of factors affecting biodiversity,” PLoS ONE, vol. 7, no. 8, 2012.
[48]
P. Bolund and S. Hunhammar, “Ecosystem services in urban areas,” Ecological Economics, vol. 29, no. 2, pp. 293–301, 1999.
[49]
G. McGranahan, P. Marcotullio, X. Bai et al., “Urban systems,” in Ecosystems and Human Well-Being: Current State and Trends, R. Hassan, R. Scholes, and N. Ash, Eds., Island Press, Washington, DC, USA, 2006.
[50]
R. Forman, Urban Regions: Ecology and Planning Beyond the City, Cambridge University Press, New York, NY, USA, 2008.
[51]
G. McGranahan and D. Satterthwaite, “Urban centers: an assessment of sustainability,” Annual Review of Environment and Resources, vol. 28, pp. 243–274, 2003.
[52]
P. Clergeau, J.-P. L. Savard, G. Mennechez, and G. Falardeau, “Bird abundance and diversity along an urban-rural gradient: a comparative study between two cities on different continents,” Condor, vol. 100, no. 3, pp. 413–425, 1998.
[53]
S. P. D. Riley, R. M. Sauvajot, T. K. Fuller et al., “Effects of urbanization and habitat fragmentation on bobcats and coyotes in southern California,” Conservation Biology, vol. 17, no. 2, pp. 566–576, 2003.
[54]
UNPD (United Nations Population Division), World Urbanization Prospects: The 2005 Revision, New York, NY, USA, 2005.
W. Thuiller, S. Lavorel, M. B. Araújo, M. T. Sykes, and I. C. Prentice, “Climate change threats to plant diversity in Europe,” Proceedings of the National Academy of Sciences of the United States of America, vol. 102, no. 23, pp. 8245–8250, 2005.
[57]
J. Memmott, P. G. Craze, N. M. Waser, and M. V. Price, “Global warming and the disruption of plant-pollinator interactions,” Ecology Letters, vol. 10, no. 8, pp. 710–717, 2007.
[58]
S. Dubey and R. Shine, “Restricted dispersal and genetic diversity in populations of an endangered montane lizard (Eulamprus leuraensis, Scincidae),” Molecular Ecology, vol. 19, no. 5, pp. 886–897, 2010.
[59]
T. J. Davies, G. F. Smith, D. U. Bellstedt et al., “Extinction risk and diversification are linked in a plant biodiversity hotspot,” PLoS Biology, vol. 9, no. 5, Article ID e1000620, 2011.
[60]
K. J. Feeley and M. R. Silman, “Land-use and climate change effects on population size and extinction risk of Andean plants,” Global Change Biology, vol. 16, no. 12, pp. 3215–3222, 2010.
[61]
B. S. P. Wang, P. J. Charest, and B. Downie, “Ex situ storage of seeds, pollen, and in vitro cultures of perennial woody plant species,” FAO Forestry Paper, vol. 113, p. 83, 1993.
[62]
L. Glowka, F. Burhene-Guilmann, H. Synge, J. A. McNeely, and L. Gündling, A Guide To the Convention on Biological Diversity (Environmental Policy and Law Paper No. 30), IUCN, Switzerland, 1994.
[63]
United Nations Conference on Environment and Development (UNCED), Convention on Biological Diversity, Geneva, Switzerland, 1992.
[64]
P. K. Pati, S. P. Rath, M. Sharma, A. Sood, and P. S. Ahuja, “In vitro propagation of rose—a review,” Biotechnology Advances, vol. 24, no. 1, pp. 94–114, 2006.
[65]
W. Grout and W. Brian, “Meristem-tip culture for propagation and virus elimination,” in Methods in Molecular Biology, R. D. Hall, Ed., pp. 115–125, Plant Cell Culture Protocol, Humana Press, Totowa, NJ, USA, 1999.
[66]
G. Faccioli, “Control of potato viruses using meristem and stem-cutting cultures, thermotherapy and chemotherapy,” in Virus and Virus-Like Diseases of Potatoes and Production of Seed Potatoes, G. Loebenstein, H. P. Berger, A. A. Brunt, and G. R. Lawsan, Eds., pp. 365–390, Kluwer Academic Publisher, Dordrecht, The Netheralands, 2001.
[67]
G. R. Rout, A. Mohapatra, and S. M. Jain, “Tissue culture of ornamental pot plant: a critical review on present scenario and future prospects,” Biotechnology Advances, vol. 24, no. 6, pp. 531–560, 2006.
[68]
C. M. Miguel, Adventitious regeneration and genetic transformation of almond (Prunus dulcis Mill.) [Ph.D. thesis], Faculty of Sciences, University of Lisbon, Lisbon, Portugal, 1998.
[69]
O. McMeans, R. M. Skirvin, A. Otterbacher, and G. Mitiku, “Assessment of tissue culture-derived 'Gala' and ‘Royal Gala’ apples (Malus x domestica Borkh.) for somaclonal variation,” Euphytica, vol. 103, no. 2, pp. 251–257, 1998.
[70]
O. Monteuuis, F. C. Baurens, D. K. S. Gogh, M. Quimado, S. Doulbeau, and J. L. Verdeil, “DNA methylation in acacia mangium in vitro and ex-vitro buds, in relation to their within-shoot position, age and leaf morphology of the shoots,” Silvae Genetica, vol. 58, no. 5-6, pp. 287–292, 2009.
[71]
B. Renau-Morata, S. G. Nebauer, I. Arrillaga, and J. Segura, “Assessments of somaclonal variation in micropropagated shoots of Cedrus: consequences of axillary bud breaking,” Tree Genetics and Genomes, vol. 1, no. 1, pp. 3–10, 2005.
[72]
V. Rani and S. N. Raina, “Genetic analysis of enhanced-axillary-branching-derived Eucalyptus tereticornis Smith and E. camaldulensis Dehn. plants,” Plant Cell Reports, vol. 17, no. 3, pp. 236–242, 1998.
[73]
F. Zhang, Y. Lv, H. Dong, and S. Guo, “Analysis of genetic stability through intersimple sequence repeats molecular markers in micropropagated plantlets of Anoectochilus formosanus Hayata, a medicinal plant,” Biological and Pharmaceutical Bulletin, vol. 33, no. 3, pp. 384–388, 2010.
[74]
P. Joshi and V. Dhawan, “Assessment of genetic fidelity of micropropagated Swertia chirayita plantlets by ISSR marker assay,” Biologia Plantarum, vol. 51, no. 1, pp. 22–26, 2007.
[75]
R. M. Devarumath, S. Nandy, V. Rani, S. Marimuthu, N. Muraleedharan, and S. N. Raina, “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 Reports, vol. 21, no. 2, pp. 166–173, 2002.
[76]
M. K. Panda, S. Mohanty, E. Subudi, L. Acharya, and S. Nayak, “Assessment of genetic stability of micropropagated plants of Curcuma longa L. by cytophotometry and RAPD analyses,” International Journal of Integrative Biology, vol. 1, no. 3, pp. 189–195, 2007.
[77]
W. Xing, M. Bao, H. Qin, and G. Ning, “Micropropagation of Rosa rugosa through axillary shoot proliferation,” Acta Biologica Cracoviensia Series Botanica, vol. 52, no. 2, pp. 69–75, 2010.
[78]
M. N. Nas, N. Mutlu, and P. E. Read, “Random amplified polymorphic DNA (RAPD) analysis of long-term cultured hybrid hazelnut,” HortScience, vol. 39, no. 5, pp. 1079–1082, 2004.
[79]
M. H. N. Mollel and E. M. A. Goyvaerts, “Micropropagation of marula, Sclerocarya birrea subsp. caffra (Anarcadiaceae) by axillary bud proliferation and random amplified polymorphic DNA (RAPD) analysis of plantlets,” African Journal of Biotechnology, vol. 11, no. 93, pp. 16003–16012, 2012.
[80]
C. Chen, J. Lan, S. Xie, S. Cui, and A. Li, “In vitro propagation and quality evaluation of long-term micro-propagated and conventionally grown Fagopyrum dibotrys Hara mutant, an important medicinal plant,” Journal of Medicinal Plants Research, vol. 6, no. 15, pp. 3003–3012, 2012.
[81]
S. Mohanty, R. K. Joshi, E. Subudi, S. Sahoo, and S. Nayak, “Genetic stability assessment of micropropagated mango Ginger (Curcuma amada Roxb.) through RAPD and ISSR markers,” Research Journal of Medicinal Plant, vol. 6, no. 7, pp. 529–536, 2012.
[82]
R. Parida, S. Mohanty, and S. Nayak, “Evaluation of genetic fidelity of in vitro propagated greater galangal (Alpinia galangal L.) using DNA based markers,” International Journal of Plant, Animal and Environmental Sciences, vol. 1, no. 3, pp. 123–133, 2011.
[83]
R. Gupta, M. Modgil, and S. K. Chakrabarti, “Assessment of genetic fidelity of micropropagated apple rootstock plants, EMLA 111, using RAPD markers,” Indian Journal of Experimental Biology, vol. 47, no. 11, pp. 925–928, 2009.
[84]
H. Pathak and V. Dhawan, “ISSR assay for ascertaining genetic fidelity of micropropagated plants of apple rootstock Merton 793,” In Vitro Cellular and Developmental Biology, vol. 48, no. 1, pp. 137–143, 2012.
[85]
H. Lata, S. Chandra, N. Techen, I. A. Khan, and M. A. Elsohly, “Assessment of the genetic stability of micropropagated plants of cannabis sativa by ISSR markers,” Planta Medica, vol. 76, no. 1, pp. 97–100, 2010.
[86]
E. L. Peredo, R. Arroyo-García, and M. á. Revilla, “Epigenetic changes detected in micropropagated hop plants,” Journal of Plant Physiology, vol. 166, no. 10, pp. 1101–1111, 2009.
[87]
S. Gantait, N. Mandal, and P. K. Das, “Field evaluation of micropropagated vs. conventionally propagated elephant garlic,” Journal of Agricultural Technology, vol. 7, no. 1, pp. 97–103, 2010.
[88]
A. Kongbangkerd, A. K?pf, P. Allacher, C. Wawrosch, and B. Kopp, “Micropropagation of squill (Charybdis numidica) through nodule culture,” Plant Cell Reports, vol. 23, no. 10-11, pp. 673–677, 2005.
[89]
L. Annarita, “Morphological evaluation of olive plants propagated in vitro culture through axillary buds and somatic embryogenesis methods,” African Journal of Plant Science, vol. 3, no. 3, pp. 037–043, 2009.
[90]
S. Goto, R. C. Thakur, and K. Ishii, “Determination of genetic stability in long-term micropropagated shoots of Pinus thunbergii Parl. using RADP markers,” Plant Cell Reports, vol. 18, no. 3-4, pp. 193–197, 1998.
[91]
E. V. Soniya and M. R. Das, “In vitro micropropagation of Piper longum—an important medicinal plant,” Plant Cell, Tissue and Organ Culture, vol. 70, no. 3, pp. 325–327, 2002.
[92]
L. A. Caro, P. A. Polci, L. I. Lindstr?m, C. V. Echenique, and L. F. Hernández, “Micropropagation of Prosopis chilensis (Mol.) Stuntz from young and mature plants,” Biocell, vol. 26, no. 1, pp. 25–33, 2002.
[93]
P. Bregitzer, S. Zhang, M.-J. Chob, and P. G. Lemaux, “Reduced somaclonal variation in barley is associated with culturing highly differentiated, meristematic tissues,” Crop Science, vol. 42, no. 4, pp. 1303–1308, 2002.
[94]
R. Medina, M. Faloci, M. A. Marassi, and L. A. Mroginski, “Genetic stability in rice micropropagation,” Biocell, vol. 28, no. 1, pp. 13–20, 2004.
[95]
M. Martins, D. Sarmento, and M. M. Oliveira, “Genetic stability of micropropagated almond plantlets, as assessed by RAPD and ISSR markers,” Plant Cell Reports, vol. 23, no. 7, pp. 492–496, 2004.
[96]
S. Dohling, S. Kumaria, and P. Tandon, “Multiple shoot induction from axillary bud cultures of the medicinal orchid, Dendrobium longicornu,” AoB PLANTS, vol. 2012, 2012.
[97]
R. K. Radha, A. M. Varghese, and S. Seeni, “Conservation through in vitro propagation and restoration of Mahonia leschenaultii, an endemic tree of the Western Ghats,” ScienceAsia, vol. 39, pp. 219–229, 2013.
[98]
H. S. Chawla, Introduction To Plant Biotechnology, Science press, 2nd edition, 2002.
[99]
T. Murashige, “Plant propagation through tissue culture,” Annual Review of Plant Physiology, vol. 25, pp. 135–166, 1974.
[100]
C. Y. Hu and P. J. Wang, “Handbook of plant cell culture,” in Meristem Shoot Tip and Bud Culture, D. A. Evans, W. R. Sharp, P. V. Ammirato, and Y. Yamada, Eds., pp. 177–227, Macmillan, New York, NY, USA, 1983.
[101]
G. J. De Klerk, “How to measure somaclonal variation,” Acta Botanica Neerlandica, vol. 39, pp. 129–144, 1990.
[102]
S. K. Sharma, G. J. Bryan, M. O. Winfield, and S. Millam, “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, vol. 226, no. 6, pp. 1449–1458, 2007.
[103]
P. J. Larkin and W. R. Scowcroft, “Somaclonal variation—a novel source of variability from cell cultures for plant improvement,” Theoretical and Applied Genetics, vol. 60, no. 4, pp. 197–214, 1981.
[104]
A. R. Gould, “Factors controlling generations of variability in vitro,” in Cell Culture and Somatic Cell Genetics in Plants. Plant Regeneration and Genetic Variability, I. K. Vasil, Ed., pp. 549–567, Academic, Orlando, Fla, USA, 1986.
[105]
S. M. Kaeppler, H. F. Kaeppler, and Y. Rhee, “Epigenetic aspects of somaclonal variation in plants,” Plant Molecular Biology, vol. 43, no. 2-3, pp. 179–188, 2000.
[106]
F. Ngezahayo, Y. Dong, and B. Liu, “Somaclonal variation at the nucleotide sequence level in rice (Oryza sativa L.) as revealed by RAPD and ISSR markers, and by pairwise sequence analysis,” Journal of Applied Genetics, vol. 48, no. 4, pp. 329–336, 2007.
[107]
H. Hirochika, K. Sugimoto, Y. Otsuki, H. Tsugawa, and M. Kanda, “Retrotransposons of rice involved in mutations induced by tissue culture,” Proceedings of the National Academy of Sciences of the United States of America, vol. 93, no. 15, pp. 7783–7788, 1996.
[108]
Z. L. Liu, F. P. Han, M. Tan et al., “Activation of a rice endogenous retrotransposon Tos17 in tissue culture is accompanied by cytosine demethylation and causes heritable alteration in methylation pattern of flanking genomic regions,” Theoretical and Applied Genetics, vol. 109, no. 1, pp. 200–209, 2004.
[109]
F. Ngezahayo, C. Xu, H. Wang, L. Jiang, J. Pang, and B. Liu, “Tissue culture-induced transpositional activity of mPing is correlated with cytosine methylation in rice,” BMC Plant Biology, vol. 9, article 91, 2009.
[110]
X. Wang, R. Wu, X. Lin et al., “Tissue culture-induced genetic and epigenetic alterations in rice pure-lines, F1 hybrids and polyploids,” BMC Plant Biology, vol. 13, p. 77, 2013.
[111]
Y. R. Mehta and D. C. Angra, “Somaclonal variation for disease resistance in wheat and production of dihaploids through wheat x maize hybrids,” Genetics and Molecular Biology, vol. 23, no. 3, pp. 617–622, 2000.
[112]
S. Predieri, “Mutation induction and tissue culture in improving fruits,” Plant Cell, Tissue and Organ Culture, vol. 64, no. 2-3, pp. 185–210, 2001.
[113]
A. Karp, “Somaclonal variation as a tool for crop improvement,” Euphytica, vol. 85, no. 1–3, pp. 295–302, 1995.
[114]
E. Unai, T. Iselen, and E. de Garcia, “Comparison of characteristics of bananas (Musa sp.) from the somaclone CIEN BTA-03 and its parental clone Williams,” Fruit, vol. 59, pp. 257–263, 2004.
[115]
T. J. Oh, M. A. Cullis, K. Kunert, I. Engelborghs, R. Swennen, and C. A. Cullis, “Genomic changes associated with somaclonal variation in banana (Musa spp.),” Physiologia Plantarum, vol. 129, no. 4, pp. 766–774, 2007.
[116]
G. Grafi and Y. Avivi, “Stem cells: a lesson from dedifferentiation,” Trends in Biotechnology, vol. 22, no. 8, pp. 388–389, 2004.
[117]
G. Grafi, A. Florentin, V. Ransbotyn, and Y. Morgenstern, “The stem cell state in plant development and in response to stress,” Frontiers in Plant Science, vol. 2, no. 53, pp. 1–10, 2011.
[118]
B. McClintock, “The significance of responses of the genome to challenge,” Science, vol. 226, no. 4676, pp. 792–801, 1984.
[119]
M. R. Ahuja, “Somaclonal genetics of forest trees,” in Somaclonal Variation and Induced Mutations in Crop Improvement, S. M. Jain, D. S. Brar, and B. S. Ahloowalia, Eds., pp. 105–121, Kluwer Academic, Dordrecht, The Netherlands, 1998.
[120]
V. Rani and S. N. Raina, “Genetic fidelity of organized meristem-derived micropropagated plants: a critical reappraisal,” In Vitro Cellular and Developmental Biology, vol. 36, no. 5, pp. 319–330, 2000.
[121]
K. Gillis, J. Gielis, H. Peeters, E. Dhooghe, and J. Oprins, “Somatic embryogenesis from mature Bambusa balcooa Roxburgh as basis for mass production of elite forestry bamboos,” Plant Cell, Tissue and Organ Culture, vol. 91, no. 2, pp. 115–123, 2007.
[122]
R. Parra, M. T. Pastor, E. Pérez-Payá, and J. B. Amo-Marco, “Effect of in vitro shoot multiplication and somatic embryogenesis on 5-methylcytosine content in DNA of Myrtus communis L.,” Plant Growth Regulation, vol. 33, no. 2, pp. 131–136, 2001.
[123]
C. Diaz-Sala, M. Rey, A. Boronat, R. Besford, and R. Rodriguez, “Variations in the DNA methylation and polypeptide patterns of adult hazel (Corylus avellana L.) associated with sequential in vitro subcultures,” Plant Cell Reports, vol. 15, no. 3-4, pp. 218–221, 1995.
[124]
X. Li, M. Xu, and S. S. Korban, “DNA methylation profiles differ between field- and in vitro-grown leaves of apple,” Journal of Plant Physiology, vol. 159, no. 11, pp. 1229–1234, 2002.
[125]
P. Smykal, L. Valledor, R. Rodríguez, and M. Griga, “Assessment of genetic and epigenetic stability in long-term in vitro shoot culture of pea (Pisum sativum L.),” Plant Cell Reports, vol. 26, no. 11, pp. 1985–1998, 2007.
[126]
M. Baránek, B. K?i?an, E. Ondru?íková, and M. Pidra, “DNA-methylation changes in grapevine somaclones following in vitro culture and thermotherapy,” Plant Cell, Tissue and Organ Culture, vol. 101, no. 1, pp. 11–22, 2010.
[127]
S. Y. Park, H. N. Murthy, D. Chakrabarthy, and K. Y. Paek, “Detection of epigenetic variation in tissue-culture-derived plants of Doritaenopsis by methylation-sensitive amplification polymorphism (MSAP) analysis,” In Vitro Cellular and Developmental Biology, vol. 45, no. 1, pp. 104–108, 2009.
[128]
A. H. C. Wong, I. I. Gottesman, and A. Petronis, “Phenotypic differences in genetically identical organisms: the epigenetic perspective,” Human Molecular Genetics, vol. 14, no. 1, pp. R11–R18, 2005.
[129]
E. A. Smith, S. B. Collette, T. A. Boynton et al., “Developmental contributions to phenotypic variation in functional leaf traits within quaking aspen clones,” Tree Physiology, vol. 31, no. 1, pp. 68–77, 2011.
[130]
S. Hirsch, R. Baumerger, and U. Grossniklaus, “Epigenetic variation,inheritance, and selection in plant populations,” Cold Spring Harbor Symposia on Quantitative Biology, vol. 77, pp. 97–104, 2012.
[131]
M. A. Palombi and C. Damiano, “Comparison between RAPD and SSR molecular markers in detecting genetic variation in kiwifruit (Actinidia deliciosa A. Chev),” Plant Cell Reports, vol. 20, no. 11, pp. 1061–1066, 2002.
[132]
J. A. Yoder, C. P. Walsh, and T. H. Bestor, “Cytosine methylation and the ecology of intragenomic parasites,” Trends in Genetics, vol. 13, no. 8, pp. 335–340, 1997.
[133]
R. A. Martienssen and V. Colot, “DNA methylation and epigenetic inheritance in plants and filamentous fungi,” Science, vol. 293, no. 5532, pp. 1070–1074, 2001.
[134]
R. I. S. Brettell and E. S. Dennis, “Reactivation of a silent Ac following tissue culture is associated with heritable alterations in its methylation pattern,” Molecular and General Genetics, vol. 229, no. 3, pp. 365–372, 1991.
[135]
M. J. M. Smulders and G. J. de Klerk, “Epigenetics in plant tissue culture,” Plant Growth Regulation, vol. 63, no. 2, pp. 137–146, 2011.
[136]
M. Alizadeh and S. K. Singh, “Molecular assessment of clonal fidelity in micropropagated grape (Vitis spp.) rootstock genotypes using RAPD and ISSR markers,” Iranian Journal of Biotechnology, vol. 7, no. 1, pp. 37–44, 2009.
[137]
A. E. Mohamed, “Somaclonal variation in micropropagated strawberry detected at the molecular level,” International Journal of Agricultural and Biological Engineering, vol. 9, no. 5, pp. 721–725, 2007.
[138]
S. Saha, T. Dey, and P. Ghosh, “Micropropagation of Ocimum kilimandscharicum guerke (Labiatae),” Acta Biologica Cracoviensia Series Botanica, vol. 52, no. 2, pp. 50–58, 2010.