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Underdeveloped embryos and kinds of dormancy in seeds of two gymnosperms: Podocarpus costalis and Nageia nagi (Podocarpaceae)

  • Shun-Ying Chen (a1), Carol C. Baskin (a2) (a3), Jerry M. Baskin (a2) and Ching-Te Chien (a1)

Although it has been speculated that seeds of the gymnosperm family Podocarpaceae have an underdeveloped embryo, no detailed studies have been done to definitively answer this question. Our purpose was to determine if embryos in seeds of two species of Podocarpaceae, Podocarpus costalis and Nageia nagi, from Taiwan are underdeveloped and to examine the kind of dormancy the seeds have. Embryos in fresh seeds of P. costalis were 4.6 ± 0.5 mm long, and they increased in length by about 54% before radicle emergence (germination), demonstrating that the embryo is underdeveloped at seed maturity. Seeds germinated to >90% at 30/20, 25/15 and 25°C in light in ≤ 4 weeks, without any cold stratification pretreatment. Thus, seeds of P. costalis have morphological dormancy (MD). Embryos in fresh seeds of N. nagi were 7.4 ± 0.8 mm long and they increased in length by about 39% before radicle emergence (germination) occurred, indicating that the embryo is underdeveloped at seed maturity. Seeds germinated to < 25% at 30/20 and 25°C in light in 4 weeks but to >90% at the same temperatures in 12 weeks. Thus, most seeds of N. nagi have morphophysiological dormancy (MPD). Although underdeveloped embryos are considered to be a primitive condition in seed plants, they also occur in the most advanced orders. The occurrence of underdeveloped embryos in Podocarpaceae documents that they are not restricted to a basal clade in gymnosperms.

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Baskin, C.C. and Baskin, J.M. (1998) Seeds: ecology, biogeography, and evolution of dormancy and germination. San Diego, Academic Press.
Baskin, C.C. and Baskin, J.M. (2005) Underdeveloped embryos in dwarf seeds and implications for assignment to dormancy class. Seed Science Research 15, 357360.
Baskin, C.C. and Baskin, J.M. (2007) A revision of Martin's seed classification system, with particular reference to his dwarf-seed type. Seed Science Research 17, 1120.
Baskin, C.C., Chien, C.T., Chen, S.Y. and Baskin, J.M. (2008) Germination of Viburnum odoratissimum seeds: a new level of morphophysiological dormancy. Seed Science Research 18, 179184.
Baskin, J.M. and Baskin, C.C. (2004) A classification system for seed dormancy. Seed Science Research 14, 116.
Baskin, J.M. and Baskin, C.C. (2008) Some considerations for adoption of Nikolaeva's formula system into seed dormancy classification. Seed Science Research 18, 131137.
Baskin, J.M., Baskin, C.C., Chien, C.T. and Chen, S.Y. (2006a) Seed dormancy in the early diverging eudicot Trochodendron aralioides (Trochodendraceae). Seed Science Research 16, 7175.
Baskin, J.M., Hidayati, S.N., Baskin, C.C., Walck, J.L., Huang, Z.Y. and Chien, C.T. (2006b) Evolutionary considerations of presence of both morphophysiological and physiological seed dormancy in the highly-advanced euasterids II order Dipsacales. Seed Science Research 16, 233242.
Cheng, G., Tang, X., Gao, H. and Shen, S. (2004) Dormancy mechanism and relieving techniques of seeds of Taxus cuspidata Sieb. et Zucc. Journal – Beijing Forestry University 26, 59(in Chinese with English abstract).
Chien, C.T., Kuo-Huang, L.L. and Lin, T.P. (1998) Changes in ultrastructure and abscisic acid level, and response to applied gibberellins in Taxus mairei seeds treated with warm and cold stratification. Annals of Botany 81, 4147.
Chien, C.T., Chen, S.Y., Chang, S.H. and Chung, J.D. (2012) Seed germination and storage of Cycas taitungensis (Cycadaceae). Taiwan Journal of Forest Science 27, 111.
Del Tredici, P. (2007) The phenology of sexual reproduction in Ginkgo biloba: ecological and evolutionary implications. The Botanical Review 73, 267278.
Devillez, F. (1978) Influence de la température sur la postmaturation et la germination des graines de l'if (Taxus baccata L.). Bulletin de la Classe des Sciences l'Academie Royale de Belgique 64, 203218.
Dodd, M.C. and Van Staden, J. (1981) Germination and viability studies on the seeds of Podocarpus henkelii Stapf. South African Journal of Science 77, 171174.
Ferrandis, P., Bonilla, M. and Osorio, L.C. (2011) Germination and soil seed bank traits of Podocarpus angustifolius (Podocarpaceae): an endemic tree species from Cuban rain forests. International Journal of Tropical Biology 59, 10611069.
Figueroa, J.A. (2003) Seed germination in temperate rain forest species of southern Chile: chilling and gap-dependency germination. Plant Ecology 166, 227240.
Figueroa, J.A. and Lusk, C.H. (2001) Germination requirements and seedling shade tolerance are not correlated in a Chilean temperate rain forest. The New Phytologist 152, 483489.
Finch-Savage, W.E. and Leubner-Metzger, G. (2006) Seed dormancy and the control of germination. The New Phytologist 171, 501523.
Forbis, T.A., Floyd, S.K. and de Querioz, A. (2002) The evolution of embryo size in angiosperms and other seed plants: implications for the evolution of seed dormancy. Evolution 56, 21122125.
Grushvitzky, I.V. (1967) After-ripening of seeds of primitive tribes of angiosperms, conditions and peculiarities. pp. 329336+8 figuresin Borris, H. (Ed.) Physiologie, ökologie und biochemie der keimung. Greifswald, Germany, Ernst-Moritz-Arndt Universität.
Hong, K.Y., Jhuang, P.H., Chien, C.T., Huang, Y.J. and Fan, Y.B. (2009) Germination and storage of seeds of the endangered plant, Nageia nagi. Scientific Agriculture 57, 170175(in Chinese with English abstract).
Hong, T.D. and Ellis, R.H. (1996) A protocol to determine seed storage behaviour. Rome, International Plant Genetic Resources Institute.
International Seed Testing Association (2007) ISTA handbook on moisture determination (1st edition). Nijënstein, H.; Nydam, J.; Don, R.; McGill, C. (Eds) Bassersdorf, Switzerland, International Seed Testing Association.
Li, H.L., Keng, H., Yang, Y.P. and Lu, S.Y. (1994) Podocarpaceae. pp. 557566in Editorial Committee of the Flora of Taiwan (Ed.) Flora of Taiwan, vol. 1, (2nd edition). Taiwan, Editorial Committee of the Flora of Taiwan.
Linkies, A., Graeber, K., Knight, C. and Leubner-Metzger, G. (2010) The evolution of seeds. The New Phytologist 186, 817831.
Liu, D., Yu, H.L. and Guo, H.H. (2011) An analysis of dormancy and dormancy release in Taxus chinensis var. mairei seeds. Seed Science and Technology 39, 2943.
Martin, A.C. (1946) The comparative internal morphology of seeds. The American Midland Naturalist 36, 513660.
Meyen, S.V. (1984) Basic features of gymnosperm systematics and phylogeny as evidenced by the fossil record. The Botanical Review 50, 1111.
Naylor, R.E.L. (1981) An evaluation of various germination indices for predicting differences in seed vigour in Italian ryegrass. Seed Science and Technology 9, 593600.
Negash, L. (1992) In vitro methods for the rapid germination of seeds of Podocarpus falcatus. Ethiopian Journal of Science 15, 8597.
Ng, F.S.P. (1992) Manual of forest fruits, seeds and seedlings, vol. 2. Kuala Lumpur, Forest Research Institute Malaysia.
Nikolaeva, M.G. (1977) Factors controlling the seed dormancy pattern. pp. 5174in Khan, A.A. (Ed.) The physiology and biochemistry of seed dormancy and germination. Amsterdam, North-Holland.
Nikolaeva, M.G., Rasumova, M.V. and Gladkova, V.N. (1985) Reference book on dormant seed germination. Danilova, M.F. (Ed.). Leningrad, ‘Nauka’ Publishers (in Russian).
Nikolaeva, M.G., Lyanguzova, I.V. and Pozdova, L.M. (1999) Biology of seeds. St. Petersburg, V.L. Komarov Botanical Institute, Russian Academy of Sciences (in Russian).
Quinn, C.J., Price, R.A. and Gadek, P.A. (2002) Familial concepts and relationships in the conifers based on rbcL and matK sequence comparisons. Kew Bulletin 57, 513531.
Royal Botanic Gardens Kew (2008) Seed Information Database (SID). Version 7.1. Available at (accessed accessed May 2008).
Shepperd, W.D. (2008) Ginkgoaceae–Ginkgo family. Ginkgo biloba L. pp. 559561in Bonner, F.T.; Karrfalt, R.P. (Eds) The woody plant seed manual. USDA Forest Service, Agriculture Handbook 727.
Wang, B.S.P., Lin, T.P. and Chang, T.T. (1998) Control of fungal growth with sphagnum for cold stratification and germination of tree seeds. Taiwan Journal of Forest Science 13, 101108.
West, W.C., Frattarelli, F.J. and Russin, K.J. (1970) Effect of stratification and gibberellin on seed germination in Ginkgo biloba. Bulletin of the Torrey Botanical Club 97, 380384.
Yang, C.J., Chien, C.T., Liao, Y.K., Chen, S.Y., Baskin, J.M., Baskin, C.C. and Kuo-Huang, L.L. (2011) Deep simple morphophysiological dormancy in seeds of the basal taxad Cephalotaxus. Seed Science Research 21, 215226.
Zhang, Z.-Q., Liao, W.-B., Zhong, L. and Chen, Z.-M. (2000) Biological study on seed germination of Taxus mairei. Forest Research 13, 280285(in Chinese with English abstract).
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Seed Science Research
  • ISSN: 0960-2585
  • EISSN: 1475-2735
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