摘要
氮素营养和水分是影响水稻生长的两个关键因素。为提高水分利用率,水稻节水栽培方式已成为目前的研究热点,在不同地区发展了覆草、覆膜等节水栽培方式。水稻旱作后,主要有两个环境条件的发生改变：田间水分状况由淹水转变为控制灌溉或相对轻度水分胁迫；土壤中氮素形态由铵态氮为主转变为以硝态氮为主或铵态氮和硝态氮混合。因此,研究不同水分条件和氮素形态对水稻生长发育、水分和光合生理、氮素吸收和代谢的影响具有重要的理论意义。
本研究以不同基因型水稻品种(汕优63、扬稻6号、86优8和武运粳7号)为材料,采用营养液培养及聚乙二醇(PEG6000)模拟水分胁迫的方法,研究不同氮素形态和水分条件对不同基因型水稻生长发育、水分和光合生理、渗透调节以及氮素代谢的影响及其生理机制。阐明不同氮素形态和水分条件对不同基因型水稻在胁迫下的响应,为指导不同地区水稻节水栽培提供适宜品种参考。主要结果如下：
1.与正常水分条件相比,水分胁迫对铵、硝营养水稻根系的生长均有促进效应,尤其在硝营养水稻中表现显著。就地上部生长而言,水分胁迫对铵营养处理籼稻的地上部干重和分蘖数无影响,但显著降低了硝营养籼稻品种；而粳稻品种铵、硝营养地上部干重均显著变化,但水分胁迫条件下铵营养粳稻的地上部干重显著高于硝营养粳稻。在两种水分条件下,供应铵营养水稻整株生物量均显著高于硝营养水稻,在水分胁迫条件下表现出较强的抗旱性。
2.与硝营养相比,水分胁迫条件下,铵营养水稻的根系具有较高的根系活力、吸收面积和水分吸收能力。无论是在正常水分还是在胁迫水分条件下,铵营养水稻根系水分吸收和运输的途径以水通道蛋白为主的细胞-细胞途径；而硝营养水稻根系在正常水分条件下以细胞-细胞途径占优势；水分胁迫后水分运输途径转为质外体途径为主。
3.与正常水分条件相比,水分胁迫条件下,铵、硝营养水稻根系通气组织(根系孔隙度)均有所增加,在硝营养根系增加尤为显著。水分胁迫显著降低了硝营养水稻根系水流导度和伤流液流速,而对铵营养水稻无影响。硝营养水稻根皮层通气组织的发育阻碍了水分的从根表面到根系中柱内的径向运输,从而降低了硝营养水稻的水分吸收。铵营养的水稻由于主要通过水通道蛋白进行水分运输,受根系通气组织的影响很小。
4.在两种水分条件下,铵营养水稻的净光合速率均显著高于硝营养水稻。水分胁迫显著降低了硝营养水稻的总叶面积和新完全展开叶的叶面积和叶绿素含量(SPAD值),而对铵营养水稻无影响,说明铵营养水稻在水分胁迫条件下能够维持较高的光合特性,从而为水稻的生长提供更多的同化产物。
5.正常水分条件下,硝营养水稻叶片中硝酸还原酶活性显著高于铵营养水稻；水分胁迫条件下,硝营养水稻硝酸还原酶显著降低,铵营养水稻基本不受水分胁迫影响。与正常条件下相比,水分胁迫条件下,硝营养水稻叶片对硝态氮的同化能力显著下降,铵营养水稻各部位对氮素的同化能力基本不受影响,从而保证了逆境下植物生长所需的充足氮源。
6.在不同水分和氮素形态条件下,水稻各部位有机渗透调节物质(脯氨酸、游离氨基酸和可溶性糖)的含量均为叶片>茎秆>根系。水分胁迫条件下,铵营养水稻各部位有机渗透调节物质含量均显著高于硝营养水稻；而铵、硝营养水稻各部位无机离子(K+、Ca2+和Mg2+)的含量无变化,说明本研究条件下无机离子(K+、Ca2+和Mg2+)参与水稻的渗透调节不显著。有机渗透调节物质是水稻的主要渗透调节物质。
7.分根实验结果表明,供应硝营养一侧根系水分胁迫显著降低水稻地上部的生长和干物质的积累,尤以两侧根系同时供应硝营养的水稻中表现明显。在两侧根系分别供应两种氮素形态营养(铵营养和硝营养)时,无论在正常还是水分胁迫条件下,单侧根系供应铵营养均有利于稳定和提高水稻叶片中叶绿素含量和光合速率。水分胁迫下供应硝营养一侧根系的通气组织与水分吸收呈显著负相关,而对供应铵营养的一侧无相关性。说明水分胁迫所造成的铵、硝营养水稻根系水分吸收的差异可能主要是由于根系自身对环境的响应。
Water supply and nitrogen nutrition are the two major restrictions for plant growth and development. At present, developing water-saving.rice cultivations, such as covering with plastic film and straw mulching, is becoming a hot topic of agricultural production research. There are two major differences between traditional cultivation and non-flooded conditions:(1) rice plants are frequently suffered from water stress in non-flooded condition;(2) nitrogen form is transferred from ammonium to nitrate or the mixture of ammonium and nitrate. Therefore, it is important to study the influence of water stress and nitrogen forms on plant growth, water relations, photosynthesis, nitrogen abosorption and metabolization.
In order to study the effects of different nitrogen forms and water stress on plant growth, water relations, photosynthesis, nitrogen absorption and utilization, hydroponic experiments supplied with different N forms and water stress simulated by adding10%(M/V) polyethylene glycol6000were conducted, four rice genotypic varieties (Shanyou63, Yangdao6,86you8and Wuyunjing7) were utilized in the experiments, which can illustrate the response of different genotypic rice plants under the water stress condition, and pro vied varietal reference for conductions in different region with rice water-saving cultivation. The results obtained are as follows:
1. Compared with non-water stress condition, water stress significantly stimulated root growth in both N forms, and the stimulation was higher in nitrate nutrition. Of indica rice plants, water stress had no negative effects on shoot biomass and tillers in ammonium nutrition, while significantly decreased them in nitrate nutrition. Of japonica rice plants, water stress significantly decreased shoot biomass and tillers in both N forms. Under water stress, shoot biomass of japonica rice plants was higher in ammonium nutrition than in nitrate nutrition. Regardless of water stress, whole plant biomass was higher in ammonium nutrition than in nitrate nutrition.
2. Under water stress, root activity, absorption area and water uptake ability were higher in ammonium nutrition than in nitrate nutrition. Regardless of water status, water uptake and transport in ammonium nutrition were mainly through cell-cell passway; in nitrate nutrition, they were mainly through cell-cell pathway under non-water stress and throught apoplastic pathway under water stress condition.
3. Water stress increased root aerenchyma formation, especially in nitrate nutrition. Water stress decreased root root hydraulic conductivity and xylem sap flow rate in nitrate nutrition, while had no negative effects in ammonium nutrition. The formation of aerenchyma in nitrate nutrition restrained radial water transport in the root cylinder and, as a result, decreased water uptake ability. Water transport occurred mainly through Hg-sensitive water channels in ammonium-supplied rice roots.
4. Regardless of water status, net photosynthetic rate (Pn) was higher in ammonium nutrition than in nitrate nutrition. Water stress significantly decreased total leaf areas, new expanded leaf area and chlorophy content (SPAD value) in nitrate nutrition, while had no negative effects in ammonium nutrition. Maintenance of relative high photosynthesis will benefit in supplying more assimilates.
5. Under non-water stress condition, leaf NR activity is significantly higher in nitrate nutrition than in ammonium nutrition. Water stress significantly decreased NR activity in nitrate nutrition, while had no negative effect in ammonium nutrition. Compared with non-water stress, nitrate assimilation ability was decreased in nitrate nutrition, while it was not decreased in ammonium nutrition.
6. Water stress decreased leaf, stem and root water content in both N forms, especially in leaves which indicated that leaves were the most sensitive part to water stress. In all conditions, organic osmoticum (proline, free amino acid and soluble sugar) of different parts were leaves> shoot> root. Under water stress condition, the organic osmoticum contents were significantly higher in ammonium nutrition than in nitrate nutrition, whereas inorganic osmoticum contents (calcium ion and magnesium ion) have no abvious effect in nitrate and ammonium nutrition. It showed that organic osmoticums may participated in ammonium nutrition, while inorganic osmoticum contents may not participate in osmotic regulating of rice growth.
7. Under water stress, the shoot biomass of rice plants were significantly decreased in nitrate supplied sections, especially in conditions where the two part of roots are both supplied with nitrate nutrition. Regardless of water stress, supplying ammonium nutrition in either or both parts is benefit for maintaining or increasing leaf chlorophy content and net photosynthetic rate. Water uptake ability was negative related to aerenchyma formation in nitrate supplied section, while there was no close relationship in ammonium supplied section. It showed that, differences in water uptake ability between the two N forms were may derived from response of root to environment.
引文
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