高温胁迫对不同耐热型马铃薯块茎形成期生长和光合特性的影响

周进华; 李有涵; 张兴; 胡荣海; 郭华春

doi:10.12357/cjea.20220658

高温胁迫对不同耐热型马铃薯块茎形成期生长和光合特性的影响

doi: 10.12357/cjea.20220658

1.
云南农业大学农学与生物技术学院薯类研究所　昆明　650201
2.
云南泵龙马铃薯种植有限公司　昆明　655200

基金项目: 国家马铃薯产业技术体系(CARS-09-15P)和云南(昆明)院士专家工作站专项(YSZJGZZ-2021058)资助

详细信息

作者简介:
周进华, 主要研究方向为马铃薯育种与栽培。E-mail: zjh75@qq.com

通讯作者:
郭华春, 主要研究方向为薯类作物育种与栽培。E-mail: ynghc@126.com

中图分类号: S532
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出版历程
- 收稿日期: 2022-08-26
- 录用日期: 2022-11-14
- 修回日期: 2022-11-14
- 网络出版日期: 2022-11-25
- 刊出日期: 2023-05-10

Growth and photosynthetic characteristics of different heat-sensitive potato genotypes during the tuberization stage under high-temperature stress

1.
Tuber and Root Crop Research Institute, College of Agronomy and Biotechnology, Yunnan Agricultural University, Kunming 650201, China
2.
Yunnan Benglong Potato Planting Co. Ltd., Kunming 655200, China

Funds: This study was supported by China Agriculture Research System (CARS-09-15P) and the Special Project of Yunnan (Kunming) Academician Expert Workstation (YSZJGZZ-2021058).

More Information

Corresponding author: E-mail: ynghc@126.com

摘要

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度显著(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, 更低的光补偿点和暗呼吸速率, 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。
- 马铃薯 /
- 高温胁迫 /
- 形态特征 /
- 光合参数 /
- 产量
Abstract: The potato (Solanum tuberosum L.) is an important grain and vegetable crop. Global warming affects its growth and production owing to its high temperature sensitivity. Investigating the physiological differences between heat-tolerant and heat-sensitive resources can help rationalize the mechanism of high-temperature resistance in potatoes. The parameters related to the 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 ℃. Under high-temperature stress, the plant height and internode length were increased, the leaves were upright, the length and area of leaves were reduced, and the plant architecture was more compact. The extent of change in the leaf number and bend angle in D187 was greater than that in QS9. The high-temperature affected potato net photosynthetic rate, water use efficiency, maximum net photosynthetic rate, apparent quantum yield, carboxylation efficiency, maximum carboxylation rate, and maximum electron transport rate, which were lower in QS9 than those in D187 under high-temperature stress. Furthermore, D187 had a lower light compensation point and dark respiration rate than the heat-sensitive cultivar (QS9), and as a result of its strong adaptability, the number of indexes with phenotypic plasticity index exceeding 0.5 in D187 was more than that in QS9. The mean phenotypic plasticity index of morphology, photosynthesis, and yield was 0.448 in D187, which was higher than that in QS9 (0.418). Furthermore, under high-temperature stress, the ability to absorb CO₂ and low-concentration CO₂ utilization were weakened, along with the acceleration of water loss and the reduction of water use efficiency in potato plants. Consequently, respiratory consumption increased, and the regeneration abilities of ribulose 1,5-diphosphate (RuBP) and chlorophyll fluorescence parameters were reduced in the dark. In contrast, chlorophyll fluorescence parameters increased under light, and the utilization ability of limited light was also enhanced. Differences in morphology and photosynthetic self-adaptation abilities are the main reasons for the difference in high-temperature resistance between heat-tolerant and heat-sensitive resources, which will help clarify the mechanism of high-temperature adaptability in potato plants and provide references for the selection of cultivars with high-temperature resistance and innovation in cultivation techniques.
- Potato /
- High-temperature stress /
- Morphological character /
- Photosynthetic parameters /
- Yield

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图 1 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下的结薯表现

Figure 1. Tuberization of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures

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图 2 ‘青薯9号’和‘滇187’马铃薯在正常温度(20 ℃)和高温(30 ℃)下的产量性状

**表示两温度间差异极显著(P<0.01, n=9)。** indicates significant difference between 20 ℃ and 30 ℃ (P<0.01, n=9).

Figure 2. Yield characters of potato materials QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures

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图 3 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯在正常温度(20 ℃)和高温(30 ℃)下植株株型和叶片性状

Figure 3. Plant architecture and leaf morphology of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures

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图 4 ‘青薯9号’和‘滇187’马铃薯在正常温度(20 ℃)和高温(30 ℃)下植株株型和叶片形态指标

*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 differences 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

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图 5 ‘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荧光特征参数的变化

Figure 5. Changes of fluorescence characteristic parameters of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures during light response and photosynthetic induction

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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

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图 7 ‘青薯9号’(QS9)和‘滇187’(D187)马铃薯高温(30 ℃)下形态、光合和产量特征等各参数的表型可塑性指数

Figure 7. Phenotypic plasticity indexes of parameters of morphology, photosynthesis, and yield of potato materials of QS9 and D187 at high (30 ℃) temperatures

P_n: net photosynthetic rate; G_s: stomatal conductance; C_i: intercellular CO₂ concentration; T_r: transpiration rate; WUE: water use efficiency; P_n·max: maximum net photosynthetic rate; LSP: light saturation point; LCP: light compensation point; R_d: dark respiration rate; AQY: apparent quantum yield; CE: carboxylation efficiency; R_L: photorespiration; CCP: carbon dioxide compensation point; V_c·max: maximum carboxylation rate; J_max: maximum electron transportation rate; T_30%P: time to reach 30% of maximum photosynthetic rate of dark adaptation ; T_60%P: time to reach 60% of maximum photosynthetic rate of dark adaptation; T_90%P: time to reach 90% of maximum photosynthetic rate of dark adaptation; IS_60s: maximum photosynthetic rate within 60 s; IS_300s: maximum photosynthetic rate within 300 s; IS_600s: maximum photosynthetic rate within 600 s; F₀: initial fluorescence intensity; F_m: maximum fluorescence intensity; F_v/F_m: potential photochemical quantum efficiency of photosystem Ⅱ; F_v/F₀: potential photochemical activity of photosystem Ⅱ.

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图 8 ‘青薯9号’(QS9)和‘滇187’(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。P_n、G_s、C_i、T_r、WUE、P_{n max}、LSP、LCP、R_d、AQY、CE、R_L、CCP、V_{c max}、J _max、F₀、F_m、F_v/F_m、F_v/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. P_n, G_s, C_i, T_r, WUE, P_{n max}, LSP, LCP, R_d, AQY, CE, R_L, CCP, V_{c max}, J_max, F₀, F_m, F_v/F_m, and F_v/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, initial 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 times 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 morphology, photosynthesis, and yield indexes of potato materials of QS9 and D187 at normal (20 ℃) and high (30 ℃) temperatures

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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 parameter	QS9			D187
光合特征参数 Photosynthetic parameter	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 differences between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively.

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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 response	QS9		D187
光响应特征参数 Parameter of light response	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±0.00	0.06±0.00^**	0.07±0.02	0.07±0.00
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 differences between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively.

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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 Parameter of CO₂ response	QS9		D187
CO₂响应特征参数 Parameter of CO₂ response	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 (μmol∙mol⁻¹)	90.50±2.06	112.92±3.61^**	119.94±4.91	135.01±18.79
最大羧化速率 Maximum carboxylation rate (V_{c max}, 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 (J_max, μmol∙mol⁻¹)	570.87±24.04	504.44±31.37	727.32±103.26	678.66±28.21
J_max/V_{c max}	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 differences between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively.

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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 Parameter of photosynthetic induction	QS9		D187
光合诱导特征参数 Parameter of photosynthetic induction	20 ℃	30 ℃	20 ℃	30 ℃
T_30%P (s)	289.37±27.11	194.11±11.47^**	327.04±46.83	193.83±10.57^*
T_60%P (s)	582.35±50.62	317.94±20.31^**	626.05±96.23	375.44±25.88^*
T_90%P (s)	1308.12±109.89	624.70±42.5^**	1366.76±227.63	825.33±65.42^*
IS_60s (%)	13.92±3.27	25.57±3.72^**	8.09±4.5	27.09±2.27^**
IS_300s (%)	45.63±4.01	74.81±3.19^**	41.97±7.11	65.26±3.25^*
IS_600s (%)	69.33±3.65	93.44±1.38^**	66.96±7.39	86.17±2.37^*
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。和分别表示两温度间差异显著(P<0.05)和极显著(P<0.01)。T_30%P, T_60%P and T_90%P are the times 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. and ** indicate significant differences between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively.

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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 parameter	QS9		D187
叶绿素荧光参数 Chlorophyll fluorescence parameter	20 ℃	30 ℃	20 ℃	30 ℃
黑暗下初始荧光强度 Initial fluorescence intensity (F₀)	199.40±1.37	163.90±24.66	134.63±1.49	131.03±2.33
黑暗下最大荧光强度 Maximum fluorescence intensity (F_m)	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 Potential photochemical quantum efficiency of photosystem Ⅱ (F_v/F_m)	0.82±0.002	0.81±0.001^**	0.82±0.001	0.81±0.002^**
PSⅡ潜在光化学活性 Potential photochemical activity of photosystem Ⅱ (F_v/F₀)	4.41±0.06	4.15±0.03^**	4.53±0.03	4.28±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 differences between 20 ℃ and 30 ℃ at P<0.05 and P<0.01, respectively.

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表 6 ‘青薯9号’(QS9)和‘滇187’(D187)形态特征、光合特征和产量特征参数初始特征值及累积贡献率

Table 6. Initial eigenvalues and accumulated variance contribution of each component of morphology, photosynthesis, and yield 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 (%)
QS9	1	26.72	65.17	65.17
	2	4.55	11.11	76.27
	3	3.00	7.31	83.58
D187	1	23.96	58.44	58.44
	2	5.27	12.84	71.28
	3	3.94	9.62	80.90

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