Lodging-resistant rice (Oryza sativa) cultivars usually show slow grain filling when nitrogen is applied in large amounts. This study investigated the possibility that a hormonal change may mediate the effect of water deficit that enhances whole plant senescence and speeds up grain filling. Two rice cultivars showing high lodging resistance and slow grain filling were field grown and applied with either normal or high amount nitrogen (HN) at heading. Well-watered and water-stressed (WS) treatments were imposed 9 days post anthesis to maturity. Results showed that WS increased partitioning of fixed 14 CO 2 into grains, accelerated the grain filling rate but shortened the grain filling period, whereas the HN did the opposite way. Cytokinin (zeatin ϩ zeatin riboside) and indole-3-acetic acid contents in the grains transiently increased at early filling stage and WS treatments hastened their declines at the late grain filling stage. Gibberellins (GAs; GA 1 ϩ GA 4 ) in the grains were also high at early grain filling but HN enhanced, whereas WS substantially reduced, its accumulation. Opposite to GAs, abscisic acid (ABA) in the grains was low at early grain filling but WS remarkably enhanced its accumulation. The peak values of ABA were significantly correlated with the maximum grain filling rates (r ϭ 0.92**, P Ͻ 0.01) and the partitioning of fixed 14 C into grains (r ϭ 0.95**, P Ͻ 0.01). Exogenously applied ABA on pot-grown HN rice showed similar results as those by WS. Results suggest that an altered hormonal balance in rice grains by water stress during grain filling, especially a decrease in GAs and an increase in ABA, enhances the remobilization of prestored carbon to the grains and accelerates the grain filling rate.

This study tested the hypothesis that a controlled water deficit during grain filling of wheat (Triticum aestivum) could accelerate grain-filling rate through regulating the key enzymes involved in Suc-to-starch pathway in the grains. Two high lodgingresistant wheat cultivars were field grown. Well-watered and water-deficit (WD) treatments were imposed from 9 DPA until maturity. The WD promoted the reallocation of prefixed 14 C from the stems to grains, shortened the grain-filling period, and increased grain-filling rate or starch accumulation rate (SAR) in the grains. Activities of Suc synthase (SuSase), soluble starch synthase (SSS), and starch branching enzyme (SBE) in the grains were substantially enhanced by WD and positively correlated with the SAR. ADP Glc pyrophosphorylase activity was also enhanced in WD grains initially and correlated with SAR with a smaller coefficient. Activities of granule-bound starch synthase and soluble and insoluble acid invertase in the grains were less affected by WD. Abscisic acid (ABA) content in the grains was remarkably enhanced by WD and very significantly correlated with activities of SuSase, SSS, and SBE. Application of ABA on well-watered plants showed similar results as those by WD. Spraying with fluridone, an ABA synthesis inhibitor, had the opposite effect. The results suggest that increased grainfilling rate is mainly attributed to the enhanced sink activity by regulating key enzymes involved in Suc-to-starch conversion, especially SuSase, SSS, and SBE, in wheat grains when subjected to a mild water deficit during grain filling, and ABA plays a vital role in the regulation of this process.Starch in the endosperm of wheat (Triticum aestivum) is the major form of carbon reserves and comprises 65% to 75% of the final dry weight of the grain (Housley et al., 1981;Dale and Housley, 1986;Hurkman et al., 2003). Grain filling is therefore mainly a process of starch biosynthesis and accumulation. It is generally accepted that four enzymes may play a key role in this process: Suc synthase (SuSase; EC 2.4.1.13), ADP Glc pyrophosphorylase (AGPase; EC 2.7.7.27), starch synthase (StSase; EC 2.4.1.21), and starch branching enzyme (SBE; EC 2.4. 1.18;Hawker and Jenner, 1993; Ahmadi and Baker, 2001;Hurkman et al., 2003). SuSase catalyses the cleavage of Suc, the main transported form of assimilates in wheat plants (Fisher and Gifford, 1986), to form UDP-Glc and Fru, which is thought to be the first step in the Suc-to-starch conversion. Its activity is considered to be linked to sink strength in the developing rice (Oryza sativa) grain and tomato (Lycopersicon esculentum) fruit (Sun et al., 1992;Wang et al., 1993;Kato, 1995). AGPase produces ADP-Glc, the primer of the starch chain (Smith and Denyer, 1992), and is regarded as the rate-limiting enzyme in starch biosynthesis (Preiss, 1988). StSase, composed of both soluble and granule-bound isoforms, elongates the amylose and amylopectin chains (Déjardin et al., 1997). Soluble StSase (SSS) activity is reported to be positively correlate...

nescence promotes remobilization of assimilates are rather obscure. Very little is known about the relation-Remobilization and transfer of the pre-stored food in vegetative ship between these two processes, a surprisingly netissues to the grains in monocarpic plants require the initiation of glected aspect given its importance to yield formation. whole plant senescence. However, mechanisms by which plant senescence promotes remobilization of assimilates are rather obscure. This

The possible regulation of senescence-initiated remobilization of carbon reserves in rice (Oryza sativa L.) by abscisic acid (ABA) and cytokinins was studied using two rice cultivars with high lodging resistance and slow remobilization. The plants were grown in pots and either well-watered (WW, soil water potential = 0 MPa) or water-stressed (WS, soil water potential = -0.05 MPa) from 9 days after anthesis until they reached maturity. Leaf water potentials of both cultivars markedly decreased at midday as a result of water stress but completely recovered by early morning. Chlorophyll (Chl) and photosynthetic rate (Pr) of the flag leaves declined faster in WS plants than in WW plants, indicating that the water deficit enhanced senescence. Water stress accelerated starch remobilization in the stems, promoted the re-allocation of pre-fixed (14)C from the stems to grains, shortened the grain-filling period and increased the grain-filling rate. Sucrose phosphate synthase (SPS, EC 2.4.1.14) activity was enhanced by water stress and positively correlated with sucrose accumulation in both the stem and leaves. Water stress substantially increased ABA but reduced zeatin (Z) + zeatin riboside (ZR) concentrations in the root exudates and leaves. ABA significantly and negatively, while Z+ZR positively, correlated with Pr and Chl of the flag leaves. ABA, not Z+ZR, was positively and significantly correlated with SPS activity and remobilization of pre-stored carbon. Spraying ABA reduced Chl in the flag leaves, and enhanced SPS activity and remobilization of carbon reserves. Spraying kinetin had the opposite effect. The results suggest that both ABA and cytokinins are involved in controlling plant senescence, and an enhanced carbon remobilization is attributed to an elevated ABA level in rice plants subjected to water stress.