Growth and photosynthetic characteristics study of different heat-sensitivity potato genotypes during tuberization stage at high-temperature stress
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摘要: 全球变暖对粮食生产的负面影响日益受到关注, 马铃薯是重要的粮菜兼用作物, 对高温敏感。探究耐热和热敏感型马铃薯资源在响应高温胁迫时的生理差异, 可为深入研究马铃薯耐热机制提供理论依据。本研究以耐热型品系‘滇187’(D187)和热敏感型品种‘青薯9号’(QS9)为材料, 在30 ℃高温胁迫处理2周后, 分析2个马铃薯材料在块茎形成期的植株形态和光合作用差异。在植株形态方面, 高温使马铃薯植株株高和节间长显著(P<0.01)增加, 叶片直立、叶片长度和面积缩小, 株型更为紧凑; 与QS9相比, D187叶片数目和披垂角更为稳定。高温胁迫下马铃薯植株水分散失加快, 水分利用率降低, 对CO2吸收和低浓度CO2利用能力减弱, 呼吸作用消耗增加, 1,5-二磷酸核酮糖(RuBP)的再生能力减弱, 黑暗下的叶绿素荧光参数降低, 光下叶绿素荧光参数升高, 对有限强光的利用能力增强。高温胁迫下, D187叶片具有更高的净光合速率、水分利用效率、最大净光合速率、表观量子效率、羧化效率、最大羧化速率、最大电子传递速率, 更低的光补偿点、暗呼吸速率, 说明D187光合能力更强、弱光利用率更高、呼吸消耗更低、碳同化能力更强。D187的形态和光合作用指标中, 可塑性指数大于0.5的参数均多于QS9, 平均可塑性指数(0.448)高于QS9 (0.418), 说明耐热型马铃薯能够更好地通过调节植株形态和光合作用来适应高温环境。Abstract: Potato (Solanum tuberosum L.) is an important grain and vegetable crop, global warming affects its growth and production because of its high-temperature-sensitive characteristic. Investigating the physiological difference between heat tolerant and heat sensitive resources can help to rationalive the mechanism of high-temperature resistance in potato. The parameters related to morphology and photosynthesis of the heat-tolerant line ‘Dian 187’ (D187) and the heat sensitive cultivar ‘Qingshu 9’ (QS9) were measured and analyzed after two weeks of high temperature stress at 30 ℃. In high temperature stress, the plant height and internode length were increased, and the leaves were upright, the length and area of leaves were reduced, and the plant architecture was more compact; the change extent of the leaves number and bend angle in D187 was greater than in QS9, indicating that potato morphology affected the net photosynthetic rate, water use efficiency, maximum net photosynthetic rate, apparent quantum yield, carboxylation efficiency, maximum carboxylation rate, maximum electron transport rate in QS9 were lower than in D187 under high temperature stress. Furthermore, D187 had lower light compensation point and dark respiration rate than the heat sensitive one (QS9), As a result of strong adaptability, the amount of indexes exceeded over 0.5 in D187 were predominant plasticity indexes of morphology and photosynthesis and the indexes mean is 0.448 in D187 higher than 0.418 in QS9. Furthermore, the ability to absorb CO2 and low concentration CO2 utilization was weakened along with the acceleration of water loss and declination of water use efficiency in potato plant. As a consequence, the respiratory consumption was increased, the regeneration ability of ribulose 1,5-diphosphate (RuBP) and the chlorophyll fluorescence parameters were reduced in the dark. In contrast, the chlorophyll fluorescence parameters were increased under light and the utilization ability of limited light was also enhanced. Difference in morphology and photosynthesis self-adaptation abilities is the main reason for difference in high-temperature resistance between heat tolerant and heat sensitive resources, these results will help clarify the mechanism of high-temperature adaptability in potato plant, and provide references basic for the selection of high-temperature-resistant cultivars and innovation in cultivation techniques.
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Key words:
- Potato /
- High temperature stress /
- Morphological character /
- Photosynthetic parameters /
- Yield
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图 4 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下植株株型和叶片的形态指标
*和**分别表示两温度间差异显著(P<0.05, n=10)和极显著(P<0.01, n=10)。* and ** indicate significant difference between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively. n=10.
Figure 4. Morphological indexes of plant architecture and leaf of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures
图 6 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下光响应和光诱导过程中的光能分配特征
Φ PSⅡ: 光化学耗散比例、Φ NPQ: 非光化学猝灭耗散比例、Φ f,d: 荧光耗散比例。Ф PSⅡ: quantum yield of photochemical dissipation, Ф NPQ: quantum yield of non-photochemical quenching dissipation, Ф f,d: quantum yield of fluorescence quenching dissipation.
Figure 6. Characteristics of light energy distribution of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures during light response and photosynthetic induction
图 8 QS9和D187在正常温度和高温下的形态、光合及产量指标的主成分分析
椭圆为不同温度下各参数的置信区间; 箭头代表各指标与主成分的关系。Pn、Gs、Ci、Tr、WUE、Pn max、LSP、LCP、Rd、AQY、CE、RL、CCP、Vc max、J max、F0、Fm、Fv/Fm、Fv/F0依次为净光合速率、气孔导度、胞间CO2浓度、蒸腾速率、水分利用效率、最大净光合速率、光饱和点、光补偿点、暗呼吸速率、表观量子效率、羧化效率、光呼吸速率、CO2补偿点、最大羧化速率、最大电子传递速率、黑暗下初始荧光强度、黑暗下最大荧光强度、PSⅡ潜在光化学量子效率、PSⅡ潜在光化学活性; T 30%P、T 60%P、T 90%P为暗适应后达到最大净光合速率30%、60%、90%所需的时间, IS 60s、IS 300s、IS 600s为暗适应后60 s、300 s、600 s所达到的最大净光合速率百分比。The ellipse is the confidence interval of each parameter under different temperature; the arrow represents the relationship between each index and the principal component. Pn、Gs、Ci、Tr、WUE、Pn max、LSP、LCP、Rd、AQY、CE、RL、CCP、Vc max、J max、F0、Fm、Fv/Fm and Fv/F0 are Net photosynthetic rate, Stomatal conductance, Intercellular CO2 concentration, Transpiration rate, Water use efficiency, Maximum net photosynthetic rate, Light saturation point, Light compensation point, Dark respiration rate, Apparent quantum yield, Carboxylation efficiency, Photorespiration, Carbon dioxide compensation point, Maximum carboxylation rate, Maximum electron transportation rate Minimum fluorescence after dark adaptation, Maximum fluorescence after dark adaptation, Maximum quantum yield of photosystem II, Potential photochemical activity of photosystem II respectively. T 30%P, T 60%P and T 90%P are the time to reach 30%, 60% and 90% of maximum photosynthetic rate of dark adaptation; IS 60s, IS 300s and IS 600s are the proportions of the maximum photosynthetic rate within 60 s, 300 s and 600 s, respectively, after dark adaptation.
Figure 8. Principal component analysis of morphological, photosynthesis, and yield indexes of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures
表 2 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下的光响应特征参数
Table 2. Parameters of light response of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures
光响应特征参数
Parameter of light responseQS9 D187 20 ℃ 30 ℃ 20 ℃ 30 ℃ 最大净光合速率 Maximum net photosynthetic rate (μmol∙m−2∙s−1) 18.03±0.14 19.09±0.88 11.98±1.32 19.19±1.00** 光饱和点 Light saturation point (μmol∙m−2∙s−1) 530.25±12.30 749.59±37.87** 537.28±98.43 694.35±17.70 光补偿点 Light compensation point (μmol∙m−2∙s−1) 35.62±3.17 50.06±6.09* 29.40±1.66 45.35±21.62 暗呼吸速率 Dark respiration rate (mmol∙m−2∙s−1) −3.25±0.28 −3.43±0.48 −2.28±0.45 −2.59±0.93 表观量子效率 Apparent quantum yield (μmol∙mol−1) 0.08±0.002 0.06±0.0007** 0.07±0.02 0.07±0.004 *和**分别表示两温度间差异显著(P<0.05)和极显著(P<0.01)。* and ** indicate significant difference between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively. 表 4 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下的光合诱导特征参数
Table 4. Parameters of photosynthetic induction of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures
光合诱导特征参数
Parameters of photosynthetic inductionQS9 D187 20 ℃ 30 ℃ 20 ℃ 30 ℃ T30%P (s) 289.37±27.11 194.11±11.47** 327.04±46.83 193.83±10.57* T60%P (s) 582.35±50.62 317.94±20.31** 626.05±96.23 375.44±25.88* T90%P (s) 1308.12±109.89 624.7±42.5** 1366.76±227.63 825.33±65.42* IS60s (%) 13.92±3.27 25.57±3.72** 8.09±4.5 27.09±2.27** IS300s (%) 45.63±4.01 74.81±3.19** 41.97±7.11 65.26±3.25* IS600s (%) 69.33±3.65 93.44±1.38** 66.96±7.39 86.17±2.37* T30%P、T60%P、T90%P为暗适应后达到最大净光合速率30%、60%、90%所需的时间; IS60s、IS300s、IS600s为暗适应后60 s、300 s、600 s所达到的最大净光合速率百分比。*和**分别表示两温度间差异显著(P<0.05)和极显著(P<0.01)。T30%P, T60%P and T90%P are the time to reach 30%, 60% and 90% of maximum photosynthetic rate of dark adaptation; IS60s, IS300s and IS600s are the proportions of the maximum photosynthetic rate within 60 s, 300 s and 600 s, respectively, after dark adaptation. * and ** indicate significant difference between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively. 表 5 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下的叶绿素荧光参数
Table 5. Chlorophyll fluorescence parameters of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures
叶绿素荧光参数
Chlorophyll fluorescence parametersQS9 D187 20 ℃ 30 ℃ 20 ℃ 30 ℃ 黑暗下初始荧光强度 (F0) 199.4±1.37 163.9±24.66 134.63±1.49 131.03±2.33 黑暗下最大荧光强度 (Fm) 1078.28±17.95 844.43±127.65* 743.99±8.02 692.32±10.38** PSⅡ潜在光化学量子效率 (Fv/Fm) 0.82±0.002 0.81±0.001** 0.82±0.001 0.81±0.002** PSⅡ潜在光化学活性 (Fv/F0) 4.41±0.06 4.15±0.03** 4.53±0.03 4.28±0.05** 表 6 QS9和D187各成分初始特征值及累积贡献率
Table 6. The initial eigenvalues and the accumulated variance contribution of each component of potato materials of QS9 and D187
品种(系)
Variety (line)主成分
Principal component特征值
Eigenvalue贡献率
Rate of contribution (%)累积贡献率
Cumulative contribution (%)青薯9号
Qingshu 91 26.71783 65.16545 65.16545 2 4.55354 11.10619 76.27164 3 2.99833 7.313 83.58464 滇187
Dian 1871 23.95874 58.43595 58.43595 2 5.26625 12.84452 71.28047 3 3.94394 9.61937 80.89985 表 1 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下的净光合速率及相关参数
Table 1. Net photosynthetic rate and related parameters of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures
光合特征参数
Photosynthetic parametersQS9 D187 20 ℃ 30 ℃ 20 ℃ 30 ℃ 净光合速率 Net photosynthetic rate (μmol∙m−2∙s−1) 9.74±0.23 7.12±0.17** 7.43±1.67 8.31±0.51 气孔导度 Stomatal conductance (mol∙m−2∙s−1) 0.16±0.02 0.14±0.03 0.10±0.02 0.12±0.02 胞间CO2浓度 Intercellular CO2 concentration (μmol∙mol−1) 289.31±13.84 279.64±23.6 274.76±15.08 257.41±7.99 蒸腾速率 Transpiration rate (mmol∙m−2∙s−1) 3.93±0.41 6.35±0.92* 2.98±0.56 7.26±0.77** 水分利用效率 Water use efficiency (μmol∙mmol−1) 2.50±0.22 1.14±0.15** 2.20±0.31 1.19±0.05** *和**分别表示两温度间差异显著(P<0.05)和极显著(P<0.01)。* and ** indicate significant difference between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively. 表 3 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下的CO2响应特征参数
Table 3. Parameters of CO2 response of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures
CO2响应特征参数
Parameters of CO2 responseQS9 D187 20 ℃ 30 ℃ 20 ℃ 30 ℃ 羧化效率 Carboxylation efficiency (mol∙mol−1) 0.12±0.01 0.12±0.01 0.15±0.002 0.15±0.002 光呼吸速率 Photorespiration (μmol∙m−2∙s−1) −10.76±0.94 −13.58±0.44* −18.44±0.83 −20.87±2.80 CO2补偿点 Carbon dioxide compensation point (μmo∙mol−1) 90.5±2.06 112.92±3.61** 119.94±4.91 135.01±18.79 最大羧化速率 Maximum carboxylation rate (μmol∙mol−1) 142.60±7.75 141.74±9.40 157.75±9.04 167.39±2.86 最大电子传递速率 Maximum electron transportation rate (μmol∙mol−1) 570.87±24.04 504.44±31.37 727.32±103.26 678.66±28.21 J max/Vc maxs 4.01±0.05 3.56±0.04** 4.65±0.9 4.05±0.13 *和**分别表示两温度间差异显著(P<0.05)和极显著(P<0.01)。* and ** indicate significant difference between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively. -
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