小麦灌浆期高温、干旱及其互作对籽粒蛋白质形成、品质性状及产量的影响
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摘要
试验于2006～2008年在河南农业大学科教示范园区进行,采用盆栽与人工气候室模拟相结合的方式,以豫麦34(强筋)和豫麦50(弱筋)为试验材料,研究了灌浆期不同时段高温、干旱胁迫及其互作对籽粒蛋白质积累调控关键酶活性、蛋白质品质性状及籽粒产量的影响。主要研究结果如下:
1.灌浆期不同时段高温胁迫使小麦旗叶谷氨酰胺合成酶(GS)、谷氨酸丙酮酸转氨酶(GPT)活性降低,不同时期相比,豫麦34和豫麦50分别受前期(花后5~9 d)和中期(花后15~19 d)高温胁迫影响较大。高温胁迫下0～10 d籽粒GS、GPT活性升高,之后明显降低,且两品种均以中期高温胁迫影响较大。土壤干旱胁迫下,小麦旗叶GS活性下降,豫麦34和豫麦50分别受前期和中期干旱胁迫影响较大;受前期干旱胁迫影响,旗叶GPT活性略有提高,而中期干旱胁迫使其活性显著降低。干旱胁迫导致两品种籽粒GS活性下降,且均以中期干旱胁迫影响较大。干旱胁迫后5~10 d内籽粒GPT活性提高,之后明显降低,豫麦34和豫麦50分别以中期和前期受影响较大。从高温与干旱效应对旗叶和籽粒GS、GPT活性的影响大小看,高温效应明显大于干旱效应。
2.灌浆期不同时段高温与干旱胁迫提高了籽粒总蛋白含量,但降低了蛋白质产量,其影响程度随胁迫程度加深而加大,而不同时段处理则随着处理时间推迟而减弱。品种间比较,豫麦50籽粒蛋白含量受高温胁迫的影响较豫麦34明显。高温胁迫下,豫麦34籽粒清蛋白和球蛋白含量下降,而豫麦50清蛋白含量增加,球蛋白含量则表现为前期和中期高温处理下增加,后期(花后25~29 d)下降;两品种籽粒醇溶蛋白和谷蛋白含量均有增加趋势。受干旱胁迫影响,豫麦34籽粒清蛋白和球蛋白含量下降,而豫麦50则在前期和中期增加,后期下降;两品种醇溶蛋白和谷蛋白含量增加。高温与干旱胁迫均使谷/醇比值不同程度降低。研究还表明,高温与干旱对小麦蛋白质品质有明显的互作效应,温度是影响其品质的主要因子。
3.对籽粒谷蛋白大聚合体(GMP)含量的研究结果表明,高温胁迫提高了两品种成熟期籽粒GMP含量;干旱胁迫下两品种籽粒GMP含量存在品种间差异:豫麦34籽粒GMP含量因干旱胁迫而降低,而豫麦50增加。方差分析结果表明,高温与干旱及其互作对两品种籽粒GMP含量均有显著或极显著影响。
4.高温胁迫下豫麦34籽粒氨基酸总含量略有下降,而豫麦50则显著增加;干旱胁迫下两品种籽粒氨基酸总含量(TAA)均有明显增加。高温与干旱胁迫均使籽粒中第一限制性氨基酸(赖氨酸)含量降低,必需氨基酸含量占氨基酸总含量的百分比(EAA/TAA)下降。品种间比较,高温与干旱胁迫对豫麦34籽粒氨基酸含量及其组分影响较豫麦50明显。研究还表明,高温与干旱对氨基酸含量及其组分的影响存在显著的互作效应,其中豫麦34和豫麦50分别以W_1T_2和W_2T_3处理组合受高温与干旱双重胁迫伤害最大。
5.研究表明,花后不同时段高温胁迫均降低籽粒灌浆速率,使最大灌浆速率出现时间提前,灌浆持续期缩短,千粒重、穗粒重和产量显著下降。不同时段高温胁迫对粒重和产量的影响在不同品种间存在差异:豫麦34和豫麦50千粒重分别以前期和中期受高温胁迫影响较大,而产量以前期影响最为严重,后期受影响相对较小。受干旱胁迫影响,豫麦34粒重以中期受影响最大,而豫麦50以前期受影响较大,而产量均以前期影响最为严重,在后期干旱胁迫下,两品种粒重均略有增加。研究还表明,高温和干旱对小麦籽粒灌浆特性及粒重的影响存在明显的水、温互作效应。渐增期平均灌浆速率的降低及快增期、缓增期持续时间的缩短是最终导致粒重降低的主要原因。两品种比较,弱筋小麦豫麦50较强筋小麦豫麦34更易受高温与干旱胁迫影响。
上述研究结果表明,不同专用型小麦品种对高温与干旱胁迫的响应存在一定的差异:总体来看,弱筋小麦豫麦50产量和蛋白质品质性状受高温与干旱影响较大。高温与干旱胁迫使小麦粒重降低,因而籽粒蛋白质含量相对提高,但导致籽粒谷/醇比值降低,使加工品质明显下降。因此,小麦实际生产中应针对不同专用型小麦品种的品质需要,采取适当的栽培调控措施,缓解小麦生育后期不同时段的干热风危害,以提高小麦产量和品质。
To evaluate effects of post-anthesis high temperature (HT) and drought stress (D) and their interaction on protein formation and protein quality traits in wheat grains, two wheat cultivars with different gluten-strengths, Yumai 34 ( a strong-gluten cultivar) and Yumai 50 ( a weak-gluten cultivar) were used under pot-cultured conditions. The experiments were carried out at Henan Agricultural University Research farm, Zhengzhou (34oN latitude, 113o E longitude), in wheat growing-seasons of 2006-2007 and 2007-2008. Plants in pot grown in open area till flowering. After anthesis, pots were divided into three groups in the artifical weather room for each variety randomly, were grown under 28℃, 2 day 38℃high temperature stress and 4 day 38℃high temperature stress, respectively. For each temperature regime, two soil water levels were established as moderate water status (relative water content, RWC = 75%±5%), drought ( RWC = 55%±5% ). Thus different temperature and water regimes were established for wheat after anthesis to evaluate post-anthesis high temperature and drought stress effects on activities of key regulatory enzymes of protein accumulation and protein traits in grains and yield of
winter wheat.
The main results were as follows:
1. Post-anthesis high temperature decreased both GS and GPT activity in flag leaves of wheat. However, the response of two cultivars to high temperature at different grain filling stages were different: when suffered from high temperature in early and middle grain filling stage, the reduction in GS and GPT activity of Yumai 34 and Yumai 50 was relatively greatest, respectively. During 0 to 10 days after high temperature treatment in early grain filling stage, GS and GPT activity in grains was increased, and then decreased in 10 days after high temperature treatment, the reduction in GS and GPT activity in middle grain filling stage of two cultivars were greatest. GS and GPT activity in flag leaves of the two wheat cultivars was decreased by drought in early and middle grain filling stages. The reduction in GS activity in flags in middle and early grain filling stage of Yumai 34 and Yumai 50 was relatively greatest under drought stress, respectively. While in early grain filling stage, GPT activity in flags was increased by drought stress, but decreased in middle grain filling stage. GS activity in grains the two wheat cultivars was decreased by drought in early and middle grain filling stages. The reduction in GS activity in grains in middle grain filling stage of two wheat cultivars was relatively greatest under drought stress. During 5 to 10 days after drought treatment in early grain filling stage, GPT activity in grains was increased, and then decreased in 10 days after high temperature treatment, when suffered from drought stress in middle and early grain filling stage, the reduction in GPT activity of Yumai 34 and Yumai 50 was relatively greatest, respectively. The results also showed that significant interactions of high temperature and drought on GS and GPT activity in both wheat cultivars. The influence of high temperature was greater than Drought stress.
2. Studies on effects of post-anthesis high temperature and drought stress and their interactions on protein concentration and its composition in grains. The results indicated that high temperature and drought stress increased protein concentration, but reduced protein yield, the influence of high temperature and drought stress on protein concentration aggravation follow their stress deepened, and the influence reduction with processing time delay. The influence of high temperature and drought stress on protein of Yumai 50 was greater than that of Yumai 34. While high temperature decreased concentration of albumin and globulin of Yumai 34, however, high temperature increased concentration of albumin of Yumai 50, and when suffered from high temperature in early and middle grain filling stage, the concentration of globulin was increased, but decreased in late grain filling stage. The concentration of gliadin and glutein were increased under high temperature stress. Under drought stress, concentration of albumin and globulin of Yumai 34 was decreased, however, when suffered from high temperature in early and middle grain filling stage, the concentration of that of Yumai 50 was increased, but decreased in late grain filling stage. The concentration of gliadin and glutein were increased under drought stress. High temperature and drought stress reduced ratio of gliadin/glutein. The results also showed that significant interactions of high temperature and drought on protein in both wheat cultivars, and the temperature were the main factor of its protein quality.
3. Studies on effects of high temperature and drought stress and their interactions on GMP concentration. The results indicated that high temperature stress increased GMP concentration in both two wheat cultivars. However, the response of two cultivars to drought stress at GMP concentration was different: when suffered from high temperature, the concentration of GMP was decreased in grains of Yumai 34, but increased in grains of Yumai 50. The result also indicated that there had a significant influence of high temperature and drought stress and their interactions on concentration of GMP in both two cultivars.
4. The results indicated that, high temperature stress decreased the Total Amino Acid in grains of Yumai 34, however increased the Total Amino Acid in grains of Yumai 50. When suffered from drought stress, Total Amino Acid was increased by high temperature and drought in both wheat cultivars. High temperature and drought stress reduced concentration of Lysine, and decreased the ratio of Essential Amino Acid to Total Amino Acid. The influence of high temperature and drought stress on Yumai 34 was greater than Yumai 50. The results also showed that there had a significant interaction of high temperature and drought stress on concentration and composition of grain Amino Acid in both cultivars. The influence of interaction of high temperature and drought stress on the treatment W_1T_2 (RWC=75%±5%, 2 day 38℃high temperature stress) and W2T3 (RWC=55%±5%, 4 day 38℃high temperature stress) was injury largest for strong and weak-gluten cultivar respectively.
5. The results showed that high temperature stress decreased both grain filling rate and period, and caused early appearance of the maximal grain filling rate, thus reduced grain dry matter accumulation. However, the response of two cultivars to high temperature at different grain filling stages were different: when suffered from high temperature in early and middle grain filling stage, the reduction in grain weight of Yumai 34 and Yumai 50 was relatively greatest, respectively. While in the late grain filling stage, grain weight was less influenced by high temperature and drought in both wheat cultivars. The dry matter accumulation of two cultivars was reduced by drought in early and middle grain filling stages. The reduction in grain weight in middle and early grain filling stage of Yumai 34 and Yumai 50 was relatively greatest under drought stress, respectively. But grain weight was slightly increased in late grain filling stage under drought stress. The results also showed that significant interactions of high temperature and drought on grain filling characteristic existed in both wheat cultivars. The decreased of grain weight was mainly caused by decrease of grain filling rate during slow-increase stage, and duration of rapid-increase and slight-increase stage. High temperature and drought stress and their interactions had a significant influence on grain yield and its components in this experiment. The influence of high temperature and drought stress on Yumai 50 was greater than Yumai 34, indicated that cultivar Yumai 34 with strong-gluten was more sensensive than Yumai 50 with weak-gluten.
In conclusion, it was showed that the response of two cultivars with different gluten to high temperature and drought stress at different grain filling stages were different: in generally, weak-gluten wheat cultivar Yumai 50 showed relative sensitive to temperature and drought stress on yield and protein property in grain filling stage. High temperature and drought stress decreased grain yield, thus relative increased grain protein concentration, but caused ratio of gliadin/glutein and processing quality traits significant declined. So, in actual production of wheat, cultivating measures should be taken according to the quality need in different cultivars to avoid or relieve damages of high temperature and drought stress at later growth stages.
1.灌浆期不同时段高温胁迫使小麦旗叶谷氨酰胺合成酶(GS)、谷氨酸丙酮酸转氨酶(GPT)活性降低,不同时期相比,豫麦34和豫麦50分别受前期(花后5~9 d)和中期(花后15~19 d)高温胁迫影响较大。高温胁迫下0～10 d籽粒GS、GPT活性升高,之后明显降低,且两品种均以中期高温胁迫影响较大。土壤干旱胁迫下,小麦旗叶GS活性下降,豫麦34和豫麦50分别受前期和中期干旱胁迫影响较大;受前期干旱胁迫影响,旗叶GPT活性略有提高,而中期干旱胁迫使其活性显著降低。干旱胁迫导致两品种籽粒GS活性下降,且均以中期干旱胁迫影响较大。干旱胁迫后5~10 d内籽粒GPT活性提高,之后明显降低,豫麦34和豫麦50分别以中期和前期受影响较大。从高温与干旱效应对旗叶和籽粒GS、GPT活性的影响大小看,高温效应明显大于干旱效应。
2.灌浆期不同时段高温与干旱胁迫提高了籽粒总蛋白含量,但降低了蛋白质产量,其影响程度随胁迫程度加深而加大,而不同时段处理则随着处理时间推迟而减弱。品种间比较,豫麦50籽粒蛋白含量受高温胁迫的影响较豫麦34明显。高温胁迫下,豫麦34籽粒清蛋白和球蛋白含量下降,而豫麦50清蛋白含量增加,球蛋白含量则表现为前期和中期高温处理下增加,后期(花后25~29 d)下降;两品种籽粒醇溶蛋白和谷蛋白含量均有增加趋势。受干旱胁迫影响,豫麦34籽粒清蛋白和球蛋白含量下降,而豫麦50则在前期和中期增加,后期下降;两品种醇溶蛋白和谷蛋白含量增加。高温与干旱胁迫均使谷/醇比值不同程度降低。研究还表明,高温与干旱对小麦蛋白质品质有明显的互作效应,温度是影响其品质的主要因子。
3.对籽粒谷蛋白大聚合体(GMP)含量的研究结果表明,高温胁迫提高了两品种成熟期籽粒GMP含量;干旱胁迫下两品种籽粒GMP含量存在品种间差异:豫麦34籽粒GMP含量因干旱胁迫而降低,而豫麦50增加。方差分析结果表明,高温与干旱及其互作对两品种籽粒GMP含量均有显著或极显著影响。
4.高温胁迫下豫麦34籽粒氨基酸总含量略有下降,而豫麦50则显著增加;干旱胁迫下两品种籽粒氨基酸总含量(TAA)均有明显增加。高温与干旱胁迫均使籽粒中第一限制性氨基酸(赖氨酸)含量降低,必需氨基酸含量占氨基酸总含量的百分比(EAA/TAA)下降。品种间比较,高温与干旱胁迫对豫麦34籽粒氨基酸含量及其组分影响较豫麦50明显。研究还表明,高温与干旱对氨基酸含量及其组分的影响存在显著的互作效应,其中豫麦34和豫麦50分别以W_1T_2和W_2T_3处理组合受高温与干旱双重胁迫伤害最大。
5.研究表明,花后不同时段高温胁迫均降低籽粒灌浆速率,使最大灌浆速率出现时间提前,灌浆持续期缩短,千粒重、穗粒重和产量显著下降。不同时段高温胁迫对粒重和产量的影响在不同品种间存在差异:豫麦34和豫麦50千粒重分别以前期和中期受高温胁迫影响较大,而产量以前期影响最为严重,后期受影响相对较小。受干旱胁迫影响,豫麦34粒重以中期受影响最大,而豫麦50以前期受影响较大,而产量均以前期影响最为严重,在后期干旱胁迫下,两品种粒重均略有增加。研究还表明,高温和干旱对小麦籽粒灌浆特性及粒重的影响存在明显的水、温互作效应。渐增期平均灌浆速率的降低及快增期、缓增期持续时间的缩短是最终导致粒重降低的主要原因。两品种比较,弱筋小麦豫麦50较强筋小麦豫麦34更易受高温与干旱胁迫影响。
上述研究结果表明,不同专用型小麦品种对高温与干旱胁迫的响应存在一定的差异:总体来看,弱筋小麦豫麦50产量和蛋白质品质性状受高温与干旱影响较大。高温与干旱胁迫使小麦粒重降低,因而籽粒蛋白质含量相对提高,但导致籽粒谷/醇比值降低,使加工品质明显下降。因此,小麦实际生产中应针对不同专用型小麦品种的品质需要,采取适当的栽培调控措施,缓解小麦生育后期不同时段的干热风危害,以提高小麦产量和品质。
To evaluate effects of post-anthesis high temperature (HT) and drought stress (D) and their interaction on protein formation and protein quality traits in wheat grains, two wheat cultivars with different gluten-strengths, Yumai 34 ( a strong-gluten cultivar) and Yumai 50 ( a weak-gluten cultivar) were used under pot-cultured conditions. The experiments were carried out at Henan Agricultural University Research farm, Zhengzhou (34oN latitude, 113o E longitude), in wheat growing-seasons of 2006-2007 and 2007-2008. Plants in pot grown in open area till flowering. After anthesis, pots were divided into three groups in the artifical weather room for each variety randomly, were grown under 28℃, 2 day 38℃high temperature stress and 4 day 38℃high temperature stress, respectively. For each temperature regime, two soil water levels were established as moderate water status (relative water content, RWC = 75%±5%), drought ( RWC = 55%±5% ). Thus different temperature and water regimes were established for wheat after anthesis to evaluate post-anthesis high temperature and drought stress effects on activities of key regulatory enzymes of protein accumulation and protein traits in grains and yield of
winter wheat.
The main results were as follows:
1. Post-anthesis high temperature decreased both GS and GPT activity in flag leaves of wheat. However, the response of two cultivars to high temperature at different grain filling stages were different: when suffered from high temperature in early and middle grain filling stage, the reduction in GS and GPT activity of Yumai 34 and Yumai 50 was relatively greatest, respectively. During 0 to 10 days after high temperature treatment in early grain filling stage, GS and GPT activity in grains was increased, and then decreased in 10 days after high temperature treatment, the reduction in GS and GPT activity in middle grain filling stage of two cultivars were greatest. GS and GPT activity in flag leaves of the two wheat cultivars was decreased by drought in early and middle grain filling stages. The reduction in GS activity in flags in middle and early grain filling stage of Yumai 34 and Yumai 50 was relatively greatest under drought stress, respectively. While in early grain filling stage, GPT activity in flags was increased by drought stress, but decreased in middle grain filling stage. GS activity in grains the two wheat cultivars was decreased by drought in early and middle grain filling stages. The reduction in GS activity in grains in middle grain filling stage of two wheat cultivars was relatively greatest under drought stress. During 5 to 10 days after drought treatment in early grain filling stage, GPT activity in grains was increased, and then decreased in 10 days after high temperature treatment, when suffered from drought stress in middle and early grain filling stage, the reduction in GPT activity of Yumai 34 and Yumai 50 was relatively greatest, respectively. The results also showed that significant interactions of high temperature and drought on GS and GPT activity in both wheat cultivars. The influence of high temperature was greater than Drought stress.
2. Studies on effects of post-anthesis high temperature and drought stress and their interactions on protein concentration and its composition in grains. The results indicated that high temperature and drought stress increased protein concentration, but reduced protein yield, the influence of high temperature and drought stress on protein concentration aggravation follow their stress deepened, and the influence reduction with processing time delay. The influence of high temperature and drought stress on protein of Yumai 50 was greater than that of Yumai 34. While high temperature decreased concentration of albumin and globulin of Yumai 34, however, high temperature increased concentration of albumin of Yumai 50, and when suffered from high temperature in early and middle grain filling stage, the concentration of globulin was increased, but decreased in late grain filling stage. The concentration of gliadin and glutein were increased under high temperature stress. Under drought stress, concentration of albumin and globulin of Yumai 34 was decreased, however, when suffered from high temperature in early and middle grain filling stage, the concentration of that of Yumai 50 was increased, but decreased in late grain filling stage. The concentration of gliadin and glutein were increased under drought stress. High temperature and drought stress reduced ratio of gliadin/glutein. The results also showed that significant interactions of high temperature and drought on protein in both wheat cultivars, and the temperature were the main factor of its protein quality.
3. Studies on effects of high temperature and drought stress and their interactions on GMP concentration. The results indicated that high temperature stress increased GMP concentration in both two wheat cultivars. However, the response of two cultivars to drought stress at GMP concentration was different: when suffered from high temperature, the concentration of GMP was decreased in grains of Yumai 34, but increased in grains of Yumai 50. The result also indicated that there had a significant influence of high temperature and drought stress and their interactions on concentration of GMP in both two cultivars.
4. The results indicated that, high temperature stress decreased the Total Amino Acid in grains of Yumai 34, however increased the Total Amino Acid in grains of Yumai 50. When suffered from drought stress, Total Amino Acid was increased by high temperature and drought in both wheat cultivars. High temperature and drought stress reduced concentration of Lysine, and decreased the ratio of Essential Amino Acid to Total Amino Acid. The influence of high temperature and drought stress on Yumai 34 was greater than Yumai 50. The results also showed that there had a significant interaction of high temperature and drought stress on concentration and composition of grain Amino Acid in both cultivars. The influence of interaction of high temperature and drought stress on the treatment W_1T_2 (RWC=75%±5%, 2 day 38℃high temperature stress) and W2T3 (RWC=55%±5%, 4 day 38℃high temperature stress) was injury largest for strong and weak-gluten cultivar respectively.
5. The results showed that high temperature stress decreased both grain filling rate and period, and caused early appearance of the maximal grain filling rate, thus reduced grain dry matter accumulation. However, the response of two cultivars to high temperature at different grain filling stages were different: when suffered from high temperature in early and middle grain filling stage, the reduction in grain weight of Yumai 34 and Yumai 50 was relatively greatest, respectively. While in the late grain filling stage, grain weight was less influenced by high temperature and drought in both wheat cultivars. The dry matter accumulation of two cultivars was reduced by drought in early and middle grain filling stages. The reduction in grain weight in middle and early grain filling stage of Yumai 34 and Yumai 50 was relatively greatest under drought stress, respectively. But grain weight was slightly increased in late grain filling stage under drought stress. The results also showed that significant interactions of high temperature and drought on grain filling characteristic existed in both wheat cultivars. The decreased of grain weight was mainly caused by decrease of grain filling rate during slow-increase stage, and duration of rapid-increase and slight-increase stage. High temperature and drought stress and their interactions had a significant influence on grain yield and its components in this experiment. The influence of high temperature and drought stress on Yumai 50 was greater than Yumai 34, indicated that cultivar Yumai 34 with strong-gluten was more sensensive than Yumai 50 with weak-gluten.
In conclusion, it was showed that the response of two cultivars with different gluten to high temperature and drought stress at different grain filling stages were different: in generally, weak-gluten wheat cultivar Yumai 50 showed relative sensitive to temperature and drought stress on yield and protein property in grain filling stage. High temperature and drought stress decreased grain yield, thus relative increased grain protein concentration, but caused ratio of gliadin/glutein and processing quality traits significant declined. So, in actual production of wheat, cultivating measures should be taken according to the quality need in different cultivars to avoid or relieve damages of high temperature and drought stress at later growth stages.
引文
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