The Inheritance of Seed Dormancy in Sinapis Arvensis L
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Heredity 56 (1986) 25—31 The Genetical Society of Great Britain Received 8 March 1985 The inheritance of seed dormancy in Sinapis arvensis L. K. Garbutt* and J. R. Witcombet School of Plant Biology, University College of North Wales, Bangor, Gwynedd, U.K. Selection for dormant and non-dormant seed in Sinapis arvensis was carried to the seventh and fourteenth generation, respectively. Crosses between the dormant and non-dormant lines clearly showed both a maternal and an embryonic component of seed dormancy. A model for the number of alleles controlling dormancy was constructed and tested. The maternal component of dormancy was shown to be controlled by a single locus with two alleles, the dormant allele being dominant to the non-dormant. No clear picture of the control of the embryonic component of dormancy was found. INTRODUCTION shown a clear response to selection for non- dormancy in several populations. They also Therehave been several reports of genetic control demonstrated a clear difference in germinability of factors affecting seed dormancy in plants. between threshed and unthreshed grains, indicat- Honing (1930) showed a genetic basis and a dis- ing the presence of inhibitors in the lemma and tinct maternal effect in the light dependence of palea, and hence a maternal component to control germination behaviour in tobacco. Harper and of dormancy. McNaughton (1960) found evidence of genetic There have been two recent attempts to estimate control of seed dormancy in species of Papaver. the number of loci involved in the control of germi- New (1958; 1959; 1961) has shown a strict genetic nation. Guenther and Borriss (1975) tried unsuc- control of seed coat types associated with differing cessfully to estimate the number of loci controlling germination rates in Spergula arvensis indicating seed dormancy in Antirrhinum spp. concluding that a strict maternal control, via the seed coat, over dormancy was controlled by a polygenic complex. seed dormancy. However, Jana, Acharya and Naylor (1979) have Naylor and his co-workers have shown genetic shown that at least three loci control after-ripening variability in seed dormancy in Avena fatua in in Avena fatua. response to several environmental variables In the field, Sinapis arvensis populations applied both whilst the seed is maturing and during usually show two "flushes" of germination each germination tests (Naylor and Jana, 1976; Naylor year, one in spring and the other in autumn. and Fedec, 1978; Sawhney and Naylor, 1979). Jam Edwards (1980) suggests that the autumnal popu- (1982) has also demonstrated genetic components lation results from the non-dormant fraction of the of seed dormancy for Arena barbata and Tr(folium seed produced in the spring. In the field, S. arvensis hirtum but in the same study was unable to demon- germination is controlled to some extent by strate a significant genetic component of dormancy environmental factors. Edwards (1980) found in Bromus mollis. germination to be associated with soil temperatures Hilu and de Wet (1980) have investigated the of 44°C at a depth of 10 cm and with high rainfall genetic basis of dormancy in Eleusine sp. and have in spring and simply with adequate moisture in autumn. There is also evidence that at least some Present addresses: *TheBiological Laboratories, Harvard Uni- seed of S. arvensis require light to germinate versity, 16 Divinity Avenue, Cambridge, MA 02138. (Frankland, 1976). In the laboratory, dormancy is t ICRISAT, Pantancheru P.O., Andhra Pradesh 502 324 India. broken by the application of gibberellic acid 26 K. GARBU1T AND J. R. WITCOMBE (Witcombe and Whittington, 1972; Edwards, the parent plants has a seed coat of generation P1 1976). and embryo of generation pe,andthe plants grown The presence of a germination inhibitor and its from this seed will be generation peSeedfrom a mode of action in seeds of S. arvensis was reported cross between two plants of pe generation will in Edwards (1968c). This inhibitor is apparently have Sgenerationseed coat and F generation produced by the seed coat (testa) (Witcombe, embryo. Hiliman and Whittington, 1969) and seems to be All plants were grown in a greenhouse in 10 cm present in the seed coat of all seeds in apparently plastic pots filled with John Innes No. 1 compost. similar quantities (see Edwards, 1968a for a Supplementary light to give a daylength of 16 hours detailed description of seed anatomy). The inhibi- minimum was provided by mercury vapour lamps tion is broken down by oxidation (Edwards, 1969) and temperature was maintained at a minimum of and the effect of the seed coat in dormant seed 18°C. Treatments of fungicide and insecticide were appears to be to prevent this oxidation taking place applied as necessary. by being impermeable to oxygen (Edwards, All pollinations were made in the same manner 1968b). The seed coat is clearly of maternal origin whether within or between lines. Dehiscing anthers and hence control of dormancy is maternal. There were removed from the male parent and placed in is evidence, however, of a further level of dor- a watch glass; a flower bud about to open, but not mancy control, possibly unrelated to this maternal yet showing the stigmatic surface was opened, and effect, and located in the embryo (Edwards, 1968c; emasculated and the newly exposed stigmatic sur- Witcombe, 1971). This points to one of the major face loaded with pollen from the anthers of the problems in studies of the inheritance of seed male parent. The pedicel was marked with paint dormancy. The seed combines parts of three dis- and 24 hours later a further application of pollen tinct generations (and genotypes)—the seed coat was made. (maternal), the endosperm (filial but with one These crosses produced generationseed, paternal and a variable number of maternal 200 individuals of which were grown on and genomes) and the embryo (filial). In the ripe seed allowed to cross at random to produce F7F of Cruciferae, however, the endosperm has disap- generation and seed 200 individuals of which, peared leaving a seed composed of two generations when grown on and crossed, produced FF seed. of tissue, seed coat and embryo. A further set of crosses was made between pe The purpose of the following investigation was plants. These comprised 50 crosses between dor- to determine the manner in which dormancy was mant males and females, 50 crosses between non- controlled genetically in Sinapis arvensis. dormant males and females, and 50 crosses between dormant males and non-dormant females. This was to confirm that the differences observed MATERIALSAND METHODS between the germination rates of seed from non- dormant maternal parents from the p1pe and the Inorder to establish the genetic basis of seed PF generations were, in fact, real and not a dormancy, two selection lines, one for dormancy function of a random difference between genera- (D) and one for non-dormancy (N.D.), were estab- tions. lished from a natural population collected in At each generation, pspePSFC Nottinghamshire by J. R. Witcombe (Witcombe germination tests were carried out on the seed in and Whittington 1972). These lines were taken to closed petri dishes each containing one sheet of the SlO generation for the ND. line and the S5 9.0 cm filter paper. Initially each dish received 5 ml generation for the D line by J. R. Witcombe and of distilled water. Thereafter, all dishes received K. Daniels. The present authors continued distilled water as necessary. The germination tests selection using the same methods as Witcombe were carried out at 25°C, with 24 hour light. The and Whittington (1972) and Witcombe (1971) to lids of the petri dishes were lifted daily to prevent the S14 generation for the N.D. line and S7 the build-up of gases which might inhibit germina- generation for the D line. At this point, 200 tion. Each dish contained 10 seeds unless otherwise reciprocal pairs of crosses were made between the stated. lines and seed produced to the third filial At the conclusion of each test (14 days), the generation. seeds that remained ungerminated were treated As mentioned in the Introduction, the seed is with high levels of G.A. (up to 3000 ig/Iitre) to made up of two distinct generations. Thus a special stimulate dormant seeds into germination and notation will be used. The seed which produces thereby detect non-viable seed. Gibberellic acid INHERITANCE OF SEED DORMANCY 27 was applied in an aqueous solution at pH 7. All generation S7 and then tended to reach a plateau, G.A. solutions were made up freshly before an though with a continued slight increase in germina- experiment and were used immediately. tion and decline in standard error (calculated from With the exception of FF bulk germination arcsine transformation) between S7 and S14 tests were performed (no record of individual plant (fig. 1). performance being kept). The results of the bulk Table 1 gives the mean percentage germination tests were based upon as many seeds as possible and 95 per cent confidence limits for the two lines (between 1000 and 2000 seeds) with only a small at the last generation of selection and the control sample being kept in reserve. Confidence intervals population. The N.D. line is clearly different from were obtained using tables from Rohif and Sokal the others. The difference between the control and (1981), when more than 1000 seeds were used the the D line is small (OO5>p>O0l) from a t-test limits for a sample of 1000 were used as a conserva- on arcsine transformed data. tive estimate. Tests of FF seed were based on Neither 100 per cent germination in the absence individual plants, five petri dishes of 20 seeds being of G.A., nor 0 per cent germination could be set up for each of 100 FF plants.