Effects of high temperature stress during pollination on plant morphology, leaf photosynthetic characteristics, and yield of summer maize
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摘要: 针对黄淮海地区花期高温影响夏玉米安全生产的问题, 本研究以热敏感型品种‘先玉335’为试验材料, 以大田常温为对照(CK), 设置授粉期高温处理(HT), 研究授粉期高温胁迫对夏玉米植株形态、叶片光合特性、干物质积累与分配及产量形成的影响。结果表明: 2021年和2022年, HT处理的冠层最高温度超过40 ℃的天数分别为7 d和8 d, 高温处理期间冠层最高温度分别较CK高1.7~6.8 ℃和1.5~4.6 ℃。HT处理显著提高了夏玉米株高和穗位高, 对茎粗和叶面积无显著影响, 但延缓了生育后期叶片衰老, 2021年和2022年成熟期的叶面积较CK分别显著提高34.69%和163.72%。高温处理期间, HT处理的玉米叶片气孔导度、蒸腾速率和胞间CO2浓度显著升高(P<0.05), 叶片羧化效率、气孔限制值和水分利用效率显著降低(P<0.05); 叶片净光合速率随处理温度而变化, 处理温度过高(一般>40 ℃时)则显著降低(P<0.05), 反之则显著升高(P<0.05)或无显著变化, 叶片整体光合性能下降。高温胁迫解除后, HT处理的叶片光合性能相关参数逐渐与CK趋于一致。经授粉期高温胁迫处理后, 玉米茎秆、叶片、苞叶、穗轴和单株干重显著, 其中穗轴干重降幅最大, 而雄穗和花丝干重显著增加, 使得干物质向茎秆、叶片、雄穗、花丝等部位的分配比例增加, 而向穗轴的分配比例显著减少(P<0.05)。至成熟期, HT处理造成玉米籽粒和单株干重显著减少48.32%和16.71% (P<0.05), 而玉米茎秆和叶片干重显著增加35.01%和9.48% (P<0.05)。HT处理的结实率和穗粒数分别显著下降54.43%和53.19% (P<0.05), 百粒重显著提高10.13% (P<0.05), 但籽粒产量显著降低46.82% (P<0.05)。综上, 授粉期高温胁迫增强了玉米叶片气孔蒸腾, 增加了胞间CO2浓度, 降低了叶片羧化效率和水分利用效率, 导致植株整体光合性能下降, 制约了光合同化物积累及向穗部的转移分配, 导致结实率显著下降, 穗粒数显著减少, 制约了花后光合同化物从“源”(茎秆和叶片)向“库”(籽粒)的转运, 最终导致籽粒产量大幅下降。Abstract: High temperatures during the flowering period seriously affect the safe production of summer maize in the Huang-Huai-Hai region. In this experiment, the heat-sensitive variety ‘Xianyu 335’ was used as the test material; the effects of high temperature stress during pollination on plant morphology, leaf photosynthetic characteristics, dry matter accumulation and distribution, and yield of summer maize were studied by setting up normal field temperature treatment (CK) and high temperature treatment during pollination (HT). The results showed that in 2021 and 2022, the number of days with maximum canopy temperatures exceeding 40 ℃ in the HT treatment group was 7 and 8 d, respectively, and the maximum canopy temperatures in the HT treatment group were higher than those in the CK group by 1.7−6.8 ℃ and 1.5−4.6 ℃, respectively. HT treatment significantly increased the plant height and ear height of summer maize but had no significant effect on stem diameter and green leaf area during the high-temperature treatment period. However, HT treatment delayed leaf senescence in the late reproductive stage of maize, and the green leaf area at maturity was 34.69% and 163.72% higher than that in the CK group in 2021 and 2022, respectively. During the high-temperature treatment period, leaf stomatal conductance, transpiration rate, and intercellular CO2 concentration were significantly higher and leaf carboxylation efficiency, stomatal limitation value, and water use efficiency were significantly lower in the HT treatment group than in the CK treatment group. The net photosynthetic rate of maize leaf in the HT treatment group varied with the treatment temperature: it significantly reduced when compared with that in the CK treatment group only when HT treatment temperature was too high (generally >40 ℃). High-temperature stress during pollination led to a decrease in the overall photosynthetic performance of the maize leaves. After exposure to 10 d of high temperature stress during pollination, the dry weights of corn stems, leaves, bracts, cobs, and individual plants decreased significantly. The dry weight of the cobs decreased the most, while those of the male ears and filaments increased significantly. HT treatment resulted in an increase in the partitioning of dry matter to stems, leaves, male ears, and filaments, and a significant decrease in partitioning to the cobs. At maturity, HT treatment significantly reduced the dry weights of corn grains by 48.32%, while those of corn stems and leaves increased by 35.01% and 9.48%. HT treatment resulted in a significant decrease of 54.43% and 53.19% in the seed setting rate and grain number per ear, respectively; a significant increase of 10.13% in 100-grain weight; and a significant decrease of 46.82% in the grain yield. In conclusion, high temperature stress during pollination enhanced the stomatal transpiration of maize leaves, increased intercellular CO2 concentration, and decreased leaf carboxylation efficiency and water use efficiency. It also led to a decline in the overall photosynthetic performance of the plant and restricted the accumulation of photosynthetic products and their transfer and partitioning to the ear, resulting in a significant decrease in seed setting rate and grain number per ear. This decline also restricted the post-flowering transport of photosynthesized assimilated compounds from the “source” (stems and leaves) to the “sink” (grains), which ultimately led to a significant decrease in grain yield.
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图 1 2021年和2022年高温处理期间不同处理的玉米冠层温度变化
CK: 对照; HT: 授粉期高温处理; CKmax: 对照日最高温度; CKavg: 对照日平均温度; HTmax: 高温处理日最高温度; HTavg: 高温处理日平均温度。CK: control; HT: high temperature treatment during pollination; CKmax: daily maximum temperature under control; CKavg: daily average temperature under control; HTmax: daily maximum temperature under high temperature treatment; HTavg: daily average temperature under high temperature treatment.
Figure 1. Daily temperature at maize canopy under different treatments during high temperature stress in 2020 and 2021
图 2 2021年和2022年高温胁迫对玉米株高、穗位高和茎粗的影响
CK: 对照; HT: 授粉期高温处理。柱上不同小写字母表示同一年份不同处理间P<0.05水平差异显著。CK: control; HT: high temperature treatment during pollination. Different lowercase letters above the bars indicate significant differences between treatments for the same year at P<0.05 level.
Figure 2. Effects of high temperature stress during pollination on plant height, ear height and stem diameter of maize in 2021 and 2022
图 3 2021年和2022年高温胁迫对玉米叶面积的影响
CK: 对照; HT: 授粉期高温处理; VT: 抽雄期。图中虚线表示高温处理结束日。*表示处理间在P<0.05水平差异显著。CK: control; HT: high temperature treatment during pollination; VT: tasseling stage.The dotted line in the figure indicates the end date of high temperature treatment. * indicates significant difference between treatments at P<0.05 level.
Figure 3. Effects of high temperature stress during pollination on maize leaf area in 2021 and 2022
图 4 2022年抽雄后第42天玉米植株航拍图
CK: 对照; HT: 授粉期高温处理。CK: control; HT: high temperature treatment during pollination.
Figure 4. Aerial photo of maize plants on the 42 d after tasseling in 2022
图 5 2021和2022年高温胁迫对玉米穗位叶净光合速率、气孔导度、胞间CO2浓度和蒸腾速率的影响
CK: 对照; HT: 授粉期高温处理; 图中虚线表示高温处理结束日; *和**分别表示处理间在P<0.05和P<0.01水平差异显著, ns表示处理间无显著性差异。CK: control; HT: high temperature treatment during pollination. The dotted line in the figure indicates the end date of high temperature treatment. * and ** indicate significant difference between treatments at P<0.05 and P<0.01 levels, respectively. ns means no significant difference between treatments.
Figure 5. Effects of high temperature stress during pollination on Pn, Gs, Ci and Tr of maize leaves in 2021 and 2022
图 6 2021和2022年高温胁迫对玉米穗位叶羧化效率、气孔限制值、瞬时和内在水分利用效率的影响
CK: 对照; HT: 授粉期高温处理; 图中虚线表示高温处理结束日; *和**分别表示处理间在P<0.05和P<0.01水平差异显著, ns表示处理间无显著性差异。CK: control; HT: high temperature treatment during pollination. The dotted line in the figure indicates the end date of high temperature treatment. * and ** indicate significant difference between treatments at P<0.05 and P<0.01 levels, respectively. ns means no significant difference between treatments.
Figure 6. Effects of high temperature stress during pollination on CE, Ls, WUEt and WUEi of maize leaves in 2021 and 2022
图 7 2021年和2022年高温胁迫对玉米干物质积累的影响
CK: 对照; HT: 授粉期高温处理; 图中虚线表示高温处理结束日。CK: control; HT: high temperature treatment during pollination. The dotted line in the figure indicates the end date of high temperature treatment.
Figure 7. Effects of high temperature stress during pollination on dry matter accumulation in 2021 and 2022
图 8 2021年和2022年高温胁迫对玉米产量构成的影响
CK: 对照; HT: 授粉期高温处理。柱上不同小写字母表示同一年份不同处理间P<0.05水平差异显著。CK: control; HT: high temperature treatment during pollination. Different lowercase letters above the bars indicate significant differences between treatments for the same year at P<0.05 level.
Figure 8. Effects of high temperature stress during pollination on maize yield and yield components in 2021 and 2022
表 1 2021年和2022年高温胁迫对玉米干物质分配的影响
Table 1. Effects of high temperature stress during pollination on dry matter distribution in 2021 and 2022
(%) 时期
Sampling time部位
Maize organ2021 2022 CK HT CK HT 抽雄后10 d 10 days after tasseling 茎秆 Stem 52.65±2.36a 53.39±0.43a 49.49±1.68a 50.13±0.96a 叶片 Leaf 25.41±0.77a 26.49±0.71a 27.62±2.63a 29.15±0.42a 雄穗 Tassel 1.02±0.11a 1.11±0.06a 1.62±0.23a 1.95±0.12a 花丝 Silk 0.93±0.06a 1.11±0.12a 0.87±0.02b 1.28±0.06a 苞叶 Bract 9.68±0.68a 9.12±0.07a 9.78±1.01a 10.30±0.47a 穗轴 Cob 10.32±0.77a 8.79±0.24a 10.61±0.24a 7.19±0.66b 抽雄后50 d 50 days after tasseling 茎秆 Stem 24.23±2.18b 29.79±0.27a 22.82±1.19b 47.41±1.80a 叶片 Leaf 12.34±0.21b 14.74±0.64a 12.29±0.09b 17.99±0.76a 雄穗 Tassel 4.81±0.14a 4.71±0.76a 4.31±1.20a 5.58±0.50a 苞叶 Bract 0.48±0.06a 0.41±0.14a 0.54±0.15a 0.76±0.14a 穗轴 Cob 8.40±0.20a 8.41±0.55a 8.03±0.23a 9.68±0.83a 籽粒 Grain 49.75±1.91a 41.93±1.72b 52.01±0.86a 18.57±2.64b CK: 对照; HT: 授粉期高温处理。同年同行数据后不同小写字母表示处理间差异达P<0.05显著水平。CK: control; HT: high temperature treatment during pollination. Values followed by different letters within the same row in the same year are significantly different at P<0.05. 
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