土壤水分亏缺对水稻茎秆贮藏碳水化合物向籽粒运转的调节

doi:10.17521/cjpe.2005.0109

摘要/Abstract

摘要：

以两个茎秆贮藏物质利用效率不同的水稻(Oryza sativa)杂交组合(‘汕优63’和‘Pc311/早献党’)为材料,进行土壤水分亏缺处理(Water-deficit),以水层灌溉为对照(Well-watered),研究水分亏缺对水稻茎贮藏性碳水化合物运转及其关键酶活性的调节作用。结果表明,水分亏缺促进了水稻茎秆贮藏物质的运转和对籽粒产量的贡献,开花前茎秆贮藏的碳水化合物对产量贡献率分别提高了1.9~3.0倍(与水层灌溉相比)。土壤水分亏缺诱导了水稻茎节间α-淀粉酶、β-淀粉酶、α-葡萄糖苷酶、D-酶活性上升,但淀粉磷酸化酶受到了抑制,说明土壤水分亏缺加强水稻茎秆贮藏淀粉水解途径,而不是磷酸解途径。就蔗糖代谢而言,土壤水分亏缺提高了蔗糖磷酸合成酶的活性和活化状态,抑制蔗糖转化酶活性,促进蔗糖合成,加速贮藏物质快速降解和转移,从而调节稻株贮藏碳水化合物向籽粒的分配。

关键词: 水分亏缺, 非结构性糖, 淀粉水解酶, 蔗糖磷酸合成酶, 水稻

Abstract:

Grain yields of rice in China are often low and vary substantially due to the shortages of available water. Assimilates stored prior to grain filling have been identified as important contributors to grain yield in drought environments, but evaluating their benefit has been hampered by an inadequate understanding of the physiological mechanism of remobilization to grains of stored carbon reserves during grain filling. A moderate soil drying applied at grain filling period can enhance plant senescence and may improve the yield through remobilizing more pre-stored food to grains. The study was designed to test the following hypothesis: if soil drying is controlled properly at the mid-late stage of grain filling, an early senescence induced by drought stress would accelerate the rate of grain filling by enhanced relocation of carbon stored reserves, and improved use of pre-stored carbon reserves may increase yields where photosynthetic assimilation is decreased by soil drying. Further evaluation of enzyme regulation in the remobilization process will help to understand the physiological mechanisms.
In the current study, two rice combinations (cvs 'Shanyou63' and 'Pc311/Zao') were grown in cement containers and pots at Yangzhou University. Controlled soil water deficit were imposed at 9 d after anthesis in consideration that the division of endosperm cell is sensitive to water deficit.
We found that, compared to well-watered treatments, the remobilization of stored carbohydrates in the rice stem was significantly enhanced under water-deficit conditions, and the increased contribution to grain yields was 1.9-3.0 times as that of rice grown under well-watered conditions. Pronounced effects of water-deficit treatment on carbohydrate metabolism were observed in rice stems. Sugar concentrations and carbon distribution among sugar components were altered under water-deficit conditions. Starch breakdown in rice stems under soil water deficit occurred through action via the hydrolytic pathway: α-amylase, β-amylase, α-glucosidase and D-enzyme were induced and enhanced by soil drying. Starch phosphorylase activity was decreased under water-deficit conditions, indicating that phosphorylase was not involved in the processes of starch remobilization. Water deficit markedly altered the pattern of sucrose metabolism by shutting down the bypass of carbon flow through the sucrose synthase (SuSy) catalyzed system and enhanced the unidirectional flow through the irreversible sucrose-phosphate synthase (SPS) and acid invertase (AINV) catalyzed pathway. SPS was induced and activated by water deficit, and played a pivotal role in enhancing synthesis of sucrose through the conversion of stored carbon into sucrose. During rapid remobilization of stored sugar, the sucrose content was increased by 10.3% in 'Shanyou63' and 20.45% in 'Pc/Zao' as compared to the well-watered treatments. It is concluded that the enhanced remobilization of stored non-structural carbohydrates (NSC) in rice stems under soil water deficit was due to induced hydrolytic enzyme activities, increased SPS activity and activation state, but decreased invertase activity.

Key words: Water deficit, Non-structural carbohydrate, Starch-hydrolytic enzymes, Sucrose-phosphate synthase, Rice

王维, 蔡一霞, 蔡昆争, 张建华, 杨建昌, 朱庆森. 土壤水分亏缺对水稻茎秆贮藏碳水化合物向籽粒运转的调节. 植物生态学报, 2005, 29(5): 819-828. DOI: 10.17521/cjpe.2005.0109

WANG Wei, CAI Yi-Xia, CAI Kun-Zheng, ZHANG Jian-Hua, YANG Jian-Chang, ZHU Qing-Sen. REGULATION OF SOIL WATER DEFICITS ON STEM-STORED CARBOHYDRATE REMOBILIZATION TO GRAINS OF RICE. Chinese Journal of Plant Ecology, 2005, 29(5): 819-828. DOI: 10.17521/cjpe.2005.0109

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图/表 8

图1 水分亏缺对水稻植株顶叶水势变化的影响(池栽试验) 结果为5次重复的平均值±SD

Fig.1 Changes in leaf water potential of rice cultivars cvs 'Shanyou63' (A) and 'Pc/zao' (B), which were subjected to water deficit from 9 d after anthesis Values are means ± SD of five replications

图2 水分亏缺对水稻茎非结构性糖含量变化的影响(池栽试验)

Fig.2 Concentration and composition of NSC in the stems of two rice cultivars cvs ‘Shanyou63’ (A, C, E, G, I) and ‘Pc/Zao’ (B, D, F, H, J), which were subjected to water deficit from 9 d after anthesis

图3 水分亏缺对水稻茎中淀粉和可溶性糖分配入的14C的影响(盆栽试验) 图例同图1 Legends see Fig.1 图注同图2 Notes see Fig.2

Fig.3 14C NSC in the stems of two rice cultivars cvs ‘Shanyou63’ (A, C) and ‘Pc/Zao’ (B, D), which were subjected to water deficit from 9 d after anthesis

图4 水分亏缺对水稻茎贮藏淀粉水解酶活性的影响(池栽试验) 图例同图1 Legends see Fig.1 图注同图2 Notes see Fig.2

Fig.4 Changes in activities of starch degradation enzymes and in the stems of two rice cultivars ‘Shanyou63’ (A, C, E, G) and ‘Pc/Zao’ (B, D, F, H), which were subjected to water deficit from 9 d after anthesis

图5 水分亏缺对水稻茎淀粉磷酸化酶活性的影响(池栽试验) 图例同图1

Fig.5 Changes in activities of starch phosphorylase in the stems of two rice cultivars 'Shanyou63' (A) and 'Pc/Zao' (B), Which were subjected to water deficit from 9 d after anthesis Legends see Fig.1

图6 水分亏缺对水稻茎蔗糖代谢酶活性的影响(池栽试验) 图例同图1

Fig.6 Changes in activities of sucrose-metabolizing enzymes in the stems of two rice cultivars ‘Shanyou63’ (A, C, E) and ‘Pc/Zao’ (B, D, F), which were subjected to water deficit from 9 d after anthesis Legends see Fig.1

表1 水分亏缺对水稻植株各器官中14C再分配的影响(盆栽试验)

Table 1 Effect of post-anthesis water deficit on the distribution of prefixed 14C in plants of two rice cultivars (cvs 'Shanyou63' and 'Pc/Zao')

品种 Cultivars	处理 Treatments	从花后9天到成熟期转运的¹⁴C量 Remobiled ¹⁴C from 9 d anthesis to harvest			成熟期籽粒中¹⁴C含量 Recovered ¹⁴C in grain at harvest stage (kBq)	呼吸损失¹⁴C Loss of respiration (%)
品种 Cultivars	处理 Treatments	叶Leaves (kBq)	茎秆Stem and sheath (kBq)	颖壳 Glumes (kBq)	成熟期籽粒中¹⁴C含量 Recovered ¹⁴C in grain at harvest stage (kBq)	呼吸损失¹⁴C Loss of respiration (%)
‘汕优63’‘Shanyou63’	水层灌溉Well-watered	17.7^b	136.4^b	7.9^b	120.7^b	25.5^b
	水分亏缺Water-deficit	22.4^a	234.9^a	9.7^a	207.4^a	22.3^a
‘Pc/早’‘Pc/Zao’	水层灌溉Well-watered	15.5^b	122.3^b	9.7^b	102.54^b	30.5^b
	水分亏缺Water-deficit	24.9^a	236.3^a	13.0^a	203.6^a	25.7^a

表2 水分亏缺对水稻茎秆碳贮藏物质的输出及籽粒产量的影响(池栽试验)

Table 2 Effect of post-anthesis water deficit on remobilization of stored reserves and grain yield of two rice cultivars (cvs ‘Shanyou63’ and ‘Pc/Zao’)

品种 Cultivars	处理 Treatments	茎中滞留糖 Residue NSC (mg·g^-1 DW)	运转率 Remobilized NSC (%)	贡献率 Contribution to grians (%)	收获指数 Harvest index	结实率 (%)	粒重 Kernel weight (mg)	产量 Yield (g·m^-2)
‘汕优63’ ‘Shanyou63’	水层灌溉 Well-watered	186.1^a	43.4^b	12.8^b	0.50^b	84.5^a	26.43^a	851.9^a
	水分亏缺 Water-deficit	92.7^b	71.8^a	24.6^a	0.56^a	79.8^b	25.87^b	771.3^b
‘Pc/早’ ‘Pc/Zao’	水层灌溉 Well-watered	265.1^a	16.3^b	6.7^b	0.40^b	77.3^a	24.57^a	945.6^a
	水分亏缺 Water-deficit	130.4^b	59.4^a	19.8^a	0.49^a	72.6^b	23.64^b	859.8^b

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