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Journal of Experimental Botany, Vol. 53, No. 374, pp. 1569-1574, July 1, 2002
© 2002 Oxford University Press

Effect of grain colour gene (R) on grain dormancy and sensitivity of the embryo to abscisic acid (ABA) in wheat

Received 18 June 2001; Accepted 15 March 2002

Eiko Himi1, Daryl J. Mares2, Akira Yanagisawa3 and Kazuhiko Noda4,1

1 Research Institute for Bioresources, Okayama University, 2-20-1 Chuo, Kurashiki, Okayama, 710-0046, Japan
2 Plant Breeding Institute, University of Sydney, PO Box 219, Narrabri, NSW 2390, Australia
3 Kitami Agricultural Experimental Station, 52 Yayoi, Kunneppu, Tokoro-gun, Hokkaido, 099-1496, Japan

4 To whom correspondence should be addressed. Fax: +81 86 434 1249. E-mail: knoda{at}rib.okayama-u.ac.jp


   Abstract
Top
 Abstract
Introduction
 Materials and methods
 Results
 Discussion
 References
 
The level of grain dormancy and sensitivity to ABA of the embryo, a key factor in grain dormancy, were examined in developing grains of a white-grained wheat line, Novosibirskaya 67 (NS-67), and its red-grained near-isogenic lines (ANK-1A to -1D); a red-grained line, AUS 1490, and its white-grained mutant line (EMS-AUS). ANK lines showed higher levels of grain dormancy than NS-67 at harvest maturity. AUS 1490 grain also showed higher dormancy than EMS-AUS grain. These results suggest that the R gene for grain colour can enhance grain dormancy. However, the dormancy effect conferred by the R gene was not large, suggesting that it plays a minor role in the development of grain dormancy. Water extracts of AUS 1490 and EMS-AUS bran contained germination inhibitors equivalent to 1–10 µM ABA, although there was no difference in the amount of inhibitors between AUS 1490 and EMS-AUS. Thus, the grain colour gene of AUS 1490 did not appear to enhance the level of grain dormancy by accumulating germination inhibitors in its bran. Sensitivity to ABA of embryos was higher in grains collected around harvest-maturity for ANK lines and AUS 1490, compared with NS-67 and EMS-AUS. The R gene might enhance grain dormancy by increasing the sensitivity of embryos to ABA.

Key words: Key words: ABA sensitivity of embryo, germination inhibitor, grain colour, grain dormancy, wheat.


   Introduction
Top
 Abstract
 Introduction
Materials and methods
 Results
 Discussion
 References
 
Grain dormancy is an important mechanism that enables the grain to remain quiescent until conditions become favourable for germination. In wheat, grain with a low level of dormancy may germinate on plants before harvest (preharvest sprouting) under wet weather conditions, which can cause a reduction in the quality of the flour produced.

Anderson et al. (1993) and Sorrells and Anderson (1996) reported that several RFLP markers for resistance to preharvest sprouting were located on chromosomes of groups 1, 2, 3, 4, 5, and 6 using quantitative trait loci (QTL) analysis of wheat. These results suggest that many genes are involved in grain dormancy.

Physiological studies on seed or grain dormancy of Arabidopsis, maize, and wheat has shown that embryo sensitivity to ABA plays a key role in the dormancy mechanism (Koornneef et al., 1984; McCarty, 1995; Walker-Simmons, 1987; Kawakami et al., 1997). Several genes for ABA sensitivity such as ABI3, ABI5 and Vp1, which were expressed mainly in seeds, have been isolated in Arabidopsis and maize (McCarty, 1995; Giraudat, 1995; Finkelstein et al., 1998; Finkelstein and Lynch, 2000). Interestingly, Maize Vp1 was also reported to be involved in flavonoid synthesis and grain maturation besides the sensitivity to ABA (McCarty, 1995). The pigment contributing to wheat grain colour (phlobaphene) is synthesized through the pathway of flavonoid synthesis (Grotewold et al., 1994; Miyamoto and Everson, 1958). Bailey et al. (1999) identified the taVp1 gene, an orthologue of Vp1, and located it in a region 30 cM from the R locus, which controls wheat grain colour and is located in the distal region of the long arm of wheat chromosome 3. McKibbin et al. (1999) suggested, however, that taVp1 expression in the embryo during its development might not be associated with the level of dormancy of wheat grain.

It is well known that red-grained wheats show a wider variation in grain dormancy than white-grained wheats. The level of grain dormancy in white-grained wheats does not exceed that in red-grained wheats, suggesting that red-grained wheats carry gene(s) for grain dormancy additional to the dormancy gene(s) in white-grained wheats (Flintham, 1993). A dormancy gene was found on chromosome 3 (Flintham et al., 1999; Mares, 1996). To elucidate further whether the R gene itself or some gene linked to the R locus affects dormancy, Flintham (2000) examined the dormancy level of after-ripened grains of the white-grained NS-67 and its near-isogenic lines carrying a single R gene on one of chromosomes 3A, 3B, and 3D. He concluded that the R gene on chromosome 3 increased levels of grain dormancy. Warner et al. (2000), using harvest-ripe grain of white-grained mutants of cv. Chinese Spring, also showed that the R gene enhanced grain dormancy. The function of the R gene on grain dormancy has been suggested to be to accumulate germination inhibitors since a water-soluble precursor of the red pigment, catechin, could inhibit grain germination (Miyamoto and Everson, 1958; Stoy and Sundin, 1976).

In the present report, embryo sensitivity to ABA was compared in developing grains of NS-67 and its near-isogenic lines for the R gene and a white-grained mutant line, EMS-AUS, derived from a red-grained wheat line, AUS 1490. The inhibitory effect of a water-soluble extract from bran on embryo germination was also examined.


   Materials and methods
Top
 Abstract
 Introduction
 Materials and methods
Results
 Discussion
 References
 
Common wheat (Triticum aestivum L.) line, Novosibirskaya-67 (NS-67, white grained, grain colour genotype R-A1a/R-B1a/R-D1a), and four ninth-backcross near-isogenic lines (ANK-1A, ANK-1B, ANK-1C, and ANK-1D) of NS-67 carrying a single R gene of wheat cultivars, Arin (R-A1a/R-B1a/R-D1b), k-28535 (R-A1a/R-B1b/R-D1a), Chinese Spring (CS, R-A1a/R-B1a/R-D1b), and Solo (R-A1b/R-B1a/R-D1a), respectively, were a kind gift of Dr A Vershinin of the Russian Academy of Sciences and provided by Dr J Flintham, John Innes Centre, Norwich, UK (Koval, 1997). Other lines used were EMS-AUS 1490 (EMS-AUS), a white-grained mutant line derived from the red-grained dormant line, AUS 1490. AUS 1490 carries a single red gene, although its grain colour genotype is unknown (Mares, 1999). These lines, a red-grained line, Kitakei 1354, grains of which were dormant and ABA-sensitive (Kawakami et al., 1997), and CS (an R gene donor to ANK-1C), were grown under a transparent vinyl roof in a field in Kurashiki. The average temperature and rainfall during grain filling period were 22.9 °C and 245.5 mm.

At anthesis, spikes were tagged and grains collected from the primary and secondary florets of the central spikelets at 5 d intervals after anthesis. Dry grain weight was measured after incubation of 100 grains at 150 °C for 3 h. Water content (%) of grains was estimated by (fresh weight–dry weight)/fresh weight. Twenty fresh grains or half grains containing embryo (embryo-half grains) were incubated in a Petri dish (9 cm in diameter) containing two pieces of filter paper (8.4 cm in diameter, filter No. 2, Toyo) moistened with 6 ml of distilled water or 10 µM aqueous solution (pH 5.7) of abscisic acid (ABA, Cat No. A-7383, Sigma) at 20 °C for 7 d. Germinated grains and embryo-half grains were counted daily. Germination was defined as pericarp rupture over the embryo. For estimating the germinability of grains and embryos, a germination index (GI) was calculated as follows.

where n1, n2, ... n7 are the number of grains or embryos that germinated on the first, second and subsequent days until the seventh day, respectively.

An aqueous extract from bran was prepared from mature AUS 1490 and EMS-AUS grains ground in a Quadrumat Junior Brabender experimental mill. The bran fraction was sieved (sieve size 231 µm) to remove the remaining endosperm, and 5 g of bran was mixed with 30 ml of distilled water and shaken with a shaker (Taitec bio-shaker, Japan) at 205 rpm and room temperature for 2 h. After centrifugation at 1000 g for 10 min, the supernatant was filtered through filter paper (Advantec No. 2) and a 0.45 µm disc filter (Steradisc 25). Twenty embryo-half grains from fresh and mature Kitakei 1354 grains were incubated in 6 ml of these extracts, or for comparison, in water, 1 µM, 10 µM or 50 µM ABA solutions at 20 °C for 7 d.

All experiments were done in triplicate. Water contents (%) of grain and GI values were transformed to arcsin{surd}. Dry weight, water content and GI values of grains and embryos were tested statistically by Duncan’s multiple range test.


   Results
Top
 Abstract
 Introduction
 Materials and methods
 Results
Discussion
 References
 
NS-67, ANK-1C, and CS
Dry weight and water content of developing grains of NS-67, ANK-1C, and CS, which was donor of the R gene to ANK-1C, were measured (Fig. 1A). No significant difference was observed in dry weight and water content among these lines. Water content decreased rapidly 30 d after pollination (DAP) and reached a minimum around 15% at 40 DAP. According to the classification of wheat grain development (Rogers and Quatrano, 1983; Noda et al., 1994), these grains appear to have reached harvest-maturity at 40 DAP and after that enter the after-ripening stage.



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  		Fig. 1. Changes during the grain development of Novosibirskaya-67 (NS-67), its near isogenic line, ANK-1C and Chinese Spring (CS). (A) Assessments of grain development (dry seed weight and water content); (B) GI of developing grains incubated with water at 20 °C for 7 d; (C) GI of embryo-half grains incubated with water or 10 µM ABA. Bars=±standard error.

 
Germinability of the developing grains was examined after imbibing for 7 d (Fig. 1B). Grains of NS-67 and CS first showed germination at 30 DAP and thereafter their germinability increased. GI values of NS-67 and CS were 25.2 and 20.5 at 40 DAP, respectively. These grains partly lost their dormancy at this stage. On the other hand, grains of ANK-1C did not germinate until after 40 DAP, and even at 45 DAP the GI of ANK-1C grains was only 5.2. The GIs of NS-67 and CS were significantly (5% level) higher than that of ANK-1C from 35–45 DAP. These results showed that grains of ANK-1C around the harvest-ripe stage were more dormant than those of its parental lines, NS-67 and CS. However, the dormancy level of ANK-1C did not appear to be high since the dormancy dissipated at 50 DAP.

Embryo-half grains of NS-67, CS and ANK-1C, when incubated in water for 7 d, germinated well after 35 DAP (Fig. 1C). The germinability of embryos appeared to develop more slowly in ANK-1C than in NS-67 and CS. The GI of embryos of ANK-1C was significantly lower at the 5% level than those of NS-67 and CS from 35–40 DAP (Fig. 1C). In an ABA solution, germination of the three lines are suppressed throughout their grain development (Fig. 1C). By comparison of ABA suppression among these lines, embryo-half grains of ANK-1C appears to exhibit higher sensitivity to ABA than those of CS and ANK-1C from DAP 35–45.

The other ANK lines
Development and germinability of whole grains and germinability of embryo-half grains in water and ABA were also examined in the three near-isogenic lines (ANK-1A, -1B, and -1D), which carried a single R gene from different red-grained wheat lines. Unfortunately, the three R gene donor lines were not available in the present experiments. There was no significant difference (at the 5% level) in their grain development until 45 DAP (Fig. 2A). Grains of all three ANK lines were significantly more dormant (at the 5% level) than those of NS-67 from 35–40 DAP (Fig. 2B). At 45 DAP, the GI value of ANK-1B grains (21.0) was significantly (at the 5% level) lower than GI values of NS-67 (51.4), ANK-1A (53.3), and ANK-1D (40.0) grains, suggesting that ANK-1B grains were more dormant than the other lines’ grains. Sensitivity of the embryos of the ANK lines to ABA appeared to be higher than that of NS-67 between 35 and 45 DAP (Fig. 2C). Among the embryos of ANK lines, ANK-1B embryos showed higher sensitivity to ABA than the other lines’ embryos.



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  		Fig. 2. Changes during the grain development of Novosibirskaya-67 (NS-67), and three near isogenic lines (ANK-1A, –1B and –1D). (A) Assessments of grain development (dry seed weight and water content); (B) GI of developing grains incubated with water at 20 °C for 7 d; (C) GI of embryo-half grains incubated with water or 10 µM ABA. Bars=±standard error.

 
Grain colour mutant
Water contents of AUS 1490 and EMS-AUS grains decreased to about 15% at 45 DAP (Fig. 3A). At 40 DAP the germinability of AUS 1490 and EMS-AUS grains increased (Fig. 3B). However, the GI of AUS 1490 grains decreased after 40 DAP while those of EMS-AUS maintained a GI of about 20 (Fig. 3B). The results show that EMS-AUS grains are less dormant than AUS 1490 grains at and after harvest maturity. However, the release of dormancy in white-grained mutant EMS-AUS was only slight. With respect to the development of embryo germinability (Fig. 3C), embryos of AUS 1490 and EMS-AUS germinated well at 40 DAP with no significant difference between them. The sensitivity of EMS-AUS embryos to ABA appeared to be lower than that of AUS 1490 embryos from 40–55 DAP except at 50 DAP (Fig. 3C).



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  		Fig. 3. Changes during the grain development of AUS 1490 and EMS-AUS. (A) Assessments of grain development (dry seed weight and water content); (B) GI of developing grains incubated with water at 20 °C for 7 d; (C) GI of embryo-half grains incubated with water or 10 µM ABA. Bars=±standard error.

 
Inhibition of germination by bran extract
The effect of water-soluble extracts from bran of AUS 1490 and EMS-AUS on germination was examined in Kitakei 1354 embryos with high sensitivity to ABA (Fig. 4). The extracts from AUS 1490 and EMS-AUS significantly (at the 1% level) inhibited the germination of Kitakei 1345 embryos, compared with the germination of Kitakei 1354 embryos in water. The effects of the AUS 1490 and EMS-AUS extracts were equivalent to 1–10 µM ABA. However, there was no statistically significant difference in the degree of germination inhibition by the extracts of AUS 1490 and EMS-AUS.



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  		Fig. 4. GI of embryo-half grains of dormant Kitakei 1354, when incubated with water, 1, 10 and 50 µM ABA, and water-soluble extracts taken from the bran of AUS 1490 and EMS-AUS brans. Bars=±standard error.

 

   Discussion
Top
 Abstract
 Introduction
 Materials and methods
 Results
 Discussion
References
 
Grains of the near-isogenic line ANK-1C (red-grained) were significantly more dormant than those of its parental lines, NS-67 and CS, from 35–45 DAP (Fig. 1). The other ANK lines (red-grained) also showed significantly higher levels of dormancy than NS-67 (white-grained) around the phase of grain maturity (Fig. 2). These results indicated that the R gene (for grain colour) and/or genes linked tightly to the R locus could confer grain dormancy to NS-67. EMS-AUS (white) grains were also less dormant than AUS 1490 (red) grains (Fig. 3B). Taken together, the results agree with those of Flintham (2000) and Warner et al. (2000) who found that the R gene enhances grain dormancy. However, the level of dormancy conferred by the R gene decreased rapidly in ANK lines during after-ripening and the reduction of dormancy level in white-grained mutant EMS-AUS was not large. The R gene might therefore act only as a minor factor in dormancy.

Among the ANK lines, grains of ANK-1C and ANK-1B appear to be more dormant than those of ANK-1A and -1B (Figs 1, 2). Koval (1997) showed that these four ANK lines were genetically different in several characteristics such as grain size and flour technological properties, suggesting that some genes located near the R locus were different in these ANK lines. Flintham et al. (1999) reported that wheat has dormancy genes which do not affect grain colour and are located on chromosome 3. The present results also suggest that additional dormancy genes might be located near the R locus.

The function of the R gene in relation to grain dormancy has been speculated to be to accumulate germination inhibitors in the grain coat tissue (Miyamoto and Everson, 1958). Stoy and Sundin (1976) reported that water-soluble precursors, catechins and catechin-tannins, of the red pigment phlobaphene in wheat grains, could inhibit embryo germination. The present results showed that water-soluble extracts from AUS 1490 and EMS-AUS bran did contain inhibitors of embryo germination, but there was no significant difference in the amount of inhibitors between AUS 1490 and EMS-AUS (Fig. 4) even though these lines showed differences in dormancy.

The rate of imbibition has been reported to be slower in coloured seeds than in white seeds of legumes and wheat (Huang et al., 1983; Powell, 1989). Huang et al. (1983) suggested that the rate of imbibition might be one of the factors that affect grain dormancy of wheat. However, the difference in water uptake was observed only within 24 h of imbibition between red and white grains (Huang et al., 1983). Unpublished data from this laboratory show that the water content of the embryos of dormant red wheat grains, when incubated in water at 20 °C, reached a plateau (56%) within 24 h of imbibition. Difference in the rate of imbibition between red and white wheat grains appears to occur only in the very early phase of imbibition. On the other hand, germination percentages of the grains of the ANK-1B and -1C collected at 45 DAP (20% and 40%, respectively) were significantly lower than the germination percentage of NS-67 (93.3%) at day 7 after incubation in water. The difference in the rate of imbibition may not explain increased dormancy level of the ANK lines observed at day 7.

It was shown that the sensitivity to ABA of embryos could be altered irrespective of the presence and absence of the R gene (Walker-Simmons, 1987; Kawakami et al., 1997). Warner et al. (2000) observed a small difference in the ABA sensitivity of harvest-ripe grains between CS and its white-grain mutants, but no difference in their after-ripening grains. They concluded that the R gene was not involved in the development of ABA sensitivity. No difference in ABA sensitivity was observed in after-ripening grains of red-grained and white-grained wheat lines. Nevertheless, the results suggest that the sensitivity to ABA of embryos of ANK lines and AUS 1490 was higher than that of NS-67, CS or EMS-AUS between 35 and 45 DAP. It is possible that the R gene might affect the sensitivity to ABA of embryos.

Walker-Simmons showed that the sensitivity to ABA of wheat embryos decreased during grain development (Walker-Simmons, 1987). Higher ABA sensitivity observed in ANK lines around harvest maturity might be a reflection of their slow development of embryo germinability (Figs 1C, 2C). However, in AUS 1490 and EMS-AUS there was no difference in the development of embryo germinability (Fig. 3C). The results suggest that the R gene is not responsible for the development of embryo germinability, which may be controlled by gene(s) linked tightly to the R gene.


   Acknowledgement
 
This work was supported in part by a grant from the Ohara Foundation for Agricultural Research.


   References
Top
 Abstract
 Introduction
 Materials and methods
 Results
 Discussion
 References
 
Anderson JA, Sorrells ME, Tanksley SD. 1993. Molecular markers for pre-harvest sprouting resistance in wheat. In: Walker-Simmons MK, Reed JL, eds. Pre-harvest sprouting in cereals 1992. St Paul, Minnesota, USA: American Association of Cereal Chemists, 121–128.

Bailey PC, McKibbin RS, Lenton JR, Holdsworth MJ, Flintham JE, Gale MD. 1999. Genetic map locations for orthologous Vp1 genes in wheat and rice. Theoretical and Applied Genetics 98, 280–284.

Finkelstein RR, Lynch TJ. 2000. The Arabidopsis abscisic acid response gene ABI5 encodes a basic leucine zipper transcription factor. The Plant Cell 12, 599–609.[Abstract/Free Full Text]

Finkelstein RR, Wang ML, Lynch TJ, Rao S, Goodman HM. 1998. The Arabidopsis abscisic acid response locus ABI4 encodes an apetala 2 domain protein. The Plant Cell 10, 1043–1054.[Abstract/Free Full Text]

Flintham JE. 1993. Grain colour and sprout-resistance in wheat. In: Walker-Simmons MK, Reed JL, eds. Pre-harvest sprouting in cereals 1992. St Paul, Minnesota, USA: American Association of Cereal Chemists, 30–36.

Flintham JE. 2000. Different genetic components control coat-imposed and embryo-imposed dormancy in wheat. Grain Science Research 10, 43–50.

Flintham JE, Adlam R, Gale MD. 1999. Grain coat and embryo dormancy in wheat. In: Weipert D, ed. Eighth international symposium on pre-harvest sprouting in cereals 1998. Detmold, Germany: Association of Cereal Research, 67–76.

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Mares DJ. 1996. Dormancy in white wheat—mechanism and location of genes. In: Noda K, Mares DJ, eds. Pre-harvest sprouting in cereals 1995. Osaka, Japan: Center for Academic Societies Japan, 179–184.

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