Responses of photosynthetic fluorescence characteristics, pollination, and yield components of maize cultivars to high temperature during flowering
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摘要: 探讨不同基因型玉米叶片光合荧光特性、授粉结实能力及产量构成对花期高温胁迫及恢复的响应, 以期为未来气候变暖条件下夏玉米稳产高产提供理论依据。采用人工模拟增温试验, 选取2个耐热型玉米品种(‘浚单20’和‘郑单958’)和2个热敏感型玉米品种(‘先玉335’和‘农华101’)为试验材料, 设置花期高温胁迫和大田常温对照2个处理, 研究花期高温胁迫及恢复对不同耐热性玉米叶片光合荧光特性、授粉结实能力、干物质积累与分配和产量构成的影响。结果表明, 花期高温胁迫显著降低了玉米穗粒数, 增加了空秆率, 进而导致籽粒产量显著下降, 且耐热型品种产量下降幅度小于热敏感型品种。与对照相比, 花期高温胁迫下耐热型和热敏感型品种穗粒数分别降低22.25%和67.18%, 百粒重分别降低2.03%和5.00%, 空秆率分别增加206.37%和283.00%, 籽粒产量分别降低31.84%和67.33%, 其中穗粒数、空秆率和籽粒产量差异均达显著水平(P<0.05)。花期高温胁迫使得4个玉米品种有效绿叶面积和叶绿素含量降低, 光系统Ⅱ受损, 光合性能显著下降; 高温胁迫结束后, 光系统Ⅱ部分参数有所恢复, 叶绿素含量和净光合速率恢复至对照水平。花期高温胁迫对雌穗总小花数、雄穗总小花数和雄穗分枝数影响较小, 但使得散粉持续期缩短, 开花吐丝间隔期拉长, 造成雌雄不遇, 结实率显著降低。花期高温导致的授粉持续期缩短是结实率降低的主要原因。花期高温胁迫显著降低了4个玉米品种干物质积累量和干物质向穗(粒)的分配比例; 高温胁迫后, 耐热型和热敏感型品种成熟期的单株干物质量分别降低16.40%和25.73%, 籽粒干物质分配比率分别降低7.08%和46.80%, 差异均达显著水平(P<0.05)。花期高温胁迫导致玉米光合性能下降, 雌雄穗协调发育受到抑制, 干物质积累量显著减少, 使得结实率下降, 穗粒数减少, 籽粒产量显著降低。花期高温胁迫对热敏感型品种的产量形成、光合荧光特性、雌雄穗发育和干物质积累与分配的影响均高于耐热型品种。Abstract: High temperatures during the flowering stage are likely to have a significant negative impact on maize growth and development, which is one of the most important factors affecting maize high and stable yield. This study aimed to evaluate the response of photosynthetic fluorescence characteristics, pollination, and yield components of different maize cultivars to high temperatures during the flowering stage to provide a theoretical basis for the stable and high yield of summer maize production under climate warming in the future. This study used heat-resistant maize cultivars, ‘XD20’ and ‘ZD958’, and heat-sensitive maize cultivars, ‘XY335’ and ‘NH101’, as research materials in a greenhouse. Then the influence of high temperature during the flowering stage (from 7 d before silking to 7 d after silking) on grain yield, leaf photosynthetic fluorescence characteristics, pollination, dry matter mass were investigated. High temperature during flowering significantly reduced the kernel number per ear and significantly increased the blank stem rate of different summer maize cultivars, leading to a significant decrease in grain yield. The grain yield decline of heat-resistant cultivars was less than that of heat-sensitive cultivars. Compared with the control, the kernel number per ear of the heat-resistant and heat-sensitive cultivars under high temperature significantly decreased by 22.25% and 67.18%, respectively, the 100-grain weight decreased by 2.03% and 5.00%, the blank stem rate significantly increased by 206.37% and 283.00%, and the grain yield significantly decreased by 31.84% and 67.33%, respectively. High temperature during flowering reduced effective green leaf area and chlorophyll content, impaired photosystem II, and significantly decreased the photosynthetic performance of the four maize cultivars. Under high-temperature stress, the leaf area of the heat-resistant and heat-sensitive cultivars decreased by 0.79% and 7.46%, the chlorophyll content decreased by 4.53% and 5.16%, the net photosynthetic rate (Pn) decreased by 19.9% and 31.6%, and the maximum photochemical efficiency (Fv/Fm) of PS Ⅱ decreased by 0.79% and 1.47%, respectively. After the high-temperature stress, some parameters of photosystem Ⅱ recovered, and chlorophyll content and Pn returned to the control level. High temperatures during flowering had little effect on the tassel branch number, tassel floret number, and ear filament number; but shortened the pollen shedding duration, lengthened the anthesis-silking interval, and significantly reduced the setting rate. Under high-temperature stress at the flowering stage, the tassel branch number of heat-resistant and heat-sensitive cultivars decreased by 4.76% and 13.66%, the tassel floret number decreased by 8.53% and 8.32%, the tear filament number decreased by 6.10% and 7.17%, the pollen shedding duration decreased by 10.81% and 26.94%, the anthesis-silking interval increased by 58.93% and 85.00%, the pollination duration decreased by 17.91% and 58.95%, and the kernel setting rate decreased by 14.77% and 63.10%, respectively. Shortening pollination duration was the main reason for the lower kernel setting rate. High temperature during flowering significantly reduced dry matter mass and the distribution ratio of dry matter to the ear or grain of four maize cultivars. After high-temperature stress, the dry matter mass per plant of heat-resistant and heat-sensitive cultivars decreased by 13.7% and 17.6%, and the distribution ratio of dry matter in the ear decreased by 49.16% and 56.51%, respectively; at maturity, the dry matter mass per plant decreased by 16.40% and 25.73%, and the distribution ratio of dry matter in the grain decreased by 7.08% and 46.80%, respectively. High temperature during flowering decreased the photosynthetic performance, inhibited the coordinated development of male and female panicles, and significantly reduced kernel setting rate, kernel number per ear, and grain yield. Compared with heat-sensitive cultivars, heat-resistant cultivars had higher photosynthetic capacity and pollination fruiting ability under high-temperature stress; their yield was less affected by high temperature.
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图 1 玉米花期高温处理(HT)和常温处理(CK)的1 d内平均温度变化(A)和15 d试验期的温度变化(B)
Figure 1. Temperature changes in one day (A) and during the 15 days experiment period (B) at maize flowering stage under normal temperature (CK) and high temperature (HT) treatments
图 2 花期高温处理(HT)和常温处理(CK)对不同夏玉米穗位叶叶绿素相对含量(SPAD)的影响
高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。7 d: 吐丝后7 d; 20 d: 吐丝后20 d。不同小写字母表示同一品种不同处理不同时间在P<0.05水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. 7 d: 7 days after silking; 20 d: 20 days after silking. Different lowercase letters above the bars represent significant differences among different treatments in different time after silking of the same cultivar at P<0.05 level.
Figure 2. Effects of the normal temperature (CK) and high temperature (HT) treatments at flowering stage on SPAD values in the ear-leaf of different summer maize cultivars
图 3 花期高温处理(HT)和常温处理(CK)对不同夏玉米品种雄穗分枝数和雄穗总小花数的影响
高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。不同小写字母表示同一品种不同处理不同时间在P<0.05水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. Different lowercase letters above the bars represent significant differences among different treatments in different time after silking of the same cultivar at P<0.05 probability level.
Figure 3. Effects of the normal temperature (CK) and high temperature (HT) treatments at flowering stage on tassel branch number and tassel floret number of different summer maize cultivars
图 4 花期高温处理(HT)和常温处理(CK)对不同夏玉米品种散粉持续期、开花吐丝间隔期和授粉持续期的影响
高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。不同小写字母表示同一品种不同处理在P<0.05水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. Different lowercase letters above the bars represent significant differences btween two treatments of the same cultivar at P<0.05 level.
Figure 4. Effects of the normal temperature (CK) and high temperature (HT) treatments at flowering stage on pollen shedding duration, anthesis-silking interval and pollination duration of different summer maize cultivars
图 5 花期高温处理(HT)和常温处理(CK)对不同夏玉米品种雌穗总小花数和结实率的影响
高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。不同小写字母表示同一品种不同处理在P<0.05水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. Different lowercase letters above the bars represent significant differences between two treatments of the same cultivar at P<0.05 level.
Figure 5. Effects of the normal temperature (CK) and high temperature (HT) treatments at flowering stage on ear filament number and kernel setting rate of different summer maize cultivars
图 6 花期高温处理结束当天高温处理(HT)和常温处理(CK)不同夏玉米品种地上部干物质量及分配比例
高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。*和**分别表示同一品种不同处理间P<0.05和P<0.01水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. * and ** represent significant difference between two treatments for the same cultivar at P<0.05 and P<0.01 levels, respectively.
Figure 6. Dry matter mass and distribution in different organs of different summer maize cultivars under the normal temperature (CK) and high temperature (HT) treatments at flowering stage at the end of high temperature treatment
图 7 花期高温处理(HT)和常温处理(CK)下不同夏玉米品种成熟期地上部干物质量及分配比例
高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。*和**分别表示同一品种不同处理间P<0.05和P<0.01水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. * and ** represent significant differences between two treatments for the same cultivar at P<0.05 and P<0.01 levels, respectively.
Figure 7. Dry matter mass and distribution in different organs of different summer maize cultivars at the maturity stage under the normal temperature (CK) and high temperature (HT) treatments at flowering stages
图 8 花期高温处理(HT)和常温处理(CK)对不同夏玉米品种产量及其构成要素的影响
高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。不同小写字母表示同一品种不同处理在P<0.05水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. Different lowercase letters above the bars represent significant differences between two treatments of the same cultivar at P<0.05 level.
Figure 8. Effect of the normal temperature (CK) and high temperature (HT) treatments at flowering stage on grain yield and its components of different summer maize cultivars
图 9 不同夏玉米品种各指标耐热系数分析
高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。A: 吐丝后7 d叶片光合荧光性状指标的耐热系数分析; B: 吐丝后20 d叶片光合荧光性状指标的耐热系数分析; C: 雌雄穗性状指标的耐热系数分析; D: 干物质积累分配及产量性状指标的耐热系数分析。LA: 叶面积; SPAD: 叶绿素相对含量; Pn: 净光合速率; Gs: 气孔导度; Ci: 胞间CO2浓度; Fv/Fm: PSⅡ最大光化学效率; Fv/Fo: PSⅡ潜在活性; PIabs: 光合性能指数; φEo: 用于电子传递的量子产额; φRo: 用于还原PSⅠ受体侧末端电子受体的量子产额; TBN: 雄穗分枝数; TFN: 雄穗总小花数; PSD: 散粉持续期; ASI: 开花吐丝间隔期; PD: 授粉持续期; EFN: 雌穗总小花数; KSR: 雌穗结实率; DM7: 吐丝后7 d总干物质量; Ear/DM7: 吐丝后7 d穗干物质分配比例; DM50: 成熟期总干物质量; Grain/DM50: 成熟期籽粒干物质分配比例; EL: 穗长; ED: 穗粗; KPE: 穗粒数; GW: 百粒重; GY: 籽粒产量。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD 958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. A: analysis of heat resistance coefficients of chlorophyll photosynthetic fluorescence parameters at 7 days after silking; B: analysis of heat resistance coefficients of chlorophyll photosynthetic fluorescence parameters at 20 days after silking; C: analysis of heat resistance coefficients of male and female spike characters; D: analysis of heat resistance coefficients of dry matter and yield characters. LA: leaf area; SPAD: chlorophyll relative content; Pn: net photosynthetic rate; Gs: stomatal conductance; Ci: intercelluar CO2 concentration; Fv/Fm: maximum photochemical efficiency; Fv/Fo: PSⅡ potential activity; PIabs: photosynthetic performance index; φEo: quantum yield for electron transport; φRo: quantum yield for reducing electron acceptor at the PSⅠ receptor side; TBN: tassel branch number; TFN: tassel floret number; PSD: pollen shedding duration; ASI: anthesis-silking interval; PD: pollination duration; EFN: ear filament number; KSR: kernel setting rate; DM7: dry matter mass at 7 d after silking; Ear/DM7: the distribution ratio of dry matter in the ear at 7 d after silking; DM50: dry matter mass at maturity stage; Grain/DM50: the distribution ratio of dry matter in the grain at the maturity stage; EL: ear length; ED: ear diameter; KPE: kernel number per ear; GW: 100-grain weight; GY: grain yield.
Figure 9. Analysis of heat resistance coefficients of each index of four summer maize cultivars
表 1 花期高温处理(HT)和常温处理(CK)对不同夏玉米品种不同生育时期单株叶面积的影响
Table 1. Effects of the normal temperature (CK) and high temperature (HT) treatments during flowering stage on leaf area per plant of different summer maize cultivars
品种
Cultivar处理
Treatment单株叶面积 Leaf area per plant (cm2) 7DAS 20DAS 30DAS 50DAS XD20 CK 6730.2±308.8a 6667.8±326.3a 5105.9±397.6a 3385.7±120.9a HT 6714.1±366.1a 6576.2±291.2a 5442.2±147.2a 3758.5±166.6a ZD958 CK 6326.6±311.6a 6285.4±278.7a 5520.5±83.6b 4547.1±187.5b HT 6241.6±177.1a 6166.6±107.9a 6027.4±116.7a 4853.7±79.9a XY335 CK 6087.3±170.5a 5956.7±151.2a 5892.5±187.3a 3595.6±228.8a HT 5934.2±392.4a 5798.0±436.5a 5535.4±346.2a 4234.7±258.8a NH101 CK 6091.6±163.7a 5933.8±84.0a 5857.1±400.2a 2972.1±237.7b HT 5335.9±273.6b 5184.0±278.6b 5406.0±315.7a 3573.5±368.6a 高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。7DAS: 吐丝后7 d; 20DAS: 吐丝后20 d; 30DAS: 吐丝后30 d; 50DAS: 吐丝后50 d。同一玉米品种同列不同小写字母表示两个处理在P<0.05水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. 7DAS: 7 days after silking, the ending time of high temperature stress; 20DAS: 20 days after silking; 30DAS: 30 days after silking; 50DAS: 50 days after silking. Values within a column followed by different lowercase letters of the same cultivar are significantly different at P<0.05 level. 
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表 2 花期高温处理(HT)和常温处理(CK)对不同夏玉米品种穗位叶光合参数的影响
Table 2. Effects of the normal temperature (CK) and high temperature (HT) treatments at flowering stage on photosynthetic parameters in the ear-leaf of different summer maize cultivars
测定时期
Sampling time品种
Cultivar处理
Treatment净光合速率
Net photosynthetic rate
[μmol(CO2)·m−2·s−1]气孔导度
Stomatal conductance
[mmol(CO2)·m−2·s−1]胞间CO2浓度
Intercelluar CO2 concentration (μmol·mmol−1)吐丝后7 d
7 days after silkingXD20 CK 26.8±1.3a 0.198±0.006a 59.3±9.9b HT 20.7±1.7b 0.156±0.013b 124.8±2.3a ZD958 CK 24.2±0.8a 0.204±0.009a 53.3±7.8b HT 20.1±0.9b 0.156±0.012b 112.9±20.2a XY335 CK 33.1±1.2a 0.210±0.012a 71.8±8.5b HT 24.9±2.0b 0.146±0.008b 199.4±4.7a NH101 CK 27.8±2.3a 0.175±0.012a 49.3±8.0b HT 17.1±3.2b 0.112±0.013b 132.4±10.1a 吐丝后20 d
20 days after silkingXD20 CK 16.9±0.3a 0.164±0.029a 41.7±3.1a HT 16.5±1.5a 0.151±0.035a 37.3±6.9a ZD958 CK 17.3±0.8a 0.189±0.017a 44.6±2.4a HT 16.5±0.3a 0.173±0.015a 40.6±4.1a XY335 CK 17.4±1.1a 0.223±0.016a 87.3±3.1a HT 16.0±1.4a 0.215±0.005a 83.9±2.7a NH101 CK 16.1±0.6a 0.188±0.007a 50.2±10.0a HT 14.9±0.7a 0.178±0.015a 47.6±0.8a 高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。同列同一玉米品种同一测定时间不同小写字母表示两个处理在P<0.05水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. Values within a column followed by different lowercase letters of the same cultivar in the same sampling time are significantly different at P<0.05 level.
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表 3 花期高温处理(HT)和常温处理(CK)对不同夏玉米品种叶绿素荧光参数的影响
Table 3. Effects of the normal temperature (CK) and high temperature (HT) treatments at flowering stage on fluorescence parameters in the ear-leaf of different summer maize cultivars
测定时期 Sampling time 品种 Cultivar 处理 Treatment Fv/Fm Fv/Fo PIabs φEo φRo 吐丝后7 d
7 days after silkingXD20 CK 0.808±0.002a 4.20±0.05a 7.05±0.20a 0.557±0.002a 0.218±0.010a HT 0.801±0.001b 4.12±0.03b 6.96±0.53a 0.554±0.007a 0.217±0.006a ZD958 CK 0.804±0.004a 4.08±0.12a 8.33±0.72a 0.573±0.014a 0.274±0.015a HT 0.798±0.004a 3.99±0.08a 8.10±0.25a 0.566±0.002a 0.267±0.013a XY335 CK 0.803±0.001a 4.08±0.02a 7.40±0.61a 0.558±0.008a 0.206±0.010a HT 0.791±0.016a 3.83±0.34a 7.00±1.40a 0.550±0.021a 0.198±0.015a NH101 CK 0.811±0.001a 4.29±0.02a 8.87±0.59a 0.568±0.006a 0.232±0.010a HT 0.800±0.001b 4.11±0.01b 8.57±0.15a 0.556±0.001a 0.213±0.008a 吐丝后20 d
20 days after silkingXD20 CK 0.737±0.008b 2.81±0.12a 2.04±0.34a 0.409±0.023a 0.208±0.009a HT 0.777±0.007a 3.53±0.34a 3.84±0.32a 0.463±0.026a 0.192±0.004a ZD958 CK 0.723±0.036a 2.74±0.49a 2.62±0.90a 0.428±0.032a 0.232±0.031a HT 0.776±0.014a 3.50±0.29a 3.30±0.59a 0.449±0.022a 0.211±0.004a XY335 CK 0.784±0.007a 3.65±0.15a 5.15±0.60a 0.515±0.007a 0.221±0.011a HT 0.801±0.004a 4.01±0.10a 5.37±0.69a 0.510±0.011a 0.196±0.005a NH101 CK 0.791±0.004b 3.78±0.08b 6.11±1.09a 0.536±0.019a 0.208±0.035a HT 0.806±0.005a 4.14±0.12a 6.22±0.23a 0.508±0.004b 0.180±0.014a 高温处理时间为吐丝前7 d至吐丝后7 d; XD20和ZD958为耐热型品种, XY335和NH101为热敏感型品种。Fv/Fm: PSⅡ最大光化学效率; Fv/Fo: PSⅡ潜在活性; PIabs: 光合性能指数; φEo: 用于电子传递的量子产额; φRo: 用于还原PSⅠ受体侧末端电子受体的量子产额。同列同一玉米品种同一测定时间不同小写字母表示两个处理在P<0.05水平差异显著。The high temperature treatment duration is from 7 days before silking to 7 days after silking. XD20 and ZD958 are heat-resistant cultivars; XY335 and NH101 are heat-sensitive cultivars. Fv/Fm: maximum photochemical efficiency; Fv/Fo: PSⅡ potential activity; PIabs: photosynthetic performance index; φEo: quantum yield for electron transport; φRo: quantum yield for reducing electron acceptor at the PSⅠ receptor side. Values within a column followed by different lowercase letters of the same cultivar in the same sampling are significantly different at P<0.05 level.
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