Skip to main content
SpringerLink
Log in

Morphogenic Aspects of Somatic Embryogenesis

  • Chapter
In Vitro Embryogenesis in Plants

Part of the book series: Current Plant Science and Biotechnology in Agriculture ((PSBA,volume 20))

Abstract

Somatic embryogenesis is the formation of an embryo from a cell other than a gamete or the product of gametic fusion. This is not an artificial phenomenon, and is known in nature as a form of apomixis called adventitious embryony, first described by Strasburger in 1878 [1] (see Sharma and Thorpe, this volume). Although Steward [2] and Reinert [3] are generally given credit for the first descriptions of somatic embryogenesis in 1958, such credit might more properly belong to Levine [4], who in 1947 reported the recovery of carrot “seedlings” from tissues exposed to low levels of α-naphthaleneacetic acid, via a process whose description sounds very much like somatic embryogenesis.

This is a preview of subscription content, log in via an institution to check access.

Access this chapter

Log in via an institution
Chapter
USD 29.95
Price excludes VAT (USA)
  • Available as PDF
  • Read on any device
  • Instant download
  • Own it forever
eBook
USD 169.00
Price excludes VAT (USA)
  • Available as PDF
  • Read on any device
  • Instant download
  • Own it forever
Softcover Book
USD 219.99
Price excludes VAT (USA)
  • Compact, lightweight edition
  • Dispatched in 3 to 5 business days
  • Free shipping worldwide - see info
Hardcover Book
USD 219.99
Price excludes VAT (USA)
  • Durable hardcover edition
  • Dispatched in 3 to 5 business days
  • Free shipping worldwide - see info

Tax calculation will be finalised at checkout

Purchases are for personal use only

Institutional subscriptions

Preview

Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

References

  1. Strasburger, E., Über Polyembryonie, Jenaische Z. Naturwiss., 12, 647, 1878.

    Google Scholar 

  2. Steward, F.C., Growth and development of cultivated cells. III. Interpretations of the growth from free cell to carrot plant, Am. J. Bot., 45, 709, 1958.

    Google Scholar 

  3. Reinert, J., Morphogenese and ihre Kotrolle an Gewebekulturen aus Carotten, Naturwiss., 45, 344, 1958.

    CAS  Google Scholar 

  4. Levine, M., Differentiation of carrot root tissue grown in culture, Bull. Torrey Bot. Club, 74, 321, 1947.

    Google Scholar 

  5. Burns, G.W., The Plant Kingdom, Macmillan Publishing Co., Inc., New York, 1974.

    Google Scholar 

  6. Wing, S.L., Hickey, L.J., and Swisher, C.C., Implications of an exceptional fossil flora for Late Cretaceous vegetation, Nature, 363, 342, 1993.

    Google Scholar 

  7. Goldberg, R.B., Barker, S.J., and Perez-Grau, L., Regulation of gene expression during plant embryogenesis, Cell, 56, 149, 1989.

    PubMed  CAS  Google Scholar 

  8. Hughes, D.W., and Galau, G.A., Temporally modular gene expression during cotyledon development, Genes Dev., 3, 358, 1989.

    PubMed  CAS  Google Scholar 

  9. Kermode, A.R., Regulatory mechanisms involved in the transition from seed development to maturation, Crit. Rev. Pl. Sci., 9, 155, 1990.

    CAS  Google Scholar 

  10. Kohlenbach, H., Comparative somatic embryogenesis, in Frontiers of Plant Tissue Culture, Thorpe, T.A., Ed., University of Calgary, Calgary, 1978, 59.

    Google Scholar 

  11. Goldberg, R.B., Regulation of plant gene expression, Phil. Trans. R. Soc. Lond. B, 314, 343, 1986.

    CAS  Google Scholar 

  12. Sharp, W.R., Evans, D.A., and Sondahl, M.R., Application of somatic embryogenesis to crop improvement, in Plant Tissue Culture 1983. Proceedings of the Fifth International Congress of Plant Tissue Culture, Fujiwara, A., Ed., Japanese Association for Plant Tissue Culture, 1982, 759.

    Google Scholar 

  13. Evans, D.A., Sharp, W.R., and Flick, C.E., Growth and behaviour of cell cultures. Embryogenesis and organogenesis, in Plant Tissue Culture: Methods and Applications in Agriculture, Thorpe, T.A., Ed., Academic Press, New York, 1981, 45.

    Google Scholar 

  14. Williams, E.G., and Maheswaran, G., Somatic embryogenesis: factors influencing coordinated behaviour of cells as an embryogenic group, Ann. Bot., 57, 443, 1986.

    Google Scholar 

  15. Carman, J.G., Embryogenic cells in plant tissue cultures: occurrence and behavior, In Vitro Cell. Dev. Biol., 26, 746, 1990.

    Google Scholar 

  16. Merkle, S.A., Parrott, W.A., and Williams, E.G., Applications of somatic embryogenesis and embryo cloning, in Plant tissue culture: Applications and limitations, Bhojwani, S.S., Ed., Elsevier Science Publishers, Amsterdam, 1990, 67.

    Google Scholar 

  17. LoSchiavo, F., Pitto, L., Giuliano, G., Torti, G., Nuti-Ronchi, V., Marazziti, D., Vergara, R., Orselli, S., and Terzi, M., DNA methylation of embryogenic carrot cell cultures and its variations as caused by mutation, differentiation, hormones and hypomethylating drugs, Theor. Appl. Genet., 77, 325, 1989.

    Google Scholar 

  18. Pechan, P.M., Bartels, D., Brown, D.C.W., and Schell, J., Messenger-RNA and protein changes associated with induction of Brassica microspore embryogenesis, Planta, 184, 161, 1991.

    CAS  Google Scholar 

  19. Kamada, H., Kobayashi, K., Kiyosue, T., and Harada, H., Stress induced somatic embryogenesis in carrot and its application to synthetic seed production, In Vitro Cell. Dev. Biol., 25, 1163, 1989.

    Google Scholar 

  20. Imamura, J., and Harada, H., Stimulation of tobacco pollen embryogenesis by anaerobic treatments, J. Plant Physiol., 103, 259, 1981.

    Google Scholar 

  21. Hakman, I., and Von Arnold, S., Plantlet regeneration through somatic embryogenesis in Picea abies (Norway spruce), J. Plant PhysioL, 121, 149, 1985.

    CAS  Google Scholar 

  22. Herberle-Bors, E., In vitro haploid formation from pollen: a critical review, Theor. Appl. Genet., 71, 361, 1985.

    Google Scholar 

  23. Nuti Ronchi, V., Giorgetti, L., Tonelli, M., and Martini, G., Ploidy reduction and genome segregation in cultured carrot cell lines. II. Somatic meiosis, Plant Cell Tissue Organ Cult., 30, 115, 1992.

    Google Scholar 

  24. Bonnelle, C., Lejeune, F., Fournier, D., and Tourte, Y., Remaniements infrastructuraux des cellules cotylédonaires en culture chez deux légumineuses; relations avec l’acquisition de potentialités embryogènes, C.R. Acad. Sci. Paris, 310, 657, 1990.

    Google Scholar 

  25. Vergara, R., Verde, F., Pitto, L., Lo Schiavo, F., and Terzi, M., Reversible variations in the methylation pattern of carrot DNA during somatic embryogenesis, Plant Cell Rep., 8, 697, 1990.

    CAS  Google Scholar 

  26. Smith, D.L., and Krikorian, A.D., Release of somatic embryogenic potential from excised zygotic embryos of carrot and maintenance of proembryonic cultures in hormone-free medium, Am. J. Bot., 76, 1832, 1989.

    PubMed  CAS  Google Scholar 

  27. Trigiano, R.N., Gray, D.J., Conger, B.V., and Mc Daniel, J.K., Origin of direct somatic embryos from cultured leaf segments of Dactylis glomerata, Bot. Gaz., 150, 72, 1989.

    Google Scholar 

  28. Dubois, T., Guedira, M., Dubois, J., and Vasseur, J., Direct somatic embryogenesis in roots of Cichorium: Is callose an early marker? Ann. Bot., 65, 539, 1990.

    Google Scholar 

  29. Dubois, T., Guedira, M., and Vasseur, J., Direct somatic embryogenesis in leaves of Cichorium, Protoplasma, 162, 120, 1991.

    Google Scholar 

  30. Wetherell, D.F., Enhanced adventive embryogenesis resulting from plasmolysis of cultured wild carrot cells, Plant Cell Tiss. Org. Cult., 3, 221, 1984.

    Google Scholar 

  31. Tsukahara, M., and Hirosawa, T., Simple dehydration treatment promotes plantlet regeneration of rice (Oryza sativa L.) callus, Plant Cell Rep., 11, 550, 1992.

    Google Scholar 

  32. Chuang, M.-J., and Chang, W.-C., Somatic embryogenesis and plant regeneration in callus culture derived from immature seeds and mature zygotic embryos of Dysosma pleintha (Hance) Woodson, Plant Cell Rep., 6, 484, 1987.

    Google Scholar 

  33. Gharyal, P.K., and Maheshwari, S.C., In vitro differentiation of somatic embryoids in a leguminous tree — Albizzia lebbeck L, Naturwiss., 68, 379, 1983.

    Google Scholar 

  34. Dhanalakshmi, S., and Lakshmanan, K.K., In vitro somatic embryogenesis and plant regeneration in Clitoria ternatea, J. Exp. Bot., 43, 213, 1992.

    Google Scholar 

  35. Street, H.E., and Withers, L.A., The anatomy of embryogenesis in culture, in Tissue Culture and Plant Science, Street, H.E., Ed., Academic Press, London/New York, 1974, 71.

    Google Scholar 

  36. Haccius, B., Question of unicellular origin of non-zygotic embryos in callus cultures, Phytomorphology, 28, 74, 1977.

    Google Scholar 

  37. Vasil, I.K., Developing cell and tissue culture systems for the improvement of cereal and grass crops, J. Plant Physiol., 128, 193, 1987.

    Google Scholar 

  38. Hartweck, L.M., Lazzeri, P.A., Cui, D., Collins, G.B., and Williams, E.G., Auxin-orientation effects on somatic embryogenesis from immature soybean cotyledons, In Vitro Cell. Dev. Biol., 24, 821, 1988.

    CAS  Google Scholar 

  39. Sato, S., Newell, C., Kolacz, K., Tredo, L., Finer, J., and Hinchee, M., Stable transformation via particle bombardment in two different soybean regeneration systems, Plant Cell Rep., 12, 408, 1993.

    CAS  Google Scholar 

  40. Polito, V.S., Mc Granahan, G., Pinney, K., and Leslie, C., Origin of somatic embryos from repetitively embryogenic cultures of walnut (Junglans regia L.): implications for Agrobacteriummediated transformation, Plant Cell Rep., 8, 219, 1989.

    Google Scholar 

  41. Norgaard, J.V., and Krogstrup, P., Cytokinin induced somatic embryogenesis from immature embryos of Abies nordmanniana Lk, Plant Cell Rep., 9, 509, 1991.

    CAS  Google Scholar 

  42. Maheswaran, G., and Williams, E.G., Direct somatic embryoid formation on immature embryos of Trifolium repens, T. pratense and Medicago sativa, and rapid clonal propagation of T. repens, Ann. Bot., 54, 201, 1984.

    Google Scholar 

  43. Maheswaran, G., and Williams, E.G., Origin and development of somatic embryoids formed directly on immature embryos of Trifolium repens in vitro, Ann. Bot., 56, 619, 1985.

    Google Scholar 

  44. Mo, L.H., and Von Arnold, S., Origin and development of embryogenic cultures from seedlings of Norway spruce, J. Plant Physiol., 138, 223, 1991.

    Google Scholar 

  45. Saunders, J.W., and Bingham, E.T., Growth regulator effects on bud initiation in callus cultures of Medicago sativa, Am. J. Bot., 62, 850, 1975.

    Google Scholar 

  46. Walker, K.A., Yu, P.C., Sato, S.J., and Jaworski, E.G., The hormonal control of organ formation in callus of Medicago sativa L. cultured in vitro, Am. J. Bot., 65, 654, 1982.

    Google Scholar 

  47. Saxena, P.K., and King, J., Morphogenesis in lentil: plant regeneration from callus cultures of Lens culinaris Medik. via somatic embryogenesis, Plant Sci., 52, 223, 1987.

    CAS  Google Scholar 

  48. Rajasekaran, K., Hein, M.B., Davis, G.C., Carnes, M.G., and Vasil, I.K., Endogenous growth regulators in leaves and tissue cultures of Pennisetum purpureum Schum, J. Plant Physiol., 130, 13, 1987.

    CAS  Google Scholar 

  49. Wenck, A.R., Conger, B.V., Trigiano, R.N., and Sams, C.E., Inhibition of somatic embryogenesis in orchardgrass by endogenous cytokinins, Plant Sci., 88, 990, 1988.

    CAS  Google Scholar 

  50. Arcioni, S., and Mariotti, D., Tissue culture and plant regeneration in the forage legumes Onobrychis viciaefolia Scop., Coronilla varia and Lotus corniculatus L, in Plant Tissue Culture, Fujiwara, A., Ed., Japanese Association for Plant Tissue Culture, Tokyo, 1982, 707.

    Google Scholar 

  51. Kysely, W., and Jacobsen, H.-J., Somatic embryogenesis from pea embryos and shoot apices, Plant Cell Tiss. Org. Cult., 20, 7, 1990.

    CAS  Google Scholar 

  52. Lippmann, B., and Lippmann, G., Induction of somatic embryos in cotyledonary tissue of soybean, Glycine max L. Merr., Plant Cell Rep., 3, 215, 1984.

    CAS  Google Scholar 

  53. Lazzeri, P.A., Hildebrand, D.F., and Collins, G.B., Soybean somatic embryogenesis: effects of hormones and culture manipulations, Plant Mol. Biol. Rep., 10, 197, 1987.

    CAS  Google Scholar 

  54. Nadolska-Orczyk, A., Somatic embryogenesis of agriculturally important lupin species (Lupinus angustifolius, L. albus, L. mutabilis), Plant Cell Tiss. Org. Cult., 28, 19, 1992.

    CAS  Google Scholar 

  55. Parrott, W.A., Merkle, S.A., and Williams, E.G., Somatic embryogenesis: potential for use in propagation and gene transfer systems, in Advanced Methods in Plant Breeding and Biotechnology, Murray, D.R., Ed., CAB International, Wallingford, 1991, 158.

    Google Scholar 

  56. Komatsuda, T., Kaneko, K., and Oka, S., Genotype x sucrose interactions for somatic embryogenesis in soybean, Crop Sci., 31, 333, 1991.

    CAS  Google Scholar 

  57. Pretova, A., and Williams, E.G., Direct somatic embryogenesis from immature embryos of flax (Linum usitatissimum L.), J. Plant Physiol., 126, 155, 1986.

    CAS  Google Scholar 

  58. Young, R., Kaul, V., and Williams, E.G., Clonal propagation in vitro from immature emryos and flower buds of Lycopersicon peruvianum and L. esculentum, Plant Sci., 52, 237, 1987.

    CAS  Google Scholar 

  59. Ghazi, T.D., Cheema, H.V., and Nabors, M.W., Somatic embryogenesis and plant regeneration from embryogenic callus of soybean, Glycine max L., Plant Cell Rep., 5, 452, 1986.

    CAS  Google Scholar 

  60. Ranch, J.P., Oglesby, L., and Zielinski, A.C., Plant regeneration from embryo-derived tissue cultures of soybean by somatic embryogenesis, In Vitro Cell. Dev. Biol., 21, 653, 1985.

    Google Scholar 

  61. Barwale, U.B., Kerns, H.R., and Widholm, J.M., Plant regeneration from callus cultures of several soybean genotypes via embryogenesis and organogenesis, Planta, 167, 473, 1986.

    CAS  Google Scholar 

  62. Roberts, D.R., Flinn, B.S., Webb, D.T., Webster, F.B., and Sutton, B.C.S., Characterization of immature embryos of interior spruce by SDS-PAGE and microscopy in relation to their competence for somatic embryogenesis, Plant Cell Rep., 8, 285, 1989.

    CAS  Google Scholar 

  63. Sotak, R.J., Sommer, H.E., and Merkle, S.A., Relation of the developmental stage of zygotic embryos of yellow-poplar to their somatic embryogenic potential, Plant Cell Rep., 10, 175, 1991.

    Google Scholar 

  64. Karunaratne, S., Gamage, C., and Kovoor, A., Leaf maturity, a critical factor in somatic embryogenesis, J. Plant Physiol., 139, 27, 1991.

    Google Scholar 

  65. Williams, E.G., Collins, G.B., and Myers, J.R., 2. Clovers (Trifolium spp.), in Biotechnology in Agriculture and Forestry, Vol. 10, Legumes and Oilseed Crops, I., Bajaj, Y.P.S., Ed., SpringerVerlag, Berlin/Heidelberg, 1990, 242.

    Google Scholar 

  66. Jacobsen, H.-J., Somatic embryogenesis in seed legumes: The possible role of soluble auxin receptors, Israel J. Bot., 40, 139, 1991.

    CAS  Google Scholar 

  67. Tremblay, F.M., Somatic embryogenesis and plantlet regeneration from embryos isolated from stored seeds of Picea glauca, Can. J. Bot., 68, 236, 1990.

    CAS  Google Scholar 

  68. Reynolds, T.L., Interactions between calcium and auxin during pollen androgenesis in anther cultures of Solanum carolinense L, Plant Sci., 72, 109, 1990.

    CAS  Google Scholar 

  69. Jansen, M.A.K., Booij, H., Schel, J.H.N., and De Vries, S.C., Calcium increases the yield of somatic embryos in carrot embryogenic suspension cultures, Plant Cell Rep., 9, 221, 1990.

    CAS  Google Scholar 

  70. Lazzeri, P.A., Hildebrand, D.F., Sunega, J., Williams, E.G., and Collins, G.B., Soybean somatic embryogenesis: interactions between sucrose and auxin, Plant Cell Rep., 7, 517, 1988.

    CAS  Google Scholar 

  71. Genga, A., and Allavena, A., Factors affecting morphogenesis from immature cotyledons of Phaseolus coccineus L, Plant Cell Tiss. Org. Cult., 27, 189, 1991.

    CAS  Google Scholar 

  72. Kochba, J., Spiegel-Roy, P., Neumann, H., and Saad, S., Effect of carbohydrates on somatic embryogenesis in subcultured nucellar callus of Citrus cultivars, Z. Pflanzenphysiol., 105, 359, 1982.

    CAS  Google Scholar 

  73. Strickland, S.G., Nichol, J.W., Mc Call, C.M., and Stuart, D.A., Effect of carbohydrate source on alfalfa somatic embryogenesis, Plant Sci., 48, 113, 1987.

    CAS  Google Scholar 

  74. Wetherell, D.F., and Dougall, D.K., Sources of nitrogen supporting growth and embryogenesis in cultured wild carrot tissue, Physiol. Plant., 37, 97, 1976.

    CAS  Google Scholar 

  75. Tremblay, L., and Tremblay, F.M., Effects of gelling agents, ammonium nitrate, and light on the development of Picea mariana (Mill) B.S.P. (black spruce) and Picea rubens Sarg. (red spruce) somatic embryos, Plant Sci., 77, 233, 1991.

    CAS  Google Scholar 

  76. Stuart, D.A., and Strickland, S.G., Somatic embryogenesis from cell cultures of Medicago sativa 1. the role of amino acid additions to the regeneration medium, Plant Sci. Lett., 34, 165, 1984.

    CAS  Google Scholar 

  77. Stuart, D.A., and Strickland, S.G., Somatic embryogenesis from cell cultures of Medicago sativa 2. The interaction of amino acids with ammonium, Plant Sci. Lett., 34, 175, 1984.

    CAS  Google Scholar 

  78. Walker, K.A., and Sato, S.J., Morphogenesis in callus tissue of Medicago sativa: the role of ammonium ion in somatic embryogenesis, Plant Cell Tiss. Org. Cult., 1, 109, 1981.

    CAS  Google Scholar 

  79. Meijer, E.G.M., and Brown, D.C.W., Role of exogenous reduced nitrogen and sucrose in rapid high frequency somatic embryogenesis in Medicago sativa, Plant Cell Tissue Organ Cult., 10, 11, 1987.

    CAS  Google Scholar 

  80. Greinwald, R., and Czygan, F.-C., Regeneration of plantlets from callus cultures of Chamaecytisus purpureus and Chamaecytisus austriacus (Leguminosae), Bot. Acta, 104, 64, 1991.

    CAS  Google Scholar 

  81. Trigiano, R.N., Beaty, R.M., and Graham, E.T., Somatic embryogenesis from immature embryos of redbud (Cercis canadensis), Plant Cell Rep., 7, 148, 1988.

    CAS  Google Scholar 

  82. Trigiano, R.N., and Conger, B.V., Regulation of growth and somatic embryogenesis by proline and serine in suspension cultures of Dactylis glomerata, J. Plant Physiol., 130, 49, 1987.

    CAS  Google Scholar 

  83. Gleddie, S., Keller, W., and Setterfield, G., Somatic embryogenesis and plant regeneration from leaf explants and cell suspensions of Solanum melongena (eggplant), Can. J. Bot., 61, 656, 1983.

    CAS  Google Scholar 

  84. Michler, C.H., and Bauer, E.O., High frequency somatic embryogenesis from leaf tissue of Populus spp, Plant Sci., 77, 111, 1991.

    CAS  Google Scholar 

  85. Arumugan, N., and Bhojwani, S.S., Somatic embryogenesis in tissue cultures of Podophyllum hexandrum, Can. J. Bot., 68, 487, 1990.

    Google Scholar 

  86. Verhagen, S.A., and Wann, S.R., Norway spruce somatic embryogenesis: high-frequency initiation from light-cultured mature embryos, Plant Cell Tiss. Org. Cult., 16, 103, 1989.

    Google Scholar 

  87. Calero, N., Actions de radiations rouges et bleues sur l’embryogenèse somatique du Palmier dattier (Phoenix dactylifera L.) en culture in vitro et sur sa teneur en leucoanthocyanes, C.R. Soc. Biol., 183, 307, 1989.

    Google Scholar 

  88. Michler, C.H., and Lineberger, R.D., Effects of light on somatic embryo development and abscisic levels in carrot suspension cultures, Plant Cell Tiss. Org. Cult., 11, 189, 1987.

    Google Scholar 

  89. Lazzeri, P.A., Hildebrand, D.F., and Collins, G.B., Soybean somatic embryogenesis: effects of nutritional, physical and chemical factors, Plant Mol. Biol. Rep., 10, 209, 1987.

    CAS  Google Scholar 

  90. Roustan, J.P., Latche, A., and Fallot, J., Stimulation of Daucus carota somatic embryogenesis by inhibitors of ethylene synthesis: cobalt and nickel, Plant Cell Rep., 8, 182, 1989.

    CAS  Google Scholar 

  91. Roustan, J.-P., Latche, A., and Fallot, J., Inhibition of ethylene production and stimulation of carrot somatic embryogenesis by salicylic acid, Biol. Plant., 32, 273, 1990.

    CAS  Google Scholar 

  92. Auboiron, E., Carron, M.-P., and Michaux-Ferrière, M., Influence of atmospheric gases, particularly ethylene, on somatic embryogenesis of Hevea brasiliensis, Plant Cell Tiss. Org. Cult., 21, 31, 1990.

    CAS  Google Scholar 

  93. Carman, J.G., The in ovulo environment and its relevance to cloning wheat via somatic embryogenesis, In Vitro Cell. Dev. Biol., 25, 1155, 1989.

    Google Scholar 

  94. Carman, J.G., Improved somatic embryogenesis in wheat by partial simulation of the in-ovulo oxygen, growth-regulator and desiccation environments, Planta, 175, 417, 1988.

    CAS  Google Scholar 

  95. Yang, Y.-S., Wada, K., Goto, M., and Futsuhara, Y., In vitro formation of nodular calli in soybean (Glycine max L.) induced by cocultivated Pseudomonas maltophilia, Japan J. Breed., 41, 595, 1991.

    Google Scholar 

  96. Wake, H., Umetsu, H., Ozeki, Y., Shimomura, K., and Matsunaga, T., Extracts of marine cyanobacteria stimulated somatic embryogenesis of Daucus carota L, Plant Cell Rep., 9, 655, 1991.

    Google Scholar 

  97. Mc Coy, T.J., Tissue culture evaluation of sodium chloride tolerance in Medicago sp cellular versus whole plant response, Plant Cell Rep., 6, 31, 1987.

    Google Scholar 

  98. Mohan Ram, H.Y., Mehta, U., Ramanuja Rao, I.V., and Narasimham, M., Haploid induction in legumes, in Plant Tissue Culture, Fujiwara A., Ed., Japanese Association for Plant Tissue Culture, Tokyo, 1982, 541.

    Google Scholar 

  99. Halperin, W., Alternative morphogenetic events in cell suspensions, Am. J. Bot., 53, 443, 1966.

    Google Scholar 

  100. Mc William, A.A., Smith, S.H., and Street, H.E., The origin and development of embryoids in suspension culutres of carrot (Daucus carota), Ann. Bot., 38, 243, 1974.

    Google Scholar 

  101. Halperin, W., and Wetherell, D.F., Adventive embryony in tissue cultures of the wild carrot, Daucus carota, Am. J. Bot., 51, 274, 1964.

    CAS  Google Scholar 

  102. Lupotto, E., Propagation of an embryogenic culture of Medicago sativa L., Z. PflanzenphysioL, 111, 95, 1983.

    CAS  Google Scholar 

  103. Merkle, S.A. and Wiecko, A.T., Regeneration of Robinia psuedoacacia via somatic embryogenesis, Can. J. For. Res., 19, 285, 1989.

    Google Scholar 

  104. Tulecke, W., and McGranahan, G., Somatic embryogenesis and plant regeneration from cotyledons of Juglans regia L, Plant Sci. Lett., 1985.

    Google Scholar 

  105. Wetzstein, H.Y., Ault, J.R., and Merkle, S.A., Further characterization of somatic embryogenesis and plantlet regeneration in pecan (Carya illinoensis), Plant Sci., 64, 193, 1989.

    Google Scholar 

  106. Smith, D.L., and Krikorian, A.D., Growth and maintenance of an embryogenic cell culture of daylily (Hemerocallis) on hormone-free medium, Ann. Bot., 67, 443, 1991.

    PubMed  CAS  Google Scholar 

  107. Loh, C.-S., Ingram, D.S., and Hanke, D.E., Cytokinins and the regeneration of plantlets from secondary embryoids of winter oilseed rape, Brassica napus ssp. oleifera, New PhytoL, 95, 349, 1983.

    CAS  Google Scholar 

  108. Smith, D.L., and Krikorian, A.D., Somatic proembryo production from excised, wounded zygotic carrot embryos on hormone-free medium: evaluation of the effects of pH, ethylene and activated charcoal, Plant Cell Rep., 9, 34, 1990.

    PubMed  CAS  Google Scholar 

  109. Becwar, M.R., Noland, T.L., and Wann, S.R., A method for quanitification of the level of somatic embryogenesis among Norway spruce callus lines, Plant Cell Rep., 6, 35, 1987.

    Google Scholar 

  110. Parrott, W.A., and Bailey, M.A., Characterization of recurrent somatic embryogenesis of alfalfa on auxin-free medium and its control by sugars and nicotinic acid, Plant Cell Tiss. Org. Cult., 32, 69, 1993.

    CAS  Google Scholar 

  111. Bailey, M.A., Boerma, H.R., and Parrott, W.A., Genotype effects on repetitive embryogenesis and plant regeneration of soybean, In Vitro Cell. Dev. Biol., 29P, 102, 1993.

    Google Scholar 

  112. Halperin, W., and Jensen, W.A., Ultrastructural changes during growth and embryogenesis in carrot cell cultures, J. Ultrastructure Res., 18, 428, 1967.

    CAS  Google Scholar 

  113. Button, J., Kochba, J., and Bornman, C.H., Fine structure of and embryoid development from embryogenic ovular callus of “Shamouti” orange (Citrus sinensis Osb.), J. Exp. Bot., 25, 446, 1974.

    Google Scholar 

  114. Finer, J.J., and Nagasawa, A., Development of an embryogenic suspension culture of soybean (Glycine max Merrill.), Plant Cell Tiss. Org. Cult., 15, 125, 1988.

    CAS  Google Scholar 

  115. Lupotto, E., The use of single somatic embryo culture in propagating and regenerating lucerne (Medicago saliva L.), Ann. Bot., 57, 19, 1986.

    Google Scholar 

  116. Merkle, S.A., Somatic embryogenesis in black locust, in Black Locust: Biology, Culture and Utilization, Hanover, J.W., Miller, K., and Plesko, S., Eds., Michigan State University, East Lansing, 1992, 136.

    Google Scholar 

  117. Gupta, P.K., and Durzan, D.J., Biotechnology of somatic polyembryogenesis and plantlet regeneration on loblolly pine, Bio/Technol., 5, 147, 1987.

    Google Scholar 

  118. Durzan, D.J., and Gupta, P.K., Somatic embryogenesis and polyembryogenesis in Douglas-fir suspension cultures, Plant Sci., 52, 229, 1987.

    CAS  Google Scholar 

  119. Gupta, P.K., and Durzan, D.J., Somatic polyembryogenesis from callus of mature sugar pine embryos, Bio/Technol., 4, 643, 1986.

    Google Scholar 

  120. Becwar, M.R., Nagmani, R., and Wann, S.R., Initiation of embryogenic cultures and somatic embryo development in loblolly pine (Pinus taeda), Can. J. For. Res., 20, 810, 1990.

    Google Scholar 

  121. Durzan, D.J., and Gupta, P.K., Somatic embryogenesis and polyembryogenesis in conifers, in Biotechnology in Agriculture, Anonymous, Ed., Alan R. Liss, Inc, 1988, 53.

    Google Scholar 

  122. Attree, S.M., Moore, D., Sawhney, V.K., and Fowke, L.C., Enhanced maturation and desiccation tolerance of white spruce [Picea glauca (Moench) Voss] somatic embryos: Effects of a non-plasmolysing water stress and abscisic acid, Ann. Bot., 68, 519, 1991.

    Google Scholar 

  123. Gupta, P.K., Timmis, R., Pullman, G., Yancey, M., Kreitinger, M., Carlson, W., and Carpenter, C., Development of an embryogenic system for automated propagation of forest trees, in Scale-Up and Automation in Plant Propagation. Cell Culture and Somatic Cell Genetics, Vol. 8, Vasil, I.K., Ed., Academic Press, Inc., San Diego, 1991, 75.

    Google Scholar 

  124. Schuller, A., Reuther, G., and Geier, T., Somatic embryogenesis from seed explants of Abies alba, Plant Cell Tiss. Org. Cult., 17, 53, 1989.

    Google Scholar 

  125. Kessell, R.H.J., and Carr, A.H., The effect of dissolved oxygen concentration on growth and differentiation of carrot (Daucus carota) tissue, J. Exp. Bot., 23, 996, 1972.

    CAS  Google Scholar 

  126. Priel, W., Florek, P., Wix, U., and Beck, A., Towards mass propagation by use of bioreactors, Acta Hort., 226, 99, 1988.

    Google Scholar 

  127. Kvaalen, H., and Von Arnold, S., Effects of various partial pressures of oxygen and carbon dioxide on different stages of somatic embryogenesis in Picea abies, Plant Cell Tiss. Org. Cult., 27, 49, 1991.

    CAS  Google Scholar 

  128. Kumar, P.P., Joy, R.W. IV, and Thorpe, T.A., Ethylene and carbon dioxide accumulation, and growth of cell suspension cultures of Picea glauca (white spruce), J. Plant Physiol., 135, 592, 1989.

    Google Scholar 

  129. Vasil, I.K., Rationale for the scale-up and automation of plant propagation, in Scale-Up and Automation in Plant Propagation. Cell Culture and Somatic Cell Genetics of Plants, Volume 8, Vasil, I.K., Ed., Academic Press, Inc., San Diego, 1991, 1.

    Google Scholar 

  130. Denchev, P.D., Kuklin, A.E., and Scragg, A.H., Somatic embryo production in bioreactors, J. Biotech., 26, 99, 1992.

    CAS  Google Scholar 

  131. Backs-Hiisemann, D., and Reinert, J., Embryobildung durch isolierte Einzelzellen aus Gewebekulturen von Daucus carota, Protoplasma, 70, 49, 1970.

    Google Scholar 

  132. Ten Hoopen, H.J.G., Van Gulik, W.M., and Meijer, Possibilities, problems, and pitfalls of large-scale plant cell cultures, in Progress in Plant Cellular and Milecular Biology, Nijkamp, H.J.J., Van der Plas, L.W.H., and Van Aatrijk, J., Eds., Kluwer Academic Publishers, Dordrecht, 1990, 673.

    Google Scholar 

  133. Fowler, M.W., Process systems and approaches for large scale plant cell culture, in Plant Tissue and Cell Culture, Green, C.E., Somers, D.A., Hackett, W.P., and Biesboer, D.D., Eds., Alan R. Liss, Inc., New York, 1987, 459.

    Google Scholar 

  134. Fowler, M.W., Large-scale cultures of cells in suspension, in Cell Culture and Somatic Cell Genetics of Plants, Vol. 1, Vasil, I.K., Ed., Academic Press, New York, 1984, 167.

    Google Scholar 

  135. Stuart, D.A., Strickland, S.G., and Walker, K.A., Bioreactor production of alfalfa somatic embryos, HortSci., 22, 800, 1987.

    Google Scholar 

  136. Chen, T.H.H., Thompson, B.G., and Gerson, D.F., In vitro production of alfalfa somatic embryos in fermentation systems, J. Ferment. Technol., 65, 353, 1987.

    Google Scholar 

  137. Terashima, T., and Nishimura, S., Mass propagation of somatic embryos in carrot (Daucus carota) using bioreactor, Japan J. Breed., 41, 234, 1991.

    Google Scholar 

  138. Preil, W., Florek, P., and Wix, U., Towards mass propagation by use of bioreactors, Acta Hort., 226, 99, 1988.

    Google Scholar 

  139. Bapat, V.A., Fulzele, D.P., Heble, M.R., and Rao, P.S., Production of sandalwood somatic embryos in bioreactors, Curr. Sci., 59, 746, 1990.

    Google Scholar 

  140. Ammirato, P.V., and Styer, D.J., Strategies for large-scale manipulation of somatic embryos in suspension culture, in Biotechnology in Plant Science, Zaitlin, M., Day, P., and Hollaender, A., Eds., Academic Press, Inc., Orlando, 1985, 161.

    Google Scholar 

  141. Styer, D.J., Bioreactor technology for plant propagation, in Tissue Culture in Forestry and Agriculture, Henke, R.R., Hughes, K.W., Constantin, M.J., and Hollaender, A., Eds., Plenum Press, New York, 1985, 117.

    Google Scholar 

  142. Levin, R., Gaba, V., Tal., Hirsch, S., De Nola, D., and Vasil, I.K., Automated plant tissue culture for mass propagation, Bio/Technol., 6, 1035, 1988.

    Google Scholar 

  143. Levin, R., and Vasil, I.K., An integrated and automated tissue culture system for mass propagation of plants, In Vitro Cell. Dev. Biol., 25, 21, 1989.

    Google Scholar 

  144. Cazzulino, D.L., Pedersen, H., Chin, C.K., and Styer, D., Kinetics of carrot somatic embryo development in suspension culture, Biotech. Bioeng., 35, 781, 1990.

    CAS  Google Scholar 

  145. Cazzulino, D., Pedersen, H., and Chin, C.-K., Bioreactors and image analysis for scale-up and plant propagation, in Scale-Up and Automation in Plant Propagation. Cell Culture and Somatic Cell Genetics of Plants, Vol. 8, Vasil, I.K., Ed., Academic Press, Inc., San Diego, 1991, 147.

    Google Scholar 

  146. McKersie, B.D., Senaratna, T., Bowley, S.R., Brown, D.C.W., Krochko, J.E., and Bewley, J.D., Application of artificial seed technology in the production of hybrid alfalfa (Medicago sativa L.), In Vitro Cell. Dev. Biol., 25, 1183, 1989.

    Google Scholar 

  147. Senaratna, T., Mc Kersie, B.D., and Bowley, S.R., Desiccation tolerance of alfalfa (Medicago sativa L.) somatic embryos. Influence of abscisic acid, stress pretreatments and drying rates, Plant Sci., 65, 253, 1989.

    CAS  Google Scholar 

  148. Senaratna, T., Mc Kersie, B.D., and Bowley, S.R., Artificial seeds of alfalfa (Medicago sativa L.). Induction of desiccation tolerance in somatic embryos, In Vitro Cell. Dev. Biol., 26, 85, 1990.

    Google Scholar 

  149. Merkle, S.A., Wiecko, A.T., Sotak, R.J., and Sommer, H.E., Maturation and conversion of Liriodendron tulipifera somatic embryos, In Vitro Cell. Dev. Biol., 26, 1086, 1990.

    Google Scholar 

  150. Walker, C.C., Growth of embryogenic Norway spruce cultures in a bioreactor. A — 190 Independent Study. The Institute of Paper Science and Technology, Atlanta, GA, 1989.

    Google Scholar 

  151. Attree, S.M., Pomeroy, M.K., and Fowke, L.C., Production of vigorous, desiccation tolerant white spruce [Picea glauca (Moench) Voss.] synthetic seeds in a bioreactor, Plant Cell Rep., 13, 601, 1994.

    Google Scholar 

  152. LoSchiavo, F., Giuliano, G., De Vries, S.C., Genga, A., Bollini, R., Pitto, L., Cozzani, F., Nuti-Ronchi, V., and Terzi, M., A carrot cell variant temperature sensitive for somatic embryogenesis reveals a defect in the glycosylation of extracellular proteins, Mol. Gen. Genet., 223, 385, 1990.

    PubMed  Google Scholar 

  153. De Jong, A.J., Cordewener, J., Lo Schiavo, F., Terzi, M., Vandekerckhove, J., Van Kammen, A., and De Vries, S.C., A carrot somatic embryo mutant is rescued by chitinase, Plant Cell, 4, 425, 1992.

    PubMed  Google Scholar 

  154. De Jong, A.J., Heidstra, R., Spaink, H.P., Hartog, M.V., Meijer, E.A., Hendriks, T., Lo Schiavo, F., Terzi, M., Bisseling, T., Van Kammen, A., and De Vries, S.C., Rhizobium lipooligosaccharides rescue a carrot somatic embryo mutant, Plant Cell, 5, 615, 1993.

    PubMed  Google Scholar 

  155. Caligo, M.A., Nuti Ronchi, V., and Nozzolini, M., Proline and serine affect polarity and development of carrot somatic embryos, Cell Differentiation, 17, 193, 1985.

    CAS  Google Scholar 

  156. Fobert, P.R., and Webb, D.T., Effect of polyamines, polyamine precursors, and polyamine biosynthetic inhibitors on somatic embryogenesis from eggplant (Solanum melongena) cotyledons, Can. J. Bot., 66, 1734, 1988.

    CAS  Google Scholar 

  157. Conger, B.V., Hovanesian, S.C., Trigiano, R.N., and Gray, D.J., Somatic embryo ontogeny in suspension cultures of orchardgrass, Crop Sci., 29, 448, 1989.

    Google Scholar 

  158. Litz, R.E., Effect of osmotic stress on somatic embryogenesis in Carica suspension cultures, J. Amer. Soc. Hort. Sci., 111, 969, 1986.

    CAS  Google Scholar 

  159. Ammirato, P.V., The effects of abscisic acid on the development of somatic embryos from cells of caraway (Carum carvi L.), Bot. Gaz., 135, 328, 1974.

    CAS  Google Scholar 

  160. Ammirato, P.V., Hormonal control of somatic embryo development from cultured cells of caraway. Interactions of abscisic acid, zeatin, and gibberellic acid, Plant Physiol., 59, 579, 1977.

    PubMed  CAS  Google Scholar 

  161. Dusková, J., Opantrny, Z., Sovová, M., and Dusek, J., Somatic embryogenesis and plant regeneration in Coronilla varia L. (crownvetch) long-term tissue cultures, Biol. Plant, 32, 8, 1990.

    Google Scholar 

  162. Parrott, W.A., Dryden, G., Vogt, S., Hildebrand, D.F., Collins, G.B., and Williams, E.G., Optimization of somatic embryogenesis and embryo germination in soybean, In Vitro Cell. Dev. Biol., 24, 817, 1988.

    CAS  Google Scholar 

  163. Vasil, I.K., Somatic embryogenesis and its consequences in the gramineae, in Tissue Culture in Agriculture and Forestry, Vol. 31, Henke, R.R., Hughes, K.W., Constantin, M.J., and Hollaender, A., Eds., Plenum Press, New York, 1985, 31.

    Google Scholar 

  164. Kearney, J.F., Parrott, W.A., and Hill, N.S., Infection of somatic embryos of tall fescue with Acremonium coenophialum, Crop Sci., 31, 979, 1991.

    Google Scholar 

  165. Zaghmout, O.M.F., and Torello, W.A., Enhanced regeneration from long-term callus cultures of red fescue by pretreatment with activated charcoal, HortSci., 23, 615, 1988.

    Google Scholar 

  166. Roberts, D.R., Flinn, B.S., Webb, D.T., Webster, F.B., and Sutton, B.C.S., Abscisic acid and indole-3-butyric acid regulation of maturation and accumulation of storage proteins in somatic embryos of interior spruce, Physiol. Plant., 78, 355, 1990.

    CAS  Google Scholar 

  167. Michalczuk, L., Cooke, T.J., and Cohen, J.D., Auxin levels at different stages of carrot somatic embryogenesis, Phytochem., 31, 1097, 1992.

    CAS  Google Scholar 

  168. Gorst, J., Overall, R.L., and Wernicke, W., Ionic currents traversing cell clusters from carrot suspension cultures reveal perpetuation of morphogenetic potential as distinct from induction of embryogenesis, Cell Differentiation, 21, 101, 1987.

    PubMed  CAS  Google Scholar 

  169. Overall, R.L., and Wernicke, W., Steady ionic currents around haploid embryos formed from tobacco pollen in culture, in Ionic Currents in Development, Nuccitelli, R., Ed., Alan R. Liss, Inc., New York, 1986, 139.

    Google Scholar 

  170. Rathore, K.S., Hodges, T.K., and Robinson, K.R., Ionic basis of currents in somatic embryos of Daucus carota, Planta, 175, 280, 1988.

    Google Scholar 

  171. Schiavone, F.M., and Cooke, T.J., Unusual patterns of somatic embryogenesis in the domesticated carrot: developmental effects of exogenous auxins and auxin transport inhibitors, Cell Differentiation, 21, 53, 1987.

    PubMed  CAS  Google Scholar 

  172. Nomura, K., and Komamine, A., Molecular mechanisms of somatic embryogenesis, Oxford Surv. Plant Molec. Cell Biol., 3, 456, 1986.

    CAS  Google Scholar 

  173. Nomura, K., and Komamine, A., Physiological and biochemical aspects of somatic embryogenesis from single cells, in Somatic Embryogenesis, Terzi, M., Pitto, L., and Sung, Z.R., Eds., Consiglio Nationale delle Recherche, 1985, 1.

    Google Scholar 

  174. Timmers, A.C.J., De Vries, S.C., and Schel, J.H.N., Distribution of membrane-bound calcium and activated calmodulin during somatic embryogenesis of carrot (Daucus carota L.), Protoplasma, 153, 24, 1989.

    Google Scholar 

  175. Chée, R.P., and Cantliffe, D.J., Embryo development from discrete cell aggregates in Ipomoea batatas (L.) Lam. in response to structural polarity, In Vitro Cell. Dev. Biol., 25, 757, 1989.

    Google Scholar 

  176. Dijak, M., Smith, D.L., Wilson, T.J., and Brown, D.C.W., Stimulation of direct embryogenesis from mesophyll protoplasts of Medicago sativa, Plant Cell Rep., 5, 468, 1986.

    Google Scholar 

  177. Okada, K., Ueda, J., Komaki, M.K., Bell, C.J., and Shimura, Y., Requirement of the auxin polar transport system in early stages of Arabidopsis floral bud formation, Plant Cell, 3, 677, 1992.

    Google Scholar 

  178. Liu, C.-m., Xu, Z.-H., and Chua, N.-H., Auxin polar transport is essential for the establishment of bilateral symmetry during early plant embryogenesis, Plant Cell, 5, 621, 1993.

    PubMed  CAS  Google Scholar 

  179. Ho, W.-J., and Vasil, I.K., Somatic embryogenesis in sugarcane (Saccharum officinarum L.) I. The morphology and physiology of callus formation and the ontogeny of somatic embryos, Protoplasma, 18, 169, 1983.

    Google Scholar 

  180. Becwar, M.R., Noland, T.L., and Wyckoff, J.L., Maturation, germination and conversion of Norway spruce (Picea abies L.) somatic embryos to plants, In Vitro Cell. Dev. Biol., 25, 575, 1989.

    Google Scholar 

  181. Buchheim, J.A., Colburn, S.M., and Ranch, J.P., Maturation of soybean somatic embryos and the transition to plantlet growth, Plant Physiol., 89, 768, 1989.

    PubMed  CAS  Google Scholar 

  182. Ebert, A., and Taylor, H.F., Assessment of the changes of 2,4-dichlorophenoxyacetic acid concentrations in plant tissue culture media in the presence of activated charcoal, Plant Cell Tiss. Org. Cult., 20, 165, 1990.

    CAS  Google Scholar 

  183. Aung, L.H., Buss, G.R., Crosby, K.E., and Brown, S.S., Changes in the hormonal levels of soybean fruit during ontogeny, Øyton, 42, 151, 1982.

    CAS  Google Scholar 

  184. Danin, M., Upfold, Si., Levin, N., Nadel, B.L., and Van Standen, J., Polyamines and cytokinins in celery embryogenic cell cultures, Plant Growth Regulation, 12, 245, 1993.

    CAS  Google Scholar 

  185. Wright, M.S., Launis, K.L., Novitzky, R., Duesing, J.H., and Harms, C.T., A simple method for the recovery of multiple fertile plants from individual somatic embryos of soybean [Glycine max (L.) Merrill], In Vitro Cell. Dev. Biol., 27P, 153, 1991.

    Google Scholar 

  186. Bewley, J.D., and Black, M., Seeds: germination, structure and composition, in Physiology of Development and Germination, Bewley, J.D., and Black, M., Eds., Plenum Press, New York, 1985, 1.

    Google Scholar 

  187. Lai, F., Senaratna, T., and Mc Kersie, B.D., Glutamine enhances storage protein synthesis in Medicago sativa L. somatic embryos, Plant Sci., 87, 69, 1992.

    CAS  Google Scholar 

  188. Verma, D.C., and Dougall, D.K., Influence of carbohydrates on quantitative aspects of growth and embryo formation in wild carrot suspension cultures, Plant Physiol., 59, 81, 1977.

    PubMed  CAS  Google Scholar 

  189. Schuller, A., and Reuther, G., Response of Abies alba embryonal-suspensor mass to various carbohydrate treatments, Plant Cell Rep., 12, 199, 1993.

    CAS  Google Scholar 

  190. Finer, J.J., and Mc Mullen, M.D., Transformation of soybean via particle bombardment of embryogenic suspension culture tissue, In Vitro Cell. Dev. Biol., 27P, 175, 1991.

    CAS  Google Scholar 

  191. Denchov, P., Velcheva, M., and Atanassov, A., A new approach to direct somatic embryogenesis in Medicago, Plant Cell Rep. 10, 338, 1991.

    Google Scholar 

  192. Kielly, G.A., and Bowley, S.R., Genetic control of somatic embryogenesis in alfalfa, Genome 35, 474, 1992.

    Google Scholar 

  193. Tremblay, L., and Tremblay, F.M., Carbohydrate requirements for the development of black spruce (Picea mariana (Mill.) B.S.P.) and red spruce (P. rubens Sarg.) somatic embryos, Plant Cell Tiss. Org. Cult. 27, 95, 1991.

    CAS  Google Scholar 

  194. Anandarajah, K., and Mc Kersie, B.D., Enhanced vigor of dry somatic embryos of Medicago sativa L. with increased sucrose, Plant Sci. 71, 261, 1990.

    CAS  Google Scholar 

  195. Lu, C.-Y., and Thorpe, T.A., Somatic embryogenesis and plantlet regeneration in cultured immature embryos of Picea glauca, J. Plant PhysioL 128, 297, 1987.

    CAS  Google Scholar 

  196. Roberts, D.R., Abscisic acid and mannitol promote early development, maturation and storage protein accumulation in somatic embryos of interior spruce, Physiol. Plant. 83, 247, 1991.

    CAS  Google Scholar 

  197. Jain, S.M., Newton, R.J., and Soltes, E.J., Enhancement of somatic embryogenesis in Norway spruce (Picea abies L.), Theor. Appl. Genet. 76, 501, 1988.

    Google Scholar 

  198. Earnshaw, B.A., and Johnson, M.A., Control of wild carrot somatic embryo development by antioxidants, Plant Physiol. 85, 273, 1987.

    PubMed  CAS  Google Scholar 

  199. Kepczynski, J., Mc Kersie, B.D., and Brown, D.C.W., Requirement of ethylene for growth of callus and somatic embryogenesis in Medicago saliva L, J. Exp. Bot. 43, 1199, 1992.

    CAS  Google Scholar 

  200. Fujii, J.A.A., Slade, D., Olsen, R., Ruzin, S.E., and Redenbaugh, K., Alfalfa somatic embryo maturation and conversion to plants, Plant Sci. 72, 93, 1990.

    CAS  Google Scholar 

  201. Crouch, M.L., Non-zygotic embryos of Brassica napus L. contain embryo-specific storage proteins, Planta 156, 520, 1982.

    CAS  Google Scholar 

  202. Flinn, B.S., Roberts, D.R., and Taylor, I.E.P., Evaluation of somatic embryos of interior spruce. Characterization and developmental regulation of storage proteins, Physiol. Plant. 82, 624, 1991.

    CAS  Google Scholar 

  203. Krochko, J.E., Pramanik, S.K., and Bewley, J.D., Contrasting storage protein synthesis and messenger RNA accumulation during development of zygotic and somatic embryos of alfalfa (Medicago sativa L.), Plant Physiol. 99, 46, 1992.

    PubMed  CAS  Google Scholar 

  204. Shoemaker, R.C., Christofferson, S.E., and Galbraith, D.W., Storage protein accumulation patterns in somatic embryos of cotton (Gossypium hirsutum L.), Plant Cell Rep. 6, 12, 1987.

    CAS  Google Scholar 

  205. Stuart, D.A., Nelsen, J., and Nichol, J.W., Expression of 7S and 11S alfalfa seed storage proteins in somatic embryos, J. Plant Physiol. 132, 134, 1988.

    CAS  Google Scholar 

  206. Misra, S., Attree, S.M., Leal, I., and Fowke, L.C., Effect of abscisic acid, osmoticum, and desiccation on synthesis of storage proteins during the development of white spruce somatic embryos, Ann. Bot., 71, 11, 1993.

    CAS  Google Scholar 

  207. Tewes, A., Manteuffel, R., Adler, K., Weber, E., and Wobus, U., Long-term cultures of barley synthesize and correctly deposit seed storage proteins, Plant Cell Rep. 10, 467, 1991.

    CAS  Google Scholar 

  208. Taylor, D.C., Weber, N., Underhill, E.W., Pomeroy, M.K., Keller, W.A., Scowcroft, W.R., Wilen, R.W., Moloney, M.M., and Holbrook, L.A., Storage protein regulation and lipid accumulation in microspore embryos of Brassica napus L, Planta 181, 18, 1990.

    CAS  Google Scholar 

  209. Dahmer, M.L., Hildebrand, D.F., and Collins, G.B., Comparative protein accumulation patterns in soybean somatic and zygotic embryos, In Vitro Cell. Dev. Biol. 28P, 106, 1992.

    CAS  Google Scholar 

  210. Joy, R.W. IV, Yeung, E.C., Kong, L., and Thorpe, T.A., Development of white spruce somatic embryos. I. Storage product deposition, In Vitro Cell. Dev. Biol. 27, 32, 1991.

    Google Scholar 

  211. Wilen, R.W., Mandel, R.M., Pharis, R.P., Holbrook, L.A., and Moloney, M.M., Effects of abscisic acid and high osmoticum on storage protein gene expression in microspore embryos of Brassica napus, Plant Physiol. 94, 875, 1990.

    CAS  Google Scholar 

  212. Pramanik, S.K., Krochko, J.E., and Bewley, J.D., Distribution of cytosolic mRNAs between polysomal and ribonucleoprotein complex fractions in alfalfa embryos, Plant Physiol. 99, 1590, 1992.

    PubMed  CAS  Google Scholar 

  213. Bray, E., and Beachy, R.N., Regulation by ABA of β-conglycinin expression in cultured developing soybean cotyledons, Plant Physiol. 79, 746, 1985.

    PubMed  CAS  Google Scholar 

  214. Finkelstein, R.R., Tenbarge, K.M., Shumway, J.E., and Crouch, M.L., Role of ABA in maturation of rapeseed embryos, Plant Physiol. 78, 630, 1985.

    PubMed  CAS  Google Scholar 

  215. Cyr, D.R., Webster, F.B., and Roberts, D.R., Biochemical events during germination and early growth of somatic embryos and seed of interior spruce (Picea glauca engelmanii complex), Seed Sci. Res. 1, 91, 1991.

    CAS  Google Scholar 

  216. Hakman, I., Stabel, P., Engstrom, P., and Eriksson, T., Storage protein accumulation during zygotic and somatic embryo development in Picea abies (Norway spruce), Physiol. Plant. 80, 441, 1990.

    CAS  Google Scholar 

  217. Flinn, B.S., Roberts, D.R., Newton, C.H., Cyr, D.R., Webster, F.B., and Taylor, I.E.P., Storage protein gene expression in zygotic and somatic embryos of interior spruce, Physiol. Plant. 1993 (in press).

    Google Scholar 

  218. Avjioglu, A., and Knox, R.B., Storage lipid accumulation by zygotic and somatic embryos in culture, Ann. Bot. 63, 409, 1989.

    CAS  Google Scholar 

  219. Hara, S., Falk, H., and Kleinig, H., Starch and triacylglycerol metabolism related to somatic embryogenesis in Papaver orientale tissue cultures, Planta 164, 303, 1985.

    CAS  Google Scholar 

  220. Pomeroy, M.K., Kramer, J.K.G., Hunt, D.J., and Keller, W.A., Fatty acid changes during development of zygotic and microspore-derived embryos of Brassica napus, Physiol. Plant. 81, 447, 1991.

    CAS  Google Scholar 

  221. Warren, G.S., and Fowler, M.W., Changing fatty acid composition during somatic embryogenesis in cultures of Daucus carota, Planta 144, 451, 1979.

    CAS  Google Scholar 

  222. Turnham, E., and Northcote, D.H., The incorporation of [1-14C] acetate into lipids during embryogenesis in oil palm tissue cultures, Phytochem. 23, 35, 1984.

    CAS  Google Scholar 

  223. Attree, S.M., Pomeroy, M.K., and Fowke, L.C., Manipulation of conditions for the culture of somatic embryos of white spruce for improved triacylglycerol biosynthesis and desiccation tolerance, Planta 187, 395, 1992.

    CAS  Google Scholar 

  224. Hildebrand, D.F., Adams, T.R., Dahmer, M.L., Williams, E.G., and Collins, G.B., Analysis of lipid composition and morphological characteristics in soybean regenerants, Plant Cell Rep. 7, 701, 1989.

    CAS  Google Scholar 

  225. Shoemaker, R.C., and Hammond, E.G., Fatty acid composition of soybean (Glycine max (L.) Merr.) somatic embryos, In Vitro Cell. Dev. Biol. 24, 829, 1988.

    CAS  Google Scholar 

  226. Kim, Y.-H., and Janick, J., Abscisic acid and proline improved desiccation tolerance and increase fatty acid content of celery somatic embryos, Plant Cell Tiss. Org. Cult. 24, 83, 1991.

    CAS  Google Scholar 

  227. Stuart, D.A., and Mc Call, C.M., Induction of somatic embryogenesis using side chain and ring modified forms of phenoxy acid growth regulators, Plant Physiol. 99, 111, 1992.

    PubMed  CAS  Google Scholar 

  228. Feirer, R.P., Conkey, J.H., and Verhagen, S.A., Triglycerides in embryogenic conifer calli: a comparison with zygotic embryos, Plant Cell Rep. 8, 207, 1989.

    CAS  Google Scholar 

  229. Dahmer, M.L., Collins, G.B., and Hildebrand, D.F., Lipid content and composition of soybean somatic embryos, Crop Sci. 31, 741, 1991.

    CAS  Google Scholar 

  230. Finkelstein, R., and Somerville, C., Abscisic acid or high osmoticum promote accumulation of long-chain fatty acids in developing embryos of Brassica napus, Plant Sci. 61, 213, 1989.

    CAS  Google Scholar 

  231. Roberts, D.R., Sutton, B.C.S., and Flinn, B.S., Synchronous and high frequency germination of interior spruce somatic embryos following partial drying at high relative humidity, Can. J. Bot. 68, 1086, 1990.

    Google Scholar 

  232. Attree, S.M., Tautorus, T.E., Dunstan, D.I., and Fowke, L.C., Somatic embryo maturation, germination and soil establishment of plants of black and white spruce (Picea mariana and Picea glauca), Can. J. Bot. 68, 2583, 1990.

    Google Scholar 

  233. Gray, D.J., and Purohit, A., Somatic embryogenesis and development of synthetic seed technology, Crit. Rev. Pl. Sci. 10, 33, 1991.

    Google Scholar 

  234. Kott, L.S., and Beversdorf, W.D., Enhanced plant regeneration from microspore-derived embryos of Brassica napus by chilling, partial desiccation and age selection, Plant Cell Tiss. Org. Cult. 23, 187, 1990.

    Google Scholar 

  235. Kott, L.S., and Kasha, K.J., Initiation and morphological development of somatic embryoids from barley cell cultures, Can. J. Bot. 62, 1245, 1984.

    Google Scholar 

  236. Finkelstein, R.R., and Crouch, M.L., Precociously germinating rapeseed embryos retain characteristics of embryogeny, Planta 162, 125, 1984.

    CAS  Google Scholar 

  237. Gray, D.J., Conger, B.V., and Songstad, D.D., Desiccated quiescent somatic embryos of orchardgrass for use as synthetic seeds, In Vitro Cell. Dev. Biol. 23, 29, 1987.

    Google Scholar 

  238. Gray, D.J., Quiescence in monocotyledonous and dicotyledonous somatic embryos induced by dehydration, HortSci. 22, 810, 1987.

    Google Scholar 

  239. Hammatt, N., and Davey, M.R., Somatic embryogenesis and plant regeneration from cultured zygotic embryos of soybean (Glycine max L. Merr.), J. Plant Physiol. 128, 219, 1987.

    Google Scholar 

  240. Roberts, D.R., Lazaroff, W.R., and Webster, F.B., Interaction between maturation and high relative humidity treatments and their effects on germination of sitka spruce somatic embryos, J. Plant Physiol. 138, 1, 1991.

    Google Scholar 

  241. Anandarajah, K., and Mc Kersie, B.D., Manipulating the desiccation tolerance and vigor of dry somatic embryos of Medicago sativa L. with sucrose, heat shock and abscisic acid, Plant Cell Rep. 9, 451, 1990.

    CAS  Google Scholar 

  242. Anandarajah, K., Kott, L., Beversdorf, W.D., and Mc Kersie, B.D., Induction of desiccation tolerance in microspore-derived embryos of Brassica napus L. by thermal stress, Plant Sci. 77, 119, 1991.

    Google Scholar 

  243. Parrott, W.A., Auxin-stimulated somatic embryogenesis from immature cotyledons of white clover, Plant Cell Rep., 10 17, 1991.

    CAS  Google Scholar 

  244. Webster, F.B., Roberts, D.R., Mc Innis, S.M., and Sutton, B.C.S., Propagation of interior spruce by somatic embryogenesis, Can. J. For. Res. 20, 1759, 1990.

    Google Scholar 

  245. Weissinger, A.K. II, and Parrott, W.A., Repetitive somatic embryogenesis and plant recovery in white clover, Plant Cell Rep. 12, 125, 1993.

    Google Scholar 

  246. Roberts, D.R., Webster, F.B., Flinn, B.S., Lazaroff, W.R., and Cyr, D.R., Somatic embryogenesis of spruce, in Synseeds. Applications of Synthetic Seeds to Crop Improvement Redenbaugh, K., Ed., CRC Press, Inc., Boca Raton, FL, 1993, 427.

    Google Scholar 

  247. Wake, H., Akasaka, A., Umetsu, H., Ozeki, Y., Shimomura, K., and Matsunaga, T., Enhanced germination of artificial seeds by marine cyanobacterial extract, Appl. Microbiol. BiotechnoL 36, 684, 1992.

    Google Scholar 

  248. Von Arnold, S., and Hakman, I., Regulation of somatic embryo development in Picea abies by abscisic acid (ABA), J. Plant Physiol. 132, 164, 1988.

    Google Scholar 

  249. Wetzstein, H.Y., and Sommer, H.E., Scanning electron microscopy of in vitro cultured Liquidambar styraciflua plantlets during acclimatization, J. Amer. Soc. Hort. Sci. 108, 475, 1983.

    Google Scholar 

  250. Komatsuda, T., and Ohyama, K., Genotypes of high competence for somatic embryogenesis and plant regeneration in soybean Glycine max, Theor. Appl. Genet. 75, 695, 1988.

    Google Scholar 

  251. Redenbaugh, K., SynSeeds: Applications of Synthetic Seeds to Crop Improvement, CRC Press, Boca Raton, FL, 1993.

    Google Scholar 

  252. Redenbaugh, K., Fujii, J.A., and Slade, D., Synthetic seed technology, in Scale-Up and Automation in Plant Propagation. Cell Culture and Somatic Cell Genetics of Plants, Vol. 8, Vasil, I.K., Ed., Academic Press, Inc., San Diego, 1991, 35.

    Google Scholar 

  253. Murashige, T., Plant cell and organ cultures as horticultural practices, Acta Hort., 78, 17, 1977.

    Google Scholar 

  254. Kitto, S., and Janick, J., Polyox as an artificial seed coat for asexual embryos, HortSci. 17, 448, 1982.

    Google Scholar 

  255. Redenbaugh, K., Nichol, J., Kossler, M.E., and Paasch, B., Encapsulation of somatic embryos for artificial seed production, In Vitro Cell. Dev. Biol. 20, 256, 1984.

    Google Scholar 

  256. Redenbaugh, K., Paasch, B.D., Nichol, J.W., Kossler, M.E., Viss, P.R., and Walker, K.A., Somatic seeds: Encapsulation of asexual plant embryos, Bio/Tech. 4, 797, 1986.

    Google Scholar 

  257. Redenbaugh, K., Slade, D., Viss, P., and Fujii, J.A., Encapsulation of somatic embryos in synthetic seed coats, HortSci. 22, 803, 1987.

    Google Scholar 

  258. Gray, D.J., Effects of dehydration and exogenous growth regulators on dormancy, quiescence and germination of grape somatic embryos, In Vitro Cell. Dev. Biol. 25, 1173, 1989.

    Google Scholar 

  259. Kitto, S.L., and Janick, J., Hardening treatments increase survival of synthetically-coated asexual embryos of carrot, J. Amer. Soc. Hort. Sci. 110, 283, 1985.

    CAS  Google Scholar 

  260. Kitto, S.L., and Janick, J., Production of synthetic seeds by encapsulating asexual embryos of carrot, J. Amer. Soc. Hort. Sci. 110, 277, 1985.

    Google Scholar 

  261. Kim, Y.-H., and Janick, J., ABA and Polyox-encapsulation or high humidity increases survival of desiccated somatic embryos of celery, HortSci. 24, 674, 1989.

    CAS  Google Scholar 

  262. Janick, J., Kitto, S.L., and Kim, Y.-H., Production of synthetic seed by desiccation and encapsulation, In Vitro Cell. Dev. Biol. 25, 1167, 1989.

    Google Scholar 

  263. Fujii, J.A., Slade, D., Aguirre-Rascon, J., and Redenbaugh, K., Field planting of alfalfa artificial seeds, In Vitro Cell. Dev. Biol. 28P, 73, 1992.

    Google Scholar 

  264. Fujii, J.A.A., Slade, D., and Redenbaugh, K., Maturation and greenhouse planting of alfalfa artificial seeds, In Vitro Cell. Dev. BioL 25, 1179, 1989.

    Google Scholar 

  265. Datta, S., and Potrykus, I., Artificial seeds of barley: encapsulation of microspore-derived embryos, Theor. Appl. Genet. 77, 820, 1989.

    Google Scholar 

  266. Shigeta, J., Mori, T., Toda, K., and Ohtake, H., Effects of capsule hardness on germination frequency of encapsulated somatic embryos of carrot, Biotechnol. Tech. 4, 21, 1990.

    Google Scholar 

  267. Bapat, V.A., and Rao, P.S., Sandalwood plantlets from “synthetic seeds”, Plant Cell Rep. 7, 434, 1988.

    Google Scholar 

  268. Bapat, V.A., and Rao, P.S., Somatic seeds of sandalwood (Santalum album L.) and mulberry (Morus indica L.), in Proceedings of the National Seminar on Advances in Seed Science and Technology Shetty, H.S., and Prakash, H.S., Eds., Univ. of Mysore, Mysore, 1989, 372.

    Google Scholar 

  269. Bapat, V.A., and Rao, P.S., Plantlet regeneration from encapsulated and nonencapsulated desiccated somatic embryos of a forest tree: sandalwood (Santalum album L.), J. Plant Biochem. Biotechnol. 1, 109, 1992.

    CAS  Google Scholar 

  270. Lulsdorf, M.M., Tautorus, T.E., Kikcio, S.I., Bethune, T.D., and Dunstan, D.I., Germination of encapsulated embryos of interior spruce (Picea glauca engelmannii complex) and black spruce (Picea mariana Mill.), Plant Cell Rep. 12, 385, 1993.

    Google Scholar 

  271. Rao, P.V.L., and Singh, B., Plantlet regeneration from encapsulated somatic embryos of hybrid Solanum melongena L, Plant Cell Rep. 10, 7, 1991.

    CAS  Google Scholar 

  272. Shigeta, J., Mori, T., and Sato, K., Storage of encapsulated somatic embryos of carrot, Biotechnol. Tech. 7, 165, 1993.

    Google Scholar 

  273. Currah, I.E., Gray, D., and Thomas, T.H., The sowing of germinating vegetable seeds using a fluid drill, Ann. Appl. Biol. 76, 311, 1974.

    Google Scholar 

  274. Baker, C.M., Synchronization and fluid sowing of carrot, Daucus carota somatic embryos. M.Sc. Thesis, University of Florida, Gainesville, 1985.

    Google Scholar 

  275. Schultheis, J.R., and Cantliffe, D.J., Plant formation of Ipomea batatas Poir. from somatic embryos in gel carriers with additives, HortSci. 23, 812, 1988.

    Google Scholar 

  276. Mc Elroy, A.R., and Brown, D.C.W., A transplant plug technique for production of alfalfa (Medicago saliva L.) plants from somatic embryos, Can. J. Plant Sci. 72, 483, 1992.

    Google Scholar 

  277. Barwale, U.B., and Widholm, J.M., Somaclonal variation in plants regenerated from cultures of soybean, Plant Cell Rep. 6, 365, 1987.

    Google Scholar 

  278. Armstrong, C.L., and Phillips, R.L., Genetic and cytogenetic variation in plants regenerated from organogenic and friable, embryogenic tissue cultures of maize, Crop Sci. 28, 363, 1988.

    Google Scholar 

  279. Caligari, P.D.S., and Shohet, S., Variability in somatic embryos, in Synseeds: Applications of Synthetic Seeds to Crop Improvement, Redenbaugh, K., Ed., CRC Press, Boca Raton, FL, 1993, 163.

    Google Scholar 

  280. Nuti Ronchi, V., Giorgetti, L., Tonelli, M., and Martini, G., Ploidy reduction and genome segregation in cultured carrot cell lines. I. Prophase chromosome reduction, Plant Cell Tiss. Org. Cult. 30, 107, 1992.

    Google Scholar 

  281. Rugh, C.L., Parrott, W.A., and Merkle, S.A., Ploidy variation in embryogenic yellow-poplar, in Proc. 22nd Southern Forest Tree Improvement Conference, Atlanta, GA, June 14–17, 1993, 493.

    Google Scholar 

  282. Kott, L.S., Flack, S., and Kasha, K.J., A comparative study of initiation and development of embryogenic callus from haploid embryos of several barley cultivars. II. Cytophotometry of embryos and callus, Can. J. Bot. 64, 2107, 1986.

    Google Scholar 

  283. Shoemaker, R.C., Amberger, L.A., Palmer, R.G., Oglesby, L., and Ranch, J.P., Effect of 2,4- dichlorophenoxyacetic acid concentration on somatic embryogenesis and heritable variation in soybean [Glycine max (L) Merr.], In Vitro Cell. Dev. Biol. 27P, 84, 1991.

    CAS  Google Scholar 

  284. Durand-Gasselin, T., Le Guen, V., Konan, K., and Duval, Y., Plantations en Côte-d’Ivoire de palmiers à huile (Elaeis guineensis Jacq.), obtenues par culture in vitro. Premiers résultats, Oleagineux 45, 1, 1990.

    Google Scholar 

  285. Le Guen, V., Samaritaan, G., Othman, A.Z., Chin, C.W., Konan, K., and Durand-Gasselin, T., Oil production in young oil palm clones, Oléagineux 46, 347, 1991.

    Google Scholar 

  286. Corley, R.H.V., Lee, C.H., Law, L.H., and Wong, C.Y., Abnormal flower development in oil palm clones, Planter 62, 233, 1986.

    Google Scholar 

  287. McKersie, B.D., and Bowley, S.R., Synthetic seeds of alfalfa, in Synseeds. Applications of Synthetic Seeds to Crop Improvement, Redenbaugh, K., Ed., CRC Press, Boca Raton, FL, 1993, 231.

    Google Scholar 

Download references

Authors
  1. S. A. Merkle
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. W. A. Parrott
    View author publications

    You can also search for this author in PubMed Google Scholar

  3. B. S. Flinn
    View author publications

    You can also search for this author in PubMed Google Scholar

Editor information

Editors and Affiliations

  1. Plant Physiology Research Group, Department of Biological Sciences, The University of Calgary, Calgary, Alberta, Canada

    Trevor A. Thorpe

Rights and permissions

Reprints and permissions

Copyright information

© 1995 Springer Science+Business Media Dordrecht

About this chapter

Cite this chapter

Merkle, S.A., Parrott, W.A., Flinn, B.S. (1995). Morphogenic Aspects of Somatic Embryogenesis. In: Thorpe, T.A. (eds) In Vitro Embryogenesis in Plants. Current Plant Science and Biotechnology in Agriculture, vol 20. Springer, Dordrecht. https://doi.org/10.1007/978-94-011-0485-2_5

Download citation

  • DOI: https://doi.org/10.1007/978-94-011-0485-2_5

  • Publisher Name: Springer, Dordrecht

  • Print ISBN: 978-94-010-4217-8

  • Online ISBN: 978-94-011-0485-2

  • eBook Packages: Springer Book Archive

Publish with us

Policies and ethics

Search

Navigation