The role of the testa during development and in establishment of dormancy of the legume seed

Frontiers in Plant Science

Volumn 5, Issue JUL, 2014, Pages 1-19

  Smýkal, Petr ^a   Vernoud, Vanessa ^b   Blair, Matthew W ^c   Soukup, Aleš ^d   Thompson, Richard D ^b  

^a PALACKÝ UNIVERSITY   (Czech Republic)

^b UNIV BOURGOGNE FRANCHE COMTÉ   (France)

^c TENNESSEE STATE UNIVERSITY   (United States)

^d CHARLES UNIVERSITY   (Czech Republic)

Author keywords

Domestication; Dormancy; Hardseededness; Legumes; Proanthocyanidins; Seed coat; Testa; Water permeability

Indexed keywords

GLYCINE MAX; MEDICAGO; PISUM SATIVUM;

EID: 84907964824     PISSN: None     EISSN: 1664462X     Source Type: Journal    
DOI: 10.3389/fpls.2014.00351     Document Type: Review

References (251)

1. Abbo, S., Lev-Yadun, S., and Gopher, A. (2012). Plant domestication and crop evolution in the Near East: On events and processes. Crit. Rev. Plant Sci. 31, 241-257. doi: 10.1080/07352689.2011.645428
2. Abbo, S., Pinhasi van-Oss, R., Gopher, A., Saranga, Y., Ofner, I., and Peleg, Z. (2014). Plant domestication versus crop evolution: A conceptual framework for cereals and grain legumes. Trends Plant Sci. 19, 351-360. doi: 10.1016/j.tplants.2013.12.002
3. Abbo, S., Rachamim, E., Zehavi, Y., Zezak, I., Lev-Yadun, S., and Gopher, A. (2011). Experimental growing of wild pea in Israel and its bearing on Near Eastern plant domestication. Ann. Bot. 107, 1399-1404. doi: 10.1093/aob/mcr081
4. Abbo, S., Saranga, Y., Peleg, Z., Lev-Yadun, S., Kerem, Z., and Gopher, A. (2009). Reconsidering domestication of legumes versus cereals in the ancient near east. Q. Rev. Biol. 84, 29-50. doi: 10.1086/596462
5. Abbo, S., Zezak, I., Schwartz, E., Lev-Yadun, S., and Gopher, A. (2008). Experimental harvesting of wild peas in Israel: Implications for the origins of near east farming. J. Arch. Sci. 35, 922-929. doi: 10.1016/j.jas.2007.06.016
6. Agbo, G. N., Hosfield, M. A., Uebersax, M. A., and Klomparens, K. (1987). Seed microstructure and its relationship to water uptake in isogenic lines and a cultivar of dry beans (Phaseolus vulgaris L.). Food Microstruct. 6, 91-102.
7. Algan, G., and Büyükkartal, H. N. B. (2000). Ultrastructure of seed coat development in the natural tetraploidTrifolium pratense L. J. Agron. Crop Sci. 184, 205-213. doi: 10.1046/j.1439-037x.2000.00373.x
8. Andriunas, F. A., Zhang, H. M., Xia, X., Patrick, J. W., and Offler, C. E. (2013). Intersection of transfer cells with phloem biology-broad evolutionary trends, function, and induction. Front. Plant Sci. 4:221. doi: 10.3389/fpls.2013.00221
9. Angosto, T., and Matilla, A. J. (1993). Variations in seeds of three endemic leguminous species at different altitudes.Physiol. Plant. 87, 329-334. doi: 10.1111/j.1399-3054.1993.tb01738.x
10. Argel, P., and Paton, C. (1999). "Overcoming legume hardseededness," in Forage Seed Production: Tropical and Subtropical Species, Vol. 2, eds D. S. Loch and J. E. Ferguson (Wallingford: CAB International), 247-265.
11. Ballard, L. A. T. (1973). Physical barriers to germination. Seed Sci. Technol. 1, 285-303.
12. Baskin, C. C. (2003). Breaking physical dormancy in seeds - focussing on the lens. New Phytol. 158, 229-232. doi: 10.1046/j.1469-8137.2003.00751.x
13. Baskin, C. C., and Baskin, J. M. (2014). Seeds: Ecology, Biogeography and Evolution of Dormancy and Germination, 2nd Edn. New York: Academic Press.
14. Baskin, C. C., Baskin, J. M., and Li, X. (2000). Taxonomy, anatomy and evolution of physical dormancy in seeds.Plant Species Biol. 15, 139-152. doi: 10.1046/j.1442-1984.2000.00034.x
15. Baskin, J. M., and Baskin, C. C. (1985). The annual dormancy cycle in buried weed seeds: A continuum. Bioscience35, 492-498. doi: 10.2307/1309817
16. Baskin, J. M., and Baskin, C. C. (1998). Seeds. Ecology, Biogeography and Evolution of Dormancy and Germination. New York: Academic Press.
17. Baskin, J. M., and Baskin, C. C. (2000). Evolutionary considerations of claims for physical dormancy-break by microbial action and abrasion by soil particles. Seed Sci. Res. 10, 409-413. doi: 10.1017/S0960258500000453
18. Baskin, J. M., Baskin, C. C., and Dixon, K. W. (2006). Physical dormancy in the endemic Australian genusStylobasium, a first report for the family Surinaceae (Fabales). Seed Sci. Rep. 16, 229-232. doi: 10.1079/SSR2006248
19. gri) a the P locus in common bean. Hortscience 29, 1178-1179.
20. Bassett, M. J. (1999). Allelism found between two common bean genes, hilum ring colour (D) and partly coloured seed coat pattern (Z), formerly assumed to be independent. J. Am. Soc. Hortic. Sci. 124, 649-653.
21. Bassett, M. J. (2002). Inheritance of reverse margo seed coat pattern and allelism between the genes J for seed coat colour and L for partly coloured seed coat pattern in common bean. J. Am. Soc. Hortic. Sci. 127, 56-61.
22. Bassett, M. J. (2007). Genetics of seed coat colour and pattern in common bean. Plant Breed. Rev. 28, 239-315. doi: 10.1002/9780470168028.ch8
23. Baudry, A., Heim, M. A., Dubreucq, B., Caboche, M., Weisshaar, B., and Lepiniec, L. (2004). TT2, TT8, and TTG1 synergistically specify the expression of BANYULS and proanthocyanidin biosynthesis in Arabidopsis thaliana.Plant J. 39, 366-380. doi: 10.1111/j.1365-313X.2004.02138.x
24. Beeckman, T., De Rycke, R., Viane, R., and Inzé, D. (2000). Histological study of seed coat development inArabidopsis thaliana. J. Plant Res. 113, 139-148. doi: 10.1007/PL00013924
25. Beisson, F., Li, Y., Bonaventure, G., Pollard, M., and Ohlrogge, J. B. (2007). The acyltransferase GPAT5 is required for the synthesis of suberin in seed coat and root of Arabidopsis. Plant Cell 19, 351-368. doi: 10.1105/tpc.106.048033
26. Bernard, R. L., and Weiss, M. G. (1973). "Qualitative genetics," in Soybeans: Improvement, Production, and Uses,1st Edn, ed. B. E. Caldwell (Madison, WI: American Society of Agronomy), 117-154.
27. Bevilacqua, L. R., Fossati, F., and Dondero, G. (1987). 'Callose' in the impermeable seed coat of Sesbania punicea.Ann. Bot. 59, 335-341.
28. Bewley, J. D., Bradford, K., Hilhorst, H., and Nonogaki, H. (2013). Seeds: Physiology of Development, Germination and Dormancy, 3rd Edn. New York: Springer-Verlag.
29. Bhalla, P. L., and Slattery, H. D. (1984). Callose deposits make clover seeds impermeable to water. Ann. Bot. 53, 125-128.
30. Bhat, J. L. (1968). Seed dormancy in Indigofera glandulosa Willd. Trop. Ecol. 9, 42-51.
31. Bibbey, R. O. (1948). Physiological studies of weed seed germination. Plant Physiol. 23, 467-484. doi: 10.1104/pp.23.4.467
32. Boersma, J. G., Buirchell, J. B., Sivasithamparam, K., and Yang, H. (2007). Development of a PCR marker tightly linked to mollis, the gene that controls seed dormancy in Lupinus angustifolius L. Plant Breed. 126, 612-616. doi: 10.1111/j.1439-0523.2007.01417.x
33. Bolingue, W., Ly Vu, B., Leprince, O., and Buitink, J. (2010). Characterization of dormancy behaviour in seeds of the model legume Medicago truncatula. Seed Sci. Res. 20, 97-107. doi: 10.1017/S0960258510000061
34. Bonnemain, J. L., Bourquin, S., Renault, S., Offler, C., and Fisher, D. G. (1991). "Transfer cells structure and physiology," in Phloem Transport and Assimilate Compartmentation, eds J. L. Bonnemain, S. Delrot, W. J. Lucas, and J. Dainty (Nantes: Quest Editions), 178-186.
35. Borisjuk, L., Weber, H., Panitz, R., Manteuffel, R., and Wobus, U. (1995). Embryogenesis of Vicia faba L: Histodifferentiation in relation to starch and storage protein synthesis. J. Plant Physiol. 147, 203-218. doi: 10.1016/S0176-1617(11)81507-5
36. Bradford, K., and Nonogaki, H. (2009). Seed Development, Dormancy and Germination. Annual Plant Reviews, Vol. 27. Oxford: Blackwell.
37. Brune, M., Rossander, L., and Halberg, L. (1989). Iron absorption and phenolic compounds: Importance of different phenolic structures. Eur. J. Clin. Nutr. 43, 547-558.
38. Burrows, G. E., Virgona, J. M., and Heady, R. D. (2009). Effect of boiling water, seed coat structure and provenance on the germination of Acacia melanoxylon seeds. Aust. J. Bot. 57, 139-147. doi: 10.1071/BT08194
39. Butler, E. A. (1988). "The SEM and seed identification, with particular reference to the Vicieae," in Scanning Electron Microscopy in Archeology, Vol. 452, ed. S. L. Olsen (Oxford: BAR International Series), 215-224.
40. Büyükkartal, H. N., Hatice Çölgeçen, H., Pinar, N. M., and Erdoǧan, N. (2013). Seed coat ultrastructure of hard-seeded and soft-seeded varieties of Vicia sativa. Turk. J. Bot. 37, 270-275.
41. Caldas, G. V., and Blair, M. W. (2009). Inheritance of condensed tannin content and relationship with seed colour and pattern genes in common bean (Phaseolus vulgaris L.). Theor. Appl. Genet. 119, 131-142. doi: 10.1007/s00122-009-1023-4
42. Chachalis, D., and Smith, M. L. (2000). Imbibition behavior of soybean (Glycine max (L.) Merrill) accessions with different testa characteristics. Seed Sci. Technol. 28, 321-331.
43. Chachalis, D., and Smith, M. L. (2001). Seed coat regulation of water uptake during imbibition in soybeans (Glycine max (L.) Merr.). Seed Sci. Technol. 29, 401-412.
44. Chernoff, M., Plitmann, U., and Kislev, M. E. (1992). Seed characters and testa texture in species of Vicieae: Their taxonomic significance. Isr. J. Bot. 41, 167-186. doi: 10.1080/0021213X.1992.10677225
45. Clarkson, D. T., and Robards, K. (1975). "The endodermis, its structural development and physiological role," inStructure and Function of Roots, eds J. G. Torrey and D. T. Clarkson (London: Academic Press), 415-436.
46. Clements, J. C., Buirchell, B. J., Yang, H., Smith, P. M. C., Sweetingham, M. W., and Smith, C. G. (2005). "Chapter 9, Lupin," in Genetic Resources, Chromosome Engineering, Crop Improvement, Vol. 1, Grain legumes, eds R. J Singh and P. P. Jauhar (Florida: CRC Press), 231-323.
47. Corner, E. J. H. (1951). The leguminous seed. Phytomorphology 1, 117-150.
48. Dalling, J. W., Davis, A. S., Schutte, B. J., and Arnold, A. E. (2011). Seed survival in soil: Interacting effects of predation, dormancy and the soil microbial community. J. Ecol. 99, 89-95. doi: 10.1111/j.1365-2745.2010.01739.x
49. Dean, G., Cao, Y. G., Xiang, D., Provart, N. J., Ramsay, L., Ahada, A.,et al. (2011). Analysis of gene expression patterns during seed coat development in Arabidopsis. Mol. Plant 4, 1074-1091. doi: 10.1093/mp/ssr040
50. Debeaujon, I., Léon-Kloosterziel, K. M., and Koornneef, M. (2000). Influence of the testa on seed dormancy, germination, and longevity in Arabidopsis. Plant Physiol. 122, 403-414. doi: 10.1104/pp.122.2.403
51. Debeaujon, I., Nesi, N., Perez, P., Devi, M., Grandjean, O., Caboche, M.,et al. (2003). Proanthocyanidin-accumulating cells in Arabidopsis testa: Regulation of differentiation and role in seed development. Plant Cell 15, 2514-2531. doi: 10.1105/tpc.014043
52. de Candolle, A. P. (1825). Mémoires sur la Famille des Légumineuses. Paris: A. Belin.
53. Dell, B. (1980). Structure and function of the strophiolar plug in seeds of Albizia lophantha. Am. J. Bot. 67, 556-563. doi: 10.2307/2442296
54. Deshpande, S. S., and Cheryan, M. (1986). Microstructure and water uptake of Phaseolus and winged beans. J. Food Sci. 51, 1218-1223. doi: 10.1111/j.1365-2621.1986.tb13089.x
55. de Souza, T. V., Voltolini, C. H., Santos, M., and Silveira Paulilo, M. T. (2012). Water absorption and dormancy-breaking requirements of physically dormant seeds of Schizolobium parahyba (Fabaceae - Caesalpinioideae). Seed Sci. Res. 22, 169-176.
56. Díaz, A. M., Caldas, G. V., and Blair, M. W. (2010). Concentrations of condensed tannins and anthocyanins in common bean seed coats. Food Res. Int. 43, 595-601. doi: 10.1016/j.foodres.2009.07.014
57. Dixon, R. A., Xie, D. Y., and Sharma, S. B. (2005). Proanthocyanidins - a final frontier in flavonoid research? New Phytol. 165, 9-28. doi: 10.1111/j.1469-8137.2004.01217.x
58. Donnelly, E. D., Watson, J. E., and McGuire, J. A. (1972). Inheritance of hard seed in Vicia. J. Hered. 63, 361-365.
59. Dorcey, E., Urbez, C., Blázquez, M. A., Carbonell, J., and Perez-Amador, M. A. (2009). Fertilization-dependent auxin response in ovules triggers fruit development through the modulation of gibberellin metabolism inArabidopsis. Plant J. 58, 318-332. doi: 10.1111/j.1365-313X.2008.03781.x
60. Dueberrn de Sousa, F. H. D., and Marcos-Filho, J. (2001). The seed coat as a modulator of seed-environment relationship in Fabaceae. Braz. J. Bot. 24, 365-375. doi: 10.1590/S0100-84042001000400002
61. Dueñas, M., Sun, B., Hernández, T., Estrella, I., and Spranger, M. I. (2003). Proanthocyanidin composition in the seed coat of lentils (Lens culinaris L.). J. Agric. Food Chem. 51, 7999-8004. doi: 10.1021/jf0303215
62. Emery, R. J., Ma, Q., and Atkins, C. A. (2000). The forms and sources of cytokinins in developing white lupine seeds and fruits. Plant Physiol. 123, 1593-1604. doi: 10.1104/pp.123.4.1593
63. Esau, K. (1965). Plant Anatomy, 2nd Edn. New York: John Wiley.
64. Fenner, M., and Thompson, K. (2005). The Ecology of Seeds. Cambridge: Cambridge University Press. doi: 10.1017/CBO9780511614101
65. Finch-Savage, W. E., and Leubner-Metzger, G. (2006). Seed dormancy and the control of germination. New Phytol.171, 501-523. doi: 10.1111/j.1469-8137.2006.01787.x
66. Foley, M. E. (2001). Seed dormancy: An update on terminology, physiological genetics, and quantitative trait loci regulating germinability. Weed Sci. 49, 305-317. doi: 10.1614/0043-1745(2001)049[0305:SDAUOT]2.0.CO;2
67. Forbes, I., and Well, H. D. (1968). Hard and soft seededness in blue lupine, Lupinus angustifolius L.: Inheritance and phenotype classification. Crop Sci. 4, 195-197. doi: 10.2135/cropsci1968.0011183X000800020018x
68. Frey, A., Godin, B., Bonnet, M., Sotta, B., and Marion-Poll, A. (2004). Maternal synthesis of abscisic acid controls seed development and yield in Nicotiana plumbaginifolia. Planta 218, 958-964. doi: 10.1007/s00425-003-1180-7
69. Fuller, D. Q., and Allaby, R. (2009). Seed dispersal and crop domestication: Shattering, germination and seasonality in evolution under cultivation. Ann. Plant Rev. 38, 238-295.
70. Galbiati, F., Sinha Roy, D., Simonini, S., Cucinotta, M., Ceccato, L., Cuesta, C.,et al. (2013). An integrative model of the control of ovule primordia formation. Plant J. 76, 446-455. doi: 10.1111/tpj.12309
71. Gallardo, K., Firnhaber, C., Zuber, H., Héricher, D., Belghazi, M., Henry, C.,et al. (2007). A combined proteome and transcriptome analysis of developing Medicago truncatula seeds. Mol. Cell. Proteomics 6, 2165-2179. doi: 10.1074/mcp.M700171-MCP200
72. Gijzen, M., Kuflu, K., Qutob, D., and Chernys, J. T. (2001). A class I chitinase from soybean seed coat. J. Exp. Bot.52, 2283-2289. doi: 10.1093/jexbot/52.365.2283
73. Gijzen, M., Miller, S. S., Kuflu, K., Buzzell, R. I., and Miki, B. L. (1999). Hydrophobic protein synthesized in the pod endocarp adheres to the seed surface. Plant Physiol. 120, 951-960. doi: 10.1104/pp.120.4.951
74. Gillikin, J. W., and Graham, J. S. (1991). Purification and developmental analysis of the major anionic peroxidases from the seed coat of Glycine max. Plant Physiol. 96, 214-220. doi: 10.1104/pp.96.1.214
75. Gillman, J. D., Tetlow, A., Lee, J. D., Shannon, J. G., and Bilyeu, K. (2011). Loss-of-function mutations affecting a specific Glycine max R2R3 MYB transcription factor result in brown hilum and brown seed coats. BMC Plant Biol.11:155. doi: 10.1186/1471-2229-11-155
76. Gogue, G. J., and Emino, E. R. (1979). Seed coat scarification of Albizia julibrissin Durazz by natural mechanisms. J. Am. Soc. Hortic. Sci. 104, 421-423.
77. Gopinathan, M. C., and Babu, C. R. (1985). Structural diversity and its adaptive significance in seeds of Vigna minima (Roxb.) Ohwi and Ohashi and allies (Leguminosae-Papilionoideae). Ann. Bot. 56, 723-732.
78. Graeber, K., Nakabayashi, K., Miatton, E., Leubner-Metzger, G., and Soppe, W. J. J. (2012). Molecular mechanisms of seed dormancy. Plant Cell Environ. 35, 1769-1786. doi: 10.1111/j.1365-3040.2012.02542.x
79. Güneş, F. (2013). Seed characteristics and testa textures of Pratensis, Orobon, Lathyrus, Orobastrum and Cicerculasections from Lathyrus (Fabaceae) in Turkey. Plant Syst. Evol. 299, 1935-1953. doi: 10.1007/s00606-013-0849-z
80. Gunn, C. R. (1981). Seed topography in the Fabaceae. Seed Sci. Technol. 9, 737-757.
81. Gunn, C. R. (1984). Fruits and Seeds of Genera in Subfamily Mimosoideae (Fabaceae). (Technical Bulletin No. 1681), (Washington, DC: United States Department of Agriculture), 1-194.
82. Gunn, C. R. (1991). Fruits and seeds of genera in subfamily Caesalpinioideae (Fabaceae). (Technical Bulletin No. 1755), (Washington, DC: United States Department of Agriculture), 1-408.
83. Guzmán-Maldonado, H., Castellanos, J., and Gonzalez, E. (1996). Relationship between theoretical and experimentally detected tannin content of common beans (Phaseolus vulgaris L.). Food Chem. 55, 333-335. doi: 10.1016/0308-8146(95)00106-9
84. Halevy, G. (1974). Effects of gazelles and seed beetles (Bruchidae) on germination and establishment of Acaciaspecies. Isr. J. Bot. 23, 120-123.
85. Hamly, D. H. (1932). Softening of the seeds of Melilotus alba. Bot. Gaz. 93, 345-375. doi: 10.1086/334269
86. Hamly, D. H. (1935). The light line in Melilotus alba. Bot. Gaz. 96, 753-757. doi: 10.1086/334521
87. Hammer, K. (1984). Das Domestikationssyndrom. Kulturpflanze 11, 11-34. doi: 10.1007/BF02098682
88. Hamphry, M. E., Lambrides, C. J., Chapman, S. C., Aitken, E. A. B., Imrie, B. C., and Lawn, R. J. (2005). Relationships between hardseededness and seed weight in mungbean (Vigna radiata) assessed by QTL analysis.Plant Breed. 124, 292-298. doi: 10.1111/j.1439-0523.2005.01084.x
89. Hancock, J. F. (2012). Plant Evolution and Origin of Species, 3rd Edn. Wallingford: CABI. doi: 10.1079/9781845938017.0000
90. Hanley, M. E., and Fenner, M. (1998). Pre-germination temperature and the survivorship and onward growth of Mediterranean fire-following plant species. Acta Oecol. 19, 181-187. doi: 10.1016/S1146-609X(98)80022-2
91. Hanna, P. J. (1984). Anatomical features of the seed coat of Acacia kempeana (Mueller) which relate to increased germination rate induced by heat treatment. New Phytol. 96, 23-29. doi: 10.1111/j.1469-8137.1984.tb03539.x
92. Hardham, A. R. (1976). Structural aspects of pathways of nutrient flow to developing embryo and cotyledons ofPisum sativum L. Aust. J. Bot. 24, 711-721. doi: 10.1071/BT9760711
93. Harlan, J. R. (1971). Agricultural origins: Centers and noncenters. Science 174, 468-474. doi: 10.1126/science.174.4008.468
94. Harper, J. L. (1957). "The ecological significance of dormancy and its importance in weed control," in Proceedings of the Fourth International Congress of Crop Protection, Hamburg, 415-420.
95. Harris, W. M. (1983). On the development of macrosclereids in seed coats of Pisum sativum L. Am. J. Bot. 70, 1528-1535. doi: 10.2307/2443351
96. Harris, W. M. (1984). On the development of osteosclereids in seed coats of Pisum sativum L. New Phytol. 98, 135-141. doi: 10.1111/j.1469-8137.1984.tb06103.x
97. Harris, W. M. (1987). Comparative ultrastructure of developing seed coats of 'hard-seeded' and 'soft-seeded' varieties of soybean, Glycine max (L.) Merr. Bot. Gaz. 148, 324-331. doi: 10.1086/337660
98. Hellens, R. P., Moreau, C., Lin-Wang, K., and Schwinn, K. E. (2010). Identification of Mendel's white flower character. PLoS ONE 5:e13230. doi: 10.1371/journal.pone.0013230
99. Hu, X. W., Wanga, Y. R., Wua, Y. P., and Baskin, C. C. (2009). Role of the lens in controlling water uptake in seeds of two Fabaceae (Papilionoideae) species treated with sulphuric acid and hot water. Seed Sci. Res. 19, 73. doi: 10.1017/S0960258509301099
100. Hyde, E. O. C. (1954). The function of the hilum in some Papilionaceae in relation to ripening of the seed and the permeability of the testa. Ann. Bot. 18, 241-256.
101. Isemura, T., Kaga, A., Konishi, S., Ando, T., Tomooka, H. O., and Vaughan, D. A. (2007). Genome dissection of traits related to domestication in azuki bean (Vigna angularis) and comparison with other warm-season legumes. Ann. Bot. 100, 1053-1071. doi: 10.1093/aob/mcm155
102. Isemura, T., Kaga, A., Tabata, S., Somta, P., Srinives, P., Shimizu, T.,et al. (2012). Construction of a genetic linkage map and genetic analysis of domestication related traits in mungbean (Vigna radiata). PLoS ONE 7:e41304. doi: 10.1371/journal.pone.0041304
103. Isemura, T., Kaga, A., Tomooka, N., Shimizu, T., and Vaughan, D. A. (2010). The genetics of domestication of rice bean, Vigna umbellata. Ann. Bot. 106, 927-944. doi: 10.1093/aob/mcq188
104. Kaga, A., Isemura, T., Tomooka, N., and Vaughan, D. A. (2008). The genetics of domestication of the azuki bean (Vigna angularis). Genetics 178, 1013-1036. doi: 10.1534/genetics.107.078451
105. Kantar, F. P., Hebblethwaite, D., and Pilbeam, C. J. (1996). Factors influencing disease resistance in high and low tannin Vicia faba. J. Agric. Sci. 127, 83-88. doi: 10.1017/S002185960007739X
106. Karaki, T., Watanabe, Y., Kondo, T., and Koike, T. (2012). Strophiole of seeds of the black locust acts as a water gap.Plant Species Biol. 27, 226-232. doi: 10.1111/j.1442-1984.2011.00343.x
107. Karssen, C. M. (1982). "Seasonal patterns of dormancy in weed seeds," in The Physiology and Biochemistry of Seed Development, Dormancy and Germination, ed. A. A. Khan (Amsterdam: Elsevier Biomedical Press), 243-270.
108. Karssen, C. M., Brinkhorst van der Swan, D. L. C., Breekland, A. E., and Koornneef, M. (1983). Induction of dormancy during seed development by endogenous abscisic-acid - studies on abscisic-acid deficient genotypes ofArabidopsis thaliana (L.) Heynh. Planta 157, 158-165. doi: 10.1007/BF00393650
109. Kikuchi, K., Koizumi, M., Ishida, N., and Kano, H. (2006). Water uptake by dry beans observed by micro-magnetic resonance imaging. Ann. Bot. 98, 545-553. doi: 10.1093/aob/mcl145
110. Koinange, E. M. K., Singh, S. P., and Gepts, P. (1996). Genetic control of the domestication syndrome in common bean. Crop Sci. 36, 1037-1045. doi: 10.2135/cropsci1996.0011183X003600040037x
111. Kongjaimun, A., Kaga, A., Tomooka, N., Somta, P., Vaughan, D. A., and Srinives, P. (2012). The genetics of domestication of yardlong bean, Vigna unguiculata (L.) Walp. ssp. unguiculata cv.-gr. sesquipedalis. Ann. Bot. 109, 1185-1200. doi: 10.1093/aob/mcs048
112. Korban, S. S., Coyne, D. P., and Weihing, J. L. (1981). Evaluation, variation, and genetic control of seed coat whiteness in dry beans (Phaseolus vulgaris L.). J. Am. Soc. Hortic. Sci. 166, 575-579.
113. Kovinich, N., Saleem, A., Arnason, J. T., and Miki, B. (2011). Combined analysis of transcriptome and metabolite data reveals extensive differences between black and brown nearly-isogenic soybean (Glycine max) seed coats enabling the identification of pigment isogenes. BMC Genomics 12:381. doi: 10.1186/1471-2164-12-381
114. Kurdyukov, S., Song, Y., Sheahan, M. B., and Rose, R. J. (2014). Transcriptional regulation of early embryo development in the model legume Medicago truncatula. Plant Cell Rep. 33, 349-362. doi: 10.1007/s00299-013-1535-x
115. Lacerda, D. R., Lemos-Filho, J. P., Goulart, M. F., Ribeiro, R. A., and Lovato, M. B. (2004). Seed-dormancy variation in natural populations of two tropical leguminous tree species: Senna multijuga (Caesalpinoideae) andPlathymenia reticulata (Mimosoideae). Seed Sci. Res. 14, 127-135. doi: 10.1079/SSR2004162
116. Lackey, J. A. (1981). Systematic significance of the epihilum in Phaseoleae (Fabaceae, Faboideae). Bot. Gaz. 142, 160-164. doi: 10.1086/337207
117. Ladizinsky, G. (1979). The origin of lentil and its wild genepool. Euphytica 28, 179-187. doi: 10.1007/BF00029189
118. Ladizinsky, G. (1985). The genetics of hard seed coat in the genus Lens. Euphytica 34, 539-543. doi: 10.1007/BF00022952
119. Ladizinsky, G. (1987). Pulse domestication before cultivation. Econ. Bot. 41, 60-65. doi: 10.1007/BF02859349
120. Ladizinsky, G. (1998). Plant Evolution Under Domestication. Dordrecht: Kluwer Academic Publishers. doi: 10.1007/978-94-011-4429-2
121. Lanfermeijer, F. C., van Oene, M. A., and Borstlap, A. C. (1992). Compartmental analysis of amino-acid release from attached and detached pea seed coats. Planta 187, 75-82. doi: 10.1007/BF00201626
122. Le, B. H., Wagmaister, J. A., Kawashima, T., Bui, A. Q., Harada, J. J., and Goldberg, R. B. (2007). Using genomics to study legume seed development. Plant Physiol. 144, 562-574. doi: 10.1104/pp.107.100362
123. Legesse, N., and Powell, A. A. (1996). Relationship between the development of seed coat pigmentation, seed coat adherence to the cotyledons and the rate of imbibition during the maturation of grain legumes. Seed Sci. Technol.24, 23-32.
124. Lepiniec, L., Debeaujon, I., Routaboul, J. M., Baudry, A., Pourcel, L., Nesi, N.,et al. (2006). Genetics and biochemistry of seed flavonoids. Annu. Rev. Plant Biol. 57, 405-430. doi: 10.1146/annurev.arplant.57.032905.105252
125. Lersten, N. R. (1982). Tracheid bar and vestured pits in legume seeds (Leguminosae: Papilionoideae). Am. J. Bot.69, 98-107. doi: 10.2307/2442834
126. Lersten N. R., and Gunn, C. R. (1981). Seed morphology and testa topography in Cicer (Fabaceae: Faboideae). Syst. Bot. 6, 223-230.
127. Lersten, N. R., and Gunn, C. R. (1982). Testa Characters in Tribe Vicieae, with Notes About Tribes Abreae, Cicereae, and Trifolieae (Fabaceae). (Technical Bulletin No. 1667), (Washington, DC: United States Department of Agriculture), 1-40.
128. Lersten, N. R., Gunn, C. R., and Brubaker, C. L. (1992). Comparative Morphology of the Lens on Legume (Fabaceae) Seeds, with Emphasis on Species in Subfamilies Caesalpinioideae and Mimosoideae. (Technical Bulletin No. 1791), (Washington, DC: United States Department of Agriculture), 44.
129. Lewis, G., Schrire, B., Mackinder, B., and Lock, M. (2005). Legumes of the World. Kew: Royal Botanic Gardens
130. Lhotská, M., and Chrtková, A. (1978). Karpologie a diasporologie: československých zástupců čeledi Fabaceae. Prague: Academia.
131. Li, X., Buirchell, B., Yan, G., and Yang, H. (2012). A molecular marker linked to the mollis gene conferring soft-seediness for marker-assisted selection applicable to a wide range of crosses in lupin (Lupinus angustifolius L.).Breed. Mol. Breed. 29, 361-370. doi: 10.1007/s11032-011-9552-3
132. Liang, M., Davis, E., Gardner, D., Cai, X., and Wu, Y. (2006). Involvement of AtLAC15 in lignin synthesis in seeds and in root elongation of Arabidopsis. Planta 224, 1185-1196. doi: 10.1007/s00425-006-0300-6
133. Liu, B., Liu, X., Wang, C., Jin, J., and Herbert, S. J. (2010). Endogenous hormones in seed, leaf, and pod wall and their relationship to seed filling in soybeans. Crop Pasture Sci. 61, 103-110. doi: 10.1071/CP09189
134. Liu, W., Peffley, E. B., Powell, R. J., Auld, D. L., and Hou, A. (2007). Association of seedcoat colour with seed water uptake, germination, and seed components in guar (Cyamopsis tetragonoloba (L.) Taub). J. Arid Environ. 70, 29-38. doi: 10.1016/j.jaridenv.2006.12.011
135. Liu, Y. H., Cao, J. S., Li, G. J., Wu, X. H., Wang, B. G., Xu, P.,et al. (2012). Genotypic differences in pod wall and seed growth relate to invertase activities and assimilate transport pathways in asparagus bean. Ann. Bot. 109, 1277-1284. doi: 10.1093/aob/mcs060
136. Lush, W. M., and Evans, L. T. (1980). The seed coats of cowpeas and other grain legumes - structure in relation to function. Field Crops Res. 3, 267-286. doi: 10.1016/0378-4290(80)90034-9
137. Ma, F., Cholewa, E., Mohamed, T., Peterson, C. A., and Gijzen, M. (2004). Cracks in the palisade cuticle of soybean seed coats correlate with their permeability to water. Ann. Bot. 94, 213-228. doi: 10.1093/aob/mch133
138. Ma, Y., and Bliss, F. A. (1978). Tannin content and inheritance in common bean. Crop Sci. 18, 201-204. doi: 10.2135/cropsci1978.0011183X001800020001x
139. Mandal, S. M., Chakraborty, D., and Dey, S. (2010). Phenolic acids act as signalling molecules in plant-microbe symbioses. Plant Signal. Behav. 5, 359-368. doi: 10.4161/psb.5.4.10871
140. Marbach, I., and Mayer, A. M. (1974). Permeability of seed coats to water as related to drying conditions and metabolism of phenolics. Plant Physiol. 54, 817-820. doi: 10.1104/pp.54.6.817
141. Marles, S. M. A., and Gruber, M. Y. (2004). Histochemical characterization of unextractable seed coat pigments and quantification of extractable lignin in the Brassicaceae. J. Sci. Food Agric. 84, 251-262. doi: 10.1002/jsfa.1621
142. Matilla, A., Gallardo, M., and Puga-Hermida, M. I. (2005). Structural, physiological and molecular aspects of heterogeneity in seeds: A review. Seed Sci. Res. 15, 63-76. doi: 10.1079/SSR2005203
143. McDonald, M. B., Vertucci, C. W., and Roos, E. E. (1988). Soybean seed imbibition: Water absorption by seed parts.Crop Sci. 28, 993-997. doi: 10.2135/cropsci1988.0011183X002800060026x
144. McKeon, G. M., and Mott, J. J. (1982). The effect of temperature on the field softening of hard seed of Stylosanthes humilis and S. hamata in dry monsoonal climate. Aust. J. Agric. Res. 33, 75-85. doi: 10.1071/AR9820075
145. Meyer, C. J., Steudle, E., and Peterson, C. A. (2007). Patterns and kinetics of water uptake by soybean seeds. J. Exp. Bot. 58, 717-732. doi: 10.1093/jxb/erl244
146. Miao, Z. H., Fortune, J. A., and Gallagher, J. (2001). Anatomical structure and nutritive value of lupin seed coats.Aust. J. Agric. Res. 52, 985-993. doi: 10.1071/AR00117
147. Miernyk, J. A., and Johnston, M. L. (2013). Proteomic analysis of the testa from developing soybean seeds. J. Proteomics 89, 265-272. doi: 10.1016/j.jprot.2013.05.013
148. Milberg, P., and Andersson, L. (1997). Seasonal variation in dormancy and light sensitivity in buried seeds of eight annual weed species. Can. J. Bot. 75, 1998-2004. doi: 10.1139/b97-911
149. Miller, S. S., Bowman, L. A., Gijzen, M., and Miki, B. L. A. (1999). Early development of the seed coat of soybean (Glycine max). Ann. Bot. 84, 297-304. doi: 10.1006/anbo.1999.0915
150. Miller, S. S., Jin, Z., Schnell, J. A., Romero, M. C., Brown, D. C. W., and Johnson, D. A. (2010). Hourglass cell development in the soybean seed coat. Ann. Bot. 106, 235-242. doi: 10.1093/aob/mcq101
151. Miranda, M., Borisjuk, L., Tewes, A., Heim, U., Sauer, N., Wobus, U.,et al. (2001). Amino acid permeases in developing seeds of Vicia faba L.: Expression precedes storage protein synthesis and is regulated by amino acid supply. Plant J. 28, 61-71. doi: 10.1046/j.1365-313X.2001.01129.x
152. Mizzotti, C., Mendes, M. A., Caporali, E., Schnittger, A., Kater, M. M., Battaglia, R.,et al. (2012). The MADS box genes SEEDSTICK and ARABIDOPSIS B sister play a maternal role in fertilization and seed development. Plant J. 70, 409-420. doi: 10.1111/j.1365-313X.2011.04878.x
153. Mohamed-Yasseen, Y., Barringer, S. A., Splittstoesser, W. E., and Costanza, S. (1994). The role of seed coats in seed viability. Bot. Rev. 60, 426-439. doi: 10.1007/BF02857926
154. Moïse, J. A., Han, S., Gudynaite-Savitch, L., Johnson, D. A., and Miki, B. L. A. (2005). Seed coats: Structure, development, composition, and biotechnology. In Vitro Cell. Dev. Biol. Plant 41, 620-644. doi: 10.1079/IVP2005686
155. Moreau, C., Ambrose, M. J., Turner, L., Hill, L., Ellis, T. H., and Hofer, J. M. (2012). The B gene of pea encodes a defective flavonoid 3′,5′-hydroxylase, and confers pink flower color. Plant Physiol. 159, 759-768. doi: 10.1104/pp.112.197517
156. Moreno-Casasola, P., Grime, J. P., and Martinez, L. (1994). A comparative study of the effects of fluctuations in temperature and moisture supply on hard coat dormancy in seeds of coastal tropical legumes in Mexico. J. Trop. Ecol. 10, 67-86. doi: 10.1017/S0266467400007720
157. Morrison, D. A., McClay, K., Porter, C., and Rish, S. (1998). The role of the lens in controlling heat-induced breakdown of testa-imposed dormancy and native Australian legumes. Ann. Bot. 82, 35-40. doi: 10.1006/anbo.1998.0640
158. Murray, D. R., and Kennedy, I. R. (1980). Changes in activities of enzymes of nitrogen-metabolism in seed coats and cotyledons during embryo development in pea seeds. Plant Physiol. 66, 782-786. doi: 10.1104/pp.66.4.782
159. Nadeau, C. D., Ozga, J. A., Kurepin, L. V., Jin, A., Pharis, R. P., and Reinecke, D. M. (2011). Tissue-specific regulation of gibberellin biosynthesis in developing pea seeds. Plant Physiol. 156, 897-912. doi: 10.1104/pp.111.172577
160. Nagai, I. (1921). A genetico-physiological study on the formation of anthocyanin and brown pigments in plants. J. Coll. Agric. Imp. Univ. Tokyo 8, 1-92.
161. Nawrath, C. (2002). The biopolymers cutin and suberin. Arabidopsis Book 1, e0021. doi: 10.1199/tab.0021
162. Nikolaeva, M. G. (1969). Physiology of Deep Dormancy in Seeds. Izdatel'stvo Nauka. Leningrad (Translated from Russian by Z. Shapiro, NSF, Washington, D.C.).
163. Nikolaeva, M. G. (1977). "Factors controlling the seed dormancy pattern," in The Physiology and Biochemistry of Seed Dormancy and Germination, ed. A. A. Khan (Amsterdam: North-Holland Publishing Co.), 51-74.
164. Noodén, L. D., Blakley, K. A., and Grzybowski, J. M. (1985). Control of seed coat thickness and permeability in soybean: A possible adaptation to stress. Plant Physiol. 79, 543-545. doi: 10.1104/pp.79.2.543
165. Nowack, M. K., Ungru, A., Bjerkan, K. N., Grini, P. E., and Schnittger, A. (2010). Reproductive cross-talk: Seed development in flowering plants. Biochem. Soc. Trans. 38, 604-612. doi: 10.1042/BST0380604
166. Offler, C. E., Nerlich, S. M., and Patrick, J. W. (1989). Pathway of photosynthate transfer in the developing seed ofVicia faba L. transfer in relation to seed anatomy. J. Exp. Bot. 40, 769-780. doi: 10.1093/jxb/40.7.769
167. Offler, C. E., and Patrick, J. W. (1984). Cellular structures, plasma-membrane surface-areas and plasmodesmatal frequencies of seed coats of Phaseolus vulgaris L. in relation to photosynthate transfer. Aust. J. Plant Physiol. 11, 79-99. doi: 10.1071/PP9840079
168. Offler, C. E., and Patrick, J. W. (1993). Pathway of photosynthate transfer in the developing seed of Vicia faba L: A structural assessment of the role of transfer cells in unloading from the seed coat. J. Exp. Bot. 44, 711-724. doi: 10.1093/jxb/44.4.711
169. Oliveira, A. E. A., Sassaki, G. L., Iacomini, M., da Cunha, M., Gomes, V. M., Fernandes, B. K. V. S.,et al. (2001). Isolation and characterization of a galactorhamnan polysaccharide from the seed coat of Canavalia ensiformis that is toxic to the cowpea weevil (Callosobruchus maculatus). Entomol. Exp. Appl. 101, 225-231. doi: 10.1046/j.1570-7458.2001.00907.x
170. Ozga, J. A., Reinecke, D. M., Ayele, B. T., Ngo, P., Nadeau, C., and Wickramarathna, A. D. (2009). Developmental and hormonal regulation of gibberellin biosynthesis and catabolism in pea fruit. Plant Physiol. 150, 448-462. doi: 10.1104/pp.108.132027
171. Palmer, R. G., and Kilen, T. C. (1987). "Quantitative genetics: Results relevant to soybean breeding," in Soybean: Improvements, Production, and Uses, 2nd Edn, ed. J. R. Wilcox (Madison, WI: Agronomy Society of America),135-209.
172. Pammel, L. H. (1899). Anatomical Characters of the Seeds of Leguminosae: Chiefly Genera of Gray's Manual, Shaw School of Botany. Ph.D. thesis, Transactions of the Academy of Science of St. Louis, Washington University, St. Louis Bibliography, St. Louis, MI.
173. Pang, Y. Z., Peel, G. J., Wright, E., Wang, Z., and Dixon, R. A. (2007). Early steps in proanthocyanidin biosynthesis in the model legume Medicago truncatula. Plant Physiol. 145, 601-615. doi: 10.1104/pp.107.107326
174. Pang, Y. Z., Wenger, J. P., Saathoff, K., Peel, G. J., Wen, J., Huhman, D.,et al. (2009). A WD40 repeat protein fromMedicago truncatula is necessary for tissue-specific anthocyanin and proanthocyanidin biosynthesis but not for trichome development. Plant Physiol. 151, 1114-1129. doi: 10.1104/pp.109.144022
175. Park, S., Ozga, J. A., Cohen, J. D., and Reinecke, D. M. (2010). Evidence for 4-Cl-IAA and IAA bound to proteins in pea fruits and seeds. J. Plant Growth Regul. 29, 184-193. doi: 10.1007/s00344-009-9123-6
176. Paulsen, T. R., Colville, L., Kranner, I., Daws, M. I., Högstedt, G., Vandvik, V.,et al. (2013). Physical dormancy in seeds: A game of hide and seek? New Phytol. 198, 496-503. doi: 10.1111/nph.12191
177. Pellew, R. A., and Southgate, B. J. (1984). The parasitism of Acacia tortilis seeds in the Serengeti. Afr. J. Ecol. 22, 73-75. doi: 10.1111/j.1365-2028.1984.tb00679.x
178. Probert, R. J. (2000). "The role of temperature in the regulation of seed dormancy and germination," in Seeds. The Ecology of Regeneration in Plant Communities, 2nd Edn, ed. M. Fenner (Wallingford: CABI publishing), 261-292. doi: 10.1079/9780851994321.0261
179. Quesnelle, P. E., and Emery, R. J. N. (2007). cis-Cytokinins that predominate in Pisum sativum during early embryogenesis will accelerate embryo growth in vitro. Can. J. Bot. 85, 91-103. doi: 10.1139/b06-149
180. Quinlivan, B. J. (1961). The effect of constant and fluctuating temperatures on the permeability of the hard seeds of some legume species. Aust. J. Agric. Res. 12, 1009-1022. doi: 10.1071/AR9611009
181. Quinlivan, B. J. (1968). The softening of hard seeds of sand-plain lupin (Lupinus varius L.). Aust. J. Agric. Res. 19, 507-515. doi: 10.1071/AR9680507
182. Quinlivan, B. J. (1971). Seed coat impermeability in legumes. J. Aust. Inst. Agric. Sci. 37, 283-295.
183. Ramsay, G. (1997). Inheritance and linkage of a gene for testa imposed seed dormancy in faba bean (Vicia faba L.).Plant Breed. 116, 287-289. doi: 10.1111/j.1439-0523.1997.tb00998.x
184. Ranathunge, K., Shao, S., Qutob, D., Gijzen, M., Peterson, C. A., and Bernards, M. A. (2010). Properties of the soybean seed coat cuticle change during development. Planta 231, 1171-1188. doi: 10.1007/s00425-010-1118-9
185. Reinecke, D. M. (1999). 4-Chloroindole-3-acetic acid and plant growth. Plant Growth Regul. 27, 3-13. doi: 10.1023/A:1006191917753
186. Reinprecht, Y., Yadegari, Z., Perry, G. E., Siddiqua, M., Wright, L. C., McClean, P. E.,et al. (2013). In silicocomparison of genomic regions containing genes coding for enzymes and transcription factors for the phenylpropanoid pathway in Phaseolus vulgaris L. and Glycine max L. Merr. Front. Plant Sci. 4:317. doi: 10.3389/fpls.2013.00317
187. Riggio Bevilacqua, L., Roti-Mihelozzi, G., and Modenesi, P. (1989). The watertight dormancy of Melilotus albaseeds: further observations on the palisade cell wall. Can. J. Bot. 67, 3453-3456. doi: 10.1139/b89-422
188. Roberts, H. A., and Boddrell, J. E. (1985). Seed survival and seasonal pattern of seedling emergence in some Leguminosae. Ann. Appl. Biol. 106, 125-132. doi: 10.1111/j.1744-7348.1985.tb03102.x
189. Rochat, C., and Boutin, J. P. (1992). Temporary storage compounds and sucrose-starch metabolism in seed coats during pea seed development (Pisum sativum). Physiol. Plant 85, 567-572. doi: 10.1111/j.1399-3054.1992.tb04756.x
190. Rolston, M. P. (1978). Water impermeable seed dormancy. Bot. Rev. 44, 365-396. doi: 10.1007/BF02957854
191. Russi, L., Cocks, P. S., and Roberts, E. H. (1992). Coat thickness and hard-seededness in some Medicago andTrifolium species. Seed Sci. Res. 2, 243-249. doi: 10.1017/S0960258500001434
192. Sabiiti, E. N., and Wein, R. W. (1987). Fire and Acacia seeds: A hypothesis of colonization success. J. Ecol. 74, 937-946. doi: 10.2307/2260305
193. Schlumbaum, A., Mauch, F., Vögeli, U., and Boller, T. (1986). Plant chitinases are potent inhibitors of fungal growth. Nature 324, 365-367. doi: 10.1038/324365a0
194. Schneitz, K., Hülskamp, M., Kopczak, S. D., and Pruitt, R. E. (1997). Dissection of sexual organ ontogenesis: A genetic analysis of ovule development in Arabidopsis thaliana. Development 124, 1367-1376.
195. Serrato Valenti, G., Cornara, L., Ferrando, M., and Modenesi, P. (1993). Structural and histochemical features ofStylosanthes scabra (Leguminosae: Papilionoideae) seed coat as related to water entry. Can. J. Bot. 71, 834-840. doi: 10.1139/b93-095
196. Serrato Valenti, G., Melone, L., Ferrando, M., and Bozzini, A. (1989). Comparative studies on testa structure of hard-seeded and soft-seeded varieties of Lupinus angustifolius L. (Leguminosae) and mechanisms of water entry.Seed Sci. Technol. 17, 563-581.
197. Serrato Valenti, G., Modenesi, P., Roti-Mihelozzi, G., and Bevilacqua, L. (1986). Structural and histochemical characters of the Prosopis tamarugo Phil. seed coat in relation to its hardness. Acta Bot. Nederland 35, 475-487.
198. Serrato-Valenti, G., De Vries, M., and Cornara, L. (1995). The hilar region in Leucaena leucocephala Lam. (De Wit) seed: Structure, histochemistry and the role of the lens in germination. Ann. Bot. 75, 569-574. doi: 10.1006/anbo.1995.1060
199. Shao, S., Meyer, C. J., Ma, F., Peterson, C. A., and Bernards, M. A. (2007). The outermost cuticle of soybean seeds: Chemical composition and function during imbibition. J. Exp. Bot. 58, 1071-1082. doi: 10.1093/jxb/erl268
200. Shen-Miller, J., Mudgett, M. B., Schopf, J. W., Clarke, R., and Berger, R. (1995). Exceptional seed longevity and robust growth: Ancient Sacred Lotus from China. Am. J. Bot. 82, 1367-1380. doi: 10.2307/2445863
201. Simao Neto, M., Jones, R. M., and Ratcliff, D. (1987). Recovery of pasture seed ingested by ruminants. 1. Seed of six tropical pasture species fed to cattle, sheep and goats. Aust. J. Exp. Agric. 27, 239-246. doi: 10.1071/EA9870239
202. Skinner, D. J., Hill, T. A., and Gasser, C. S. (2004). Regulation of ovule development. Plant Cell 16, S32-S45. doi: 10.1105/tpc.015933
203. Slater, S. M., Yuan, H. Y., Lulsdorf, M. M., Vandenberg, A., Zaharia, L. I., Han, X.,et al. (2013). Comprehensive hormone profiling of the developing seeds of four grain legumes. Plant Cell Rep. 32, 1939-1952. doi: 10.1007/s00299-013-1505-3
204. Slattery, H. D., Atwell, B. J., and Kuo, J. (1982). Relationship between colour, phenolic content and impermeability in the seed coat of various Trifolium subterraneum L. genotypes. Ann. Bot. 50, 373-378.
205. Smýkal, P., Coyne, C. J., Ambrose, M. J., Maxted, N., Schaefer, H., Blair, M.,et al. (2014). Legume crops phylogeny and genetic diversity for science and breeding. Crit. Rev. Plant Sci. doi: 10.1080/07352689.2014.897904 [Epub ahead of print].
206. Spurny, M. (1963). Cell wall structure of epidermal cells of the pea seed coat (Pisum sativum L.) studied by microcinematography. Mikroskopie 18, 272-279.
207. Spurny, M. (1964). Changes in the permeability of the seed coat in connection with the development of suberin adcrustations of the macrosclereids from the seed coat of the pea (Pisum sativum L.). Flora 154, 547-567.
208. Stevenson, T. M. (1937). Sweet clover studies on habit of growth, seed pigmentation and permeability of the seed coat. Sci. Agric. 17, 627-654.
209. Taylor, G. B. (1981). Effect of constant temperature treatments followed by alternating temperatures on the softening of hard seeds of Trifolium subterraneum L. Aust. J. Plant Physiol. 8, 547-558. doi: 10.1071/PP9810547
210. Taylor, G. B. (2005). Hardseededness in Mediterranean annual pasture legumes in Australia: A review. Aust. J. Agric. Res. 56, 645-661. doi: 10.1071/AR04284
211. Taylor, G. B., and Ewing, M. A. (1996). Effects of extended (4-12 years) burial on seed softening in subterranean clover and annual medics. Aust. J. Exp. Agric. 36, 145-150. doi: 10.1071/EA9960145
212. Taylor, G. B., and Revell, C. K. (1999). Effect of pod burial, light, and temperature on seed softening in yellow serradella. Aust. J. Agric. Res. 50, 1203-1209. doi: 10.1071/AR98206
213. Taylor, G. B., and Revell, C. K. (2002). Seed softening, imbibition time, and seedling establishment in yellow serradella. Aust. J. Agric. Res. 53, 1011-1018. doi: 10.1071/AR01201
214. Telewski, F. W., and Zeevaart, J. A. (2002). The 120-yr period for Dr. Beal's seed viability experiment. Am. J. Bot.89, 1285-1288. doi: 10.3732/ajb.89.8.1285
215. Thompson, K., and Ooi, M. K. J. (2013). Germination and dormancy breaking: Two different things. Seed Sci. Res.23, 1. doi: 10.1017/S0960258512000190
216. Thompson, R., Burstin, J., and Gallardo, K. (2009). Post-genomics studies of developmental processes in legume seeds. Plant Physiol. 151, 1023-1029. doi: 10.1104/pp.109.143966
217. Thompson, R. D., Hueros, G., Becker, H., and Maitz, M. (2001). Development and functions of seed transfer cells.Plant Sci. 160, 775-783. doi: 10.1016/S0168-9452(01)00345-4
218. Thorne, J. H. (1981). Morphology and ultrastructure of maternal seed tissues of soybean in relation to the import of photosynthate. Plant Physiol. 67, 1016-1025. doi: 10.1104/pp.67.5.1016
219. Thorne, J. H., and Rainbird, R. M. (1983). An in vivo technique for the study of phloem unloading in seed coats of developing soybean seeds. Plant Physiol. 72, 268-271. doi: 10.1104/pp.72.1.268
220. Tuteja, J. H., Clough, S. J., Chan, W. C., and Vlodking, L. O. (2004). Tissue specific gene silencing mediated by a naturally occurring chalcone synthase gene cluster in Glycine max. Plant Cell 16, 819-835. doi: 10.1105/tpc.021352
221. Van Assche, J. A., Debucquoy, K. L. A., and Rommens, W. A. F. (2003). Seasonal cycles in the germination capacity of buried seeds of some Leguminosae (Fabaceae). New Phytol. 158, 315-323. doi: 10.1046/j.1469-8137.2003.00744.x
222. Van Dongen, J. T., Ammerlaan, A. M., Wouterlood, M., Van Aelst, A. C., and Borstlap, A. C. (2003). Structure of the developing pea seed coat and the post-pholem transport pathway of nutrients. Ann. Bot. 93, 729-737. doi: 10.1093/aob/mcg066
223. van Klinken, K. D., and Goulier, J. B. (2013). Habitat-specific seed dormancy-release mechanisms in four legume species. Seed Sci. Res. 23, 181-188. doi: 10.1017/S0960258513000196
224. Van Staden, J., Manning, J. C., and Kelly, K. M. (1989). Legume seeds - the structure: function equation. Monogr. Syst. Bot. Missouri Bot. Gard. 29, 417-450.
225. Vaughan, D. A., Bernard, R. L., Sinclair, J. B., and Kunwar, I. K. (1987). Soybean seed coat development. Crop Sci.27, 759-765. doi: 10.2135/cropsci1987.0011183X002700040031x
226. Vázquez-Yanes, C., and Orozco-Segovia, A. (1982). Seed germination of tropical rain forest pioneer tree (Heliocarpus donnell-smithii) in response to diurnal fluctuation in temperature. Physiol. Plant. 56, 295-298. doi: 10.1111/j.1399-3054.1982.tb00341.x
227. Verdier, J., Dessaint, F., Schneider, C., and Abirached-Darmency, M. (2013a). A combined histology and transcriptome analysis unravels novel questions on Medicago truncatula seed coat. J. Exp. Bot. 64, 459-470. doi: 10.1093/jxb/ers304
228. Verdier, J., Lalanne, D., Pelletier, S., Torres-Jerez, I., Righetti, K., Bandyopadhyay, K.,et al. (2013b). A regulatory network-based approach dissects late maturation processes related to the acquisition of desiccation tolerance and longevity of Medicago truncatula seeds. Plant Physiol. 163, 757-774. doi: 10.1104/pp.113.222380
229. Verdier, J., Kakar, K., Gallardo, K., Le Signor, C., Aubert, G., Schlereth, A.,et al. (2008). Gene expression profiling ofM. truncatula transcription factors identifies putative regulators of grain legume seed filling. Plant Mol. Biol. 67, 567-580. doi: 10.1007/s11103-008-9320-x
230. Verdier, J., Zhao, J., Torres-Jerez, I., Ge, S., Liu, C., He, X.,et al. (2012). MtPAR MYB transcription factor acts as an on switch for proanthocyanidin biosynthesis in Medicago truncatula. Proc. Natl. Acad. Sci. U.S.A. 109, 1766-1771. doi: 10.1073/pnas.1120916109
231. Vleeshouwers, L. M., Bouwmeester, H. J., and Karssen, C. M. (1995). Redefining seed dormancy: An attempt to integrate physiology and ecology. J. Ecol. 83, 1031-1037. doi: 10.2307/2261184
232. Wang, H. L., and Grusak, M. A. (2005). Structure and development of Medicago truncatula pod wall and seed coat.Ann. Bot. 95, 737-747. doi: 10.1093/aob/mci080
233. Warrag, E. I., and Eltigani, M. A. (2005). Breaking seed coat dormancy of Acacia nilotica seeds under simulated natural habit conditions in Sudan. Trop. Ecol. 46, 127-132.
234. Weber, H., Borisjuk, L., Heim, U., Sauer, N., and Wobus, U. (1997). A role for sugar transporters during seed development: Molecular characterization of a hexose and a sucrose carrier in fava bean seeds. Plant Cell 9, 895-908. doi: 10.1105/tpc.9.6.895
235. Weber, H., Borisjuk, L., and Wobus, U. (2005). Molecular physiology of legume seed development. Annu. Rev. Plant Biol. 56, 253-279. doi: 10.1146/annurev.arplant.56.032604.144201
236. Weeden, N. F. (2007). Genetic changes accompanying the domestication of Pisum sativum: Is there a common genetic basis to the 'domestication syndrome' for legumes? Ann. Bot. 100, 1017-1025. doi: 10.1093/aob/mcm122
237. Weiss, E., Kislev, M. E., and Hartmann, A. (2006). Autonomous cultivation before domestication. Science 312, 1608-1610. doi: 10.1126/science.1127235
238. Werker, E., Marbach, I., and Mayer, A. M. (1979). Relation between the anatomy of the testa, water permeability and the presence of phenolics in the genus Pisum. Ann. Bot. 43, 765-771.
239. White, J. (1908). The occurrence of an impermeable cuticle on the exterior of certain seeds. Proc. Royal Soc. Victoria 21, 203-210.
240. Winkel-Shirley, B. (2001). Flavonoid biosynthesis. a colourful model for genetics, biochemistry, cell biology, and biotechnology. Plant Physiol. 126, 485-493. doi: 10.1104/pp.126.2.485
241. Wolf, W., Baker, F. L., and Bernard, R. L. (1981). Soybean seed-coat structural features: pits, deposits and cracks.Scan. Electron Microsc. 3, 531-544.
242. Wyatt, J. E. (1977). Seed coat and water absorption properties of seed of near-isogenic snap bean lines differing in seed coat color. J. Am. Soc. Hortic. Sci. 102, 478-480.
243. Yaklich, R. W., Vigil, E. L., and Wergin, W. P. (1986). Pore development and seed coat permeability in soybean.Crop Sci. 26, 616-624. doi: 10.2135/cropsci1986.0011183X002600030041x
244. Yang, K., Jeong, N., Moon, J. K., Lee, Y. H., Lee, S. H., Kim, H. M.,et al. (2010). Genetic analysis of genes controlling natural variation of seed coat and flower colours in soybean. J. Hered. 101, 757-768. doi: 10.1093/jhered/esq078
245. Zabala, G., and Vodkin, L. O. (2003). Cloning of the pleiotropic T locus in soybean and two recessive alleles that differentially affect structure and expression of the encoded flavonoid 3 hydroxylase. Genetics 163, 295-309.
246. Zabala, G., and Vodkin, L. O. (2005). The wp mutation of Glycine max carries a gene fragment-rich transposon of the CACTA superfamily. Plant Cell 17, 2619-2632. doi: 10.1105/tpc.105.033506
247. Zabala, G., and Vodkin, L. O. (2007). A rearrangement resulting in small tandem repeats in the F3′5′H gene of white flower genotypes is associated with the soybean W1 locus. Crop Sci. 47, S113-S124. doi: 10.2135/cropsci2006.12.0838tpg
248. Zhao, J., and Dixon, R. A. (2009). MATE transporters facilitate vacuolar uptake of epicatechin 3′-o-glucoside for proanthocyanidin biosynthesis in Medicago truncatula and Arabidopsis. Plant Cell 21, 2323-2340. doi: 10.1105/tpc.109.067819
249. Zhao, J., Pang, Y., and Dixon, R. A. (2010). The mysteries of proanthocyanidin transport and polymerization. Plant Physiol. 153, 437-443. doi: 10.1104/pp.110.155432
250. Zhou, S., Sekizaki, H., Yang, Z., Sawa, S., and Pan, J. (2010). Phenolics in the seed coat of wild soybean (Glycine soja) and their significance for seed hardness and seed germination. J. Agric. Food Chem. 58, 10972-10978. doi: 10.1021/jf102694k
251. Zohary, D., Hopf, M., and Weiss, E. (2012). Domestication of Plants in the Old World, 4th Edn. Oxford: Oxford University Press.