摘要
干旱胁迫是影响水稻生产的重要非生物胁迫因子之一。杂草稻具有较强的抗非生物胁迫的能力,是栽培稻遗传改良的重要种质资源。本文以收集于中国北方杂草稻为试材,进行芽期耐早性研究及苗期耐旱性鉴定,并以鉴定出的耐旱性1级的杂草稻HEB07-2及杂草稻WR04-6、巴西陆稻、旱敏感品种越富为试材。研究了不同程度干旱胁迫下(20%PEG胁迫0h、5h、24h、72h)不同杂草稻和栽培稻幼苗叶片和根系抗氧化防御系统、渗透调节系统的变化规律。并分析幼苗叶片光合系统在干旱胁迫下的变化,利用Affymetrix水稻表达谱芯片(GeneChip Rice Genome Array)比较不同杂草稻和栽培稻幼苗干旱胁迫下根系基因表达差异状况。主要研究结果如下：
利用15%PEG-6000(聚乙二醇6000)模拟干旱胁迫,对供试杂草稻和栽培稻最终发芽率无显著影响,杂草稻和栽培稻的发芽指数显著降低,即延迟了杂草稻和栽培稻的发芽时间。供试杂草稻和栽培稻的生长速度显著降低,对杂草稻的影响小于栽培稻,表现为正常条件下杂草稻和栽培稻地上和地下部生长量相当,在胁迫条件下杂草稻各生长指标有大于栽培稻的趋势。杂草稻的根长胁迫指数、茎长胁迫指数、根数胁迫指数、根干重胁迫指数、茎干重胁迫指数变异范围较大,而栽培稻变异范围较小。干旱胁迫下杂草稻根冠比呈增大趋势,而栽培稻根冠比降低,说明干旱胁迫对杂草稻根系抑制程度较栽培稻小。
利用苗期反复干旱胁迫法对杂草稻和栽培稻苗期耐早性进行了鉴定,结果表明第一次干旱胁迫对杂草稻和栽培稻的存活率影响较小,而第二次干旱胁迫后杂草稻和栽培稻存活率下降幅度较大,杂草稻变异范围大于栽培稻,苗期反复干旱存活率最大的杂草稻为LYG07-10。干旱胁迫期间,杂草稻叶片抗衰度变异范围大于栽培稻,抗衰度最大的杂草稻为HEB07-2。按苗期抗旱性综合系数划分抗旱级别,杂草稻抗旱性达到1级、3级、5级、7级的数量分别为8、25、15、6份,分别占供试杂草稻材料的12.7%、39.7%、23.8%、9.5%,属于正态分布。即耐旱性极强和极弱的杂草稻较少。
干旱胁迫期间,供试杂草稻和栽培稻叶片和根系丙二醛含量和超氧阴离子自由基产生速率显著上升,HEB07-2叶片和根系丙二醛上升幅度显著小于其他试材,超氧阴离子自由基产生速率最小,显著小于其他供试材料,说明HEB07-2在干旱胁迫下积累活性氧较少,膜质过氧化程度较低,细胞膜较为完整,受伤害程度较小。
各试材在正常条件下抗氧化酶活性相当,巴西陆稻略低,干旱胁迫使杂草稻HEB07-2叶片和根系超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)、谷胱甘肽还原酶(GR)、脱氢抗坏血酸还原酶(DHAR)活性显著上升,非酶抗氧化剂谷胱甘肽和抗坏血酸含量显著增加,而旱敏感品种越富胁迫72h后抗氧化酶活性和抗氧化剂含量均下降,巴西陆稻和WR04-6上升幅度小于HEB07-2,说明干旱胁迫下HEB07-2能够有效清除过量产生的活性氧(ROS),缓解干旱胁迫对生物膜的损伤。
干旱胁迫下,杂草稻HEB07-2叶片和根系渗透调节含量显著增加,而旱敏感品种越富叶片和根系可溶性糖、可溶性蛋白含量先升后降。WR04-6和巴西陆稻渗透调节物质上升幅度均小于HEB07-2。说明HEB07-2在干旱胁迫下通过积累更多渗透调节物质维持细胞水势,从而达到耐旱的目的。本研究通过对干旱胁迫下叶片和根系的ROS积累和抗氧化系统及渗透调节物质变化规律比较发现,叶片对PEG模拟的干旱胁迫更加敏感。
PEG模拟干旱胁迫72h后,旱敏感品种越富总叶绿素含量显著降低,而杂草稻HEB07-2叶绿素含量上升,WR04-6和巴西陆稻总叶绿素含量无显著变化。干旱胁迫期间,供试材料初始荧光(Fo)上升,最大荧光(Fm)下降。旱敏感品种越富上升幅度最大,而HEB07-2的Fo和Fm上升和下降时间较其他试材晚,说明其受伤害时间延后。干旱胁迫使净光合速率、气孔导度下降,旱敏感品种越富下降幅度最大,HEB07-2最小。利用红外热像仪测定群体温度变化发现,越富和WR04-6在干旱胁迫下叶温显著上升,而HEB07-2和巴西陆稻叶温无显著变化。
利用Affymetrix水稻表达谱芯片(GeneChip Rice Genome Array)将极抗旱的杂草稻样本HEB07-2与巴西陆稻样本进行表达谱差异分析,结果表明,杂草稻HEB07-2转录组对于干旱信号的响应程度与方向均与巴西陆稻存在很大差异,HEB07-2以正向调控为主,而巴西陆稻以负调控为主,在差异基因表达的倍数上也体现出了HEB07-2高于巴西陆稻的趋势。通过进一步的GO分析发现,杂草稻与巴西陆稻的三类差异表达基因,即生物过程相关基因、细胞组成相关基因、以及分子功能相关基因,均可以与两者间生理与生活指标的差异相互印证。
Drought stress is an important type of abiotic stress.15%PEG-6000(polyethylene glycol6000) was applied in this study. Results showed that it had no significant effect on the final germination rate of the test weedy rice and cultivated rice, but germination indexex of the weedy rice and cultivated rice were significantly reduced, while germination time of weedy rice and cultivated rice germination time were prolonged. Drought stress significantly reduced the growth rates of weedy rice and cultivated rice, but the reduction of the growth rates of weedy rice was comparatively less than cultivated rice. This suggested that under control treatments, weedy rice and cultivated rice had almost the same growth amounts in terms of ground and underground parts. However, this experiment showed of a trend that under drought stress, growth indexes of weedy rice was greater than that of the cultivated rice. Meanwhile, the weedy rice exhibited an much larger variation range in root length stress index, stem length stress index, the number of stress index, root dry weight stress index, stem heavy stress index than the cultivated rice. Under drought stress, weedy rice root to shoot ratio tends to increase while the cultivated rice root to shoot ratio tends were decrdasing, indicating drought stress had more significantly effect on the root inhibition of weedy rice than on that of the cultivated rice.
Repeated drought stress was adopted during seedling stage and drought tolerance was determined. Results showed that the survival rates of weedy rice and of cultivated rice were influenced slightly by the first drought stress, while they were influenced significantly by the second drought stress. In the second drought stress, weedy rice had much larger variation range than the cultivated rice survival rates. During the repeated drought stress experiment, weedy rice LYG07-10had the highest survival rate and weedy rice. HEB07-2had the highest anti-degree range. Drought level was divided according to drought resistance coefficient. The amounts of weedy rice drought resistance of1,3,5,7were8,25,15,6, respectively, making up of the test weedy rice materials12.7%,39.7%,23.8%,9.5%, respectively, which belonged to the normal distribution. That was the weedy rice with the strongest and the weakest drought tolerance was relatively less.
During drought stress, leaves and roots mda content and superoxide anion radical production rate of test weedy rice and cultivated rice increased significantly. Among them, the increased of mda content in leaves and roots of HEB07-2was significantly smaller than that of the other tested materials, and the superoxide anion radical production rate of it was the slowest, significantly smaller than the other tested materials, which indicated that the cell membrane HEB07-2was more complete and was injured less. Thus, under drought stress, HEB07-2had less accumulation of reactive oxygen species and lower degree of lipid peroxidation; Under normal conditions, each test material had similar activity of antioxidant enzymes, which was slightly lower in upland rice IRPARG Under tihe influence of drought stress, leaves and roots of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), ascorbate peroxidase (APX), glutathione reductase (GR), dehydroascorbate reductase (DHAR) activity increased significantly and the non-enzymatic antioxidants glutathione and ascorbic acid content also increased significantly. However,72hours later, the antioxidant enzyme activity and antioxidant content of drought-sensitive varieties decreased, while the increase rates of the upland rice IRPARG and WR04-6less than that of HEB07-2, which indicated that HEB07-2can effectively scavenge the excessive production of reactive oxygen species (ROS) to ease drought stress damage to the biological membrane. Leaf and root osmotic adjustment of HEB07-2was significantly increased by the impact of drought stress, and leaves and roots soluble sugar showed an earlier increase and later decrease. The increase of osmotic adjustment substances in WR04-6and IRPARG was smaller than that in HEB07-2, which indicated that under drought stress, HEB07-2can maintain cell water potential by accumulating more osmotic adjustment substances, so as to achieve the purpose of the drought resistance.
72h after the PEG simulated drought stress, total chlorophyll conten tin drought-sensitive varieties significantly reduced, while the chlorophyll content of HEB07-2increased. Total chlorophyll contentin WR04-6and Brazilian Upland did not change significantly. During drought stress, initial fluorescence (Fo) increased, while the maximum fluorescence (Fm) decreased. Drought-sensitive variety Yuefu had the highest increase. While the beginning time of Fo increased and decreased of Fm was later than others', which indicated that HEB07-2was injured later than others. The net photosynthetic rate and stomatal conductance were decreased due to drought stress. Among them, drought-sensitive variety Yuefu had the highest decrease of net photosynthetic rate and stomatal conductance, while HEB07-2had the lowest decrease. Groups temperature changes was determined by infrared camera measurement, leaf temperature in Yuefu and WR04-6increased significantly under drought stress, while leaf temperature in HEB07-2and upland rice IRPARG had no significant change.
Comparative analysis of terminal drought tolerance weedy rice HEB07-2between drought tolerance upland rice IRPARG by using expression profiles showed that the response to drought of transcriptome of weedy rice HEB07-2between upland rice IRPARG had significant difference in the respects of response directions and degrees. The majority of genes expressed in HEB07-2were most up-regulated, while in IRPARG were down-regulated, and the fold change of HEB07-2genes were larger. Gene ontology(GO) analysis suggested that weedy rice HEB07-2and upland rice IRPARG three categories of differently expressed genes related to biological process, cell component, molecular function, can be verified with physiological indexes.
引文
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