Skip to main content Accessibility help

Free cyclitol unloading from seed coats on stem–leaf–pod explants of low-raffinose, low-stachyose, low-phytin soybean

  • Suzanne M. Kosina (a1), Steven R. Schnebly (a2) and Ralph L. Obendorf (a1)


Raffinose, stachyose and phytin are undesirable compounds for soybean food and animal feed products. In seeds, raffinose and stachyose are believed to contribute to desiccation and cold stress tolerance. Thus, removal of these compounds from soybean by genetic mutation has resulted in a more commercially desirable composition, but potentially less physiologically viable seeds. In an effort to develop a method to improve viability and seed storability in soybean, stem–leaf–pod explants of three low raffinose, low stachyose lines, two of which were also low in phytin, and a check line were fed solutions containing d-chiro-inositol, myo-inositol or d-pinitol, free cyclitols which unload through the seed coat to the developing embryo where they accumulate as fagopyritols, galactinol and galactopinitols, respectively, during seed maturation. Increased galactopinitol and fagopyritol accumulation may substitute for the roles of raffinose and stachyose in low raffinose, stachyose and phytin seeds. Explants of all lines unloaded d-chiro-inositol, myo-inositol and d-pinitol. Fed d-chiro-inositol accumulated in leaf tissues demonstrating uptake into explants. Fed d-chiro-inositol and myo-inositol accumulated in pod wall and seed coat tissues of one or more lines. The results indicate that d-chiro-inositol was unloaded from the seed coat to the embryo in increased amounts after feeding. The potential use of increased maternal d-chiro-inositol for synthesis of fagopyritols in embryos to improve seed performance in low-stachyose and low-phytin soybean seeds is supported. The seed coat cup unloading of fed free cyclitols may provide a model system to test effective unloading of upregulated maternally synthesized cyclitols.


Corresponding author

*Correspondence Fax: +1-607-255-2644 Email:


Hide All
Blackman, S.A., Obendorf, R.L. and Leopold, A.C. (1992) Maturation proteins and sugars in desiccation tolerance of developing soybean seeds. Plant Physiology 100, 225230.
Buitink, J., Thomas, M., Gissot, L. and Leprince, O. (2004) Starvation, osmotic stress and desiccation tolerance lead to expression of different genes of the regulatory beta and gamma subunits of the SnRK1 complex in germinating seeds of Medicago truncatula. Plant Cell and Environment 27, 5567.
Caffrey, M., Fonseca, V. and Leopold, A.C. (1988) Lipid–sugar interactions: relevance to anhydrous biology. Plant Physiology 86, 754758.
Chappell, A.S., Scaboo, A.M., Wu, X., Nguyen, H., Pantalone, V.R. and Bilyeu, K.D. (2006) Characterization of the MIPS gene family in Glycine max. Plant Breeding 125, 493500.
Chien, C.T., Lin, T.P., Juo, C.G. and Her, G.R. (1996) Occurrence of a novel galactopinitol and its changes with other non-reducing sugars during development of Leucaena leucocephala seeds. Plant and Cell Physiology 37, 539544.
Chiera, J.M. and Grabau, E.A. (2007) Localization of myo-inositol phosphate synthase (GmMIPS-1) during the early stages of soybean seed development. Journal of Experimental Botany 58, 22612268.
Chiera, J.M., Streeter, J.G. and Finer, J.J. (2006) Ononitol and pinitol production in transgenic soybean containing the inositol methyl transferase gene from Mesembryanthemum crystallinum. Plant Science 171, 647654.
Dittrich, P. and Brandl, A. (1987) Revision of the pathway of d-pinitol formation in Leguminosae. Phytochemistry 26, 19251926.
Ellis, E.C. and Spanswick, R.M. (1987) Sugar efflux from attached seed coats of Glycine max (L.) Merr. Journal of Experimental Botany 38, 14701483.
Gomes, C.I., Obendorf, R.L. and Horbowicz, M. (2005) myo-Inositol, d-chiro-inositol, and d-pinitol synthesis, transport, and galactoside formation in soybean explants. Crop Science 45, 13121319.
Heaney, R.P., Weaver, C.M. and Fitzsimmons, M.L. (1991) Soybean phytate content – effect on calcium absorption. American Journal of Clinical Nutrition 53, 745747.
Hegeman, C.E., Good, L.L. and Grabau, E.A. (2001) Expression of d-myo-inositol-3-phosphate synthase in soybean. Implications for phytic acid biosynthesis. Plant Physiology 125, 19411948.
Hitz, W.D., Carlson, T.J., Kerr, P.S. and Sebastian, S.A. (2002) Biochemical and molecular characterization of a mutation that confers a decreased raffinosaccharide and phytic acid phenotype on soybean seeds. Plant Physiology 128, 650660.
Horbowicz, M. and Obendorf, R.L. (1994) Seed desiccation tolerance and storability: dependence on flatulence-producing oligosaccharides and cyclitols – review and survey. Seed Science Research 4, 385405.
Horbowicz, M., Brenac, P. and Obendorf, R.L. (1998) Fagopyritol B1, O-α-d-galactopyranosyl-(1 → 2)-d-chiro-inositol, a galactosyl cyclitol in maturing buckwheat seeds associated with desiccation tolerance. Planta 205, 111.
Htoo, J.K., Sauer, W.C., Zhang, Y., Cervantes, M., Liao, S.F., Araiza, B.A., Morales, A. and Torrentera, N. (2007) The effect of feeding low-phytate barley–soybean meal diets differing in protein content to growing pigs on the excretion of phosphorus and nitrogen. Journal of Animal Science 85, 700705.
Kosina, S.M., Castillo, A., Schnebly, S.R. and Obendorf, R.L. (2009) Soybean seed coat cup unloading on plants with low-raffinose, low-stachyose seeds. Seed Science Research 19, 145153.
Koster, K.L. and Leopold, A.C. (1988) Sugars and desiccation tolerance in seeds. Plant Physiology 88, 829832.
Lahuta, L.B., Górecki, R.J. and Horbowicz, M. (2005) High concentrations of d-pinitol or d-chiro-inositol inhibit the biosynthesis of raffinose family oligosaccharides in maturing smooth tare (Vicia tetrasperma [L.] Schreb.) seeds. Acta Physiologiae Plantarum 27, 505513.
Loewus, F.A. and Murthy, P.P.N. (2000) myo-Inositol metabolism in plants. Plant Science 150, 119.
Lynch, S.R., Dassenko, S.A., Cook, J.D., Juillerat, M.A. and Hurrell, R.F. (1994) Inhibitory effect of a soybean-protein related moiety on iron-absorption in humans. American Journal of Clinical Nutrition 60, 567572.
Ma, J.M., Horbowicz, M. and Obendorf, R.L. (2005) Cyclitol galactosides in embryos of buckwheat stem–leaf–seed explants fed d-chiro-inositol, myo-inositol, or d-pinitol. Seed Science Research 15, 329338.
Meis, S.J., Fehr, W.R. and Schnebly, S.R. (2003) Seed source effect on field emergence of soybean lines with reduced phytate and raffinose saccharides. Crop Science 43, 13361339.
Mitsuhashi, N., Kondo, M., Nakaune, S., Ohnishi, M., Hayashi, M., Hara-Nishimura, I., Richardson, A., Fukaki, H., Nishimura, M. and Mimura, T. (2008) Localization of myo-inositol-1-phosphate synthase to the endosperm in developing seeds of Arabidopsis. Journal of Experimental Botany 59, 30693076.
Neus, J.D., Fehr, W.R. and Schnebly, S.R. (2005) Agronomic and seed characteristics of soybean with reduced raffinose and stachyose. Crop Science 45, 589592.
Nunes, A.C.S., Vianna, G.R., Cuneo, F., Amaya-Farfán, J., de Capdeville, G., Rech, E.L. and Aragão, F.J.L. (2006) RNAi-mediated silencing of the myo-inositol-1-phosphate synthase gene (GmMIPS1) in transgenic soybean inhibited seed development and reduced phytate content. Planta 224, 125132.
Obendorf, R.L. (1997) Oligosaccharides and galactosyl cyclitols in seed desiccation tolerance (Review Update). Seed Science Research 7, 6374.
Obendorf, R.L., Horbowicz, M., Dickerman, A.M., Brenac, P. and Smith, M.E. (1998) Soluble oligosaccharides and galactosyl cyclitols in maturing soybean seeds in planta and in vitro. Crop Science 38, 7884.
Obendorf, R.L., Odorcic, S., Ueda, T., Coseo, M.P. and Vassallo, E. (2004) Soybean galactinol synthase forms fagopyritol B1 but not galactopinitols: substrate feeding of isolated embryos and heterologous expression. Seed Science Research 14, 321333.
Obendorf, R.L., Sensenig, E.M., Wu, J., Ohashi, M., O'Sullivan, T.E., Kosina, S.M. and Schnebly, S.R. (2008 a) Soluble carbohydrates in mature soybean seed after feeding d-chiro-inositol, myo-inositol, or d-pinitol to stem-leaf-pod explants of low-raffinose, low-stachyose lines. Plant Science 175, 650655.
Obendorf, R.L., Zimmerman, A.D., Ortiz, P.A., Taylor, A.G. and Schnebly, S.R. (2008 b) Imbibitional chilling sensitivity and soluble carbohydrate composition of low raffinose, low stachyose soybean seed. Crop Science 48, 23962403.
Obendorf, R.L., Zimmerman, A.D., Zhang, Q., Castillo, A., Kosina, S.M., Bryant, E.G., Sensenig, E.M., Wu, J. and Schnebly, S.R. (2009) Accumulation of soluble carbohydrates during seed development and maturation of low-raffinose, low-stachyose soybean. Crop Science 49, 329341.
Rosnoblet, C., Aubry, C., Leprince, O., Vu, B.L., Rogniaux, H. and Buitink, J. (2007) The regulatory gamma subunit SNF4b of the sucrose non-fermenting-related kinase complex is involved in longevity and stachyose accumulation during maturation of Medicago truncatula seeds. Plant Journal 51, 4759.
Sebastian, S.A., Kerr, P.S., Pearlstein, R.W. and Hitz, W.D. (2000) Soybean germplasm with novel genes for improved digestibility. pp. 5673in Drackley, J.K. (Ed.) Soy in animal nutrition. Savoy, Illinois, Federation of Animal Science Societies.
Sharpley, A.N., Daniel, T., Sims, T., Lemunyon, J., Stevens, R. and Perry, R. (2003) Agricultural phosphorus and eutrophication (2nd edition). Agricultural Research Service, ARS 149. Washington, DC, United States Department of Agriculture.
Streeter, J.G. (2001) Simple partial purification of d-pinitol from soybean leaves. Crop Science 41, 19851987.
Streeter, J.G., Lohnes, D.G. and Fioritto, R.J. (2001) Patterns of pinitol accumulation in soybean plants and relationships to drought tolerance. Plant Cell and Environment 24, 429438.
Suarez, F.L., Springfield, J., Furne, J.K., Lohrmann, T.T., Kerr, P.S. and Levitt, M.D. (1999) Gas production in humans ingesting a soybean flour derived from beans naturally low in oligosaccharides. American Journal of Clinical Nutrition 69, 135139.
Thorne, J.H. (1980) Kinetics of 14C-photosynthate uptake by developing soybean fruit. Plant Physiology 65, 975979.
Thorne, J.H. (1981) Morphology and ultrastructure of maternal seed tissues of soybean in relation to the import of photosynthate. Plant Physiology 67, 10161025.
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 Physiology 72, 268271.


Free cyclitol unloading from seed coats on stem–leaf–pod explants of low-raffinose, low-stachyose, low-phytin soybean

  • Suzanne M. Kosina (a1), Steven R. Schnebly (a2) and Ralph L. Obendorf (a1)


Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed