Analysis of photosynthetic and fluorescence characteristics of low-light tolerant genotype potato under shade condition
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摘要: 为探寻遮阴对马铃薯光合和荧光特性及吸收光能分配的影响，本研究采用大田试验，以马铃薯不耐弱光品种‘丽薯6号’和耐弱光品种‘中薯20’为材料，出苗后用遮光率70%的黑色遮阳网进行遮阴处理（T），以自然光照为对照（CK），测定了遮阴30 d后马铃薯叶片的光合作用、光响应曲线、CO2响应曲线、光诱导曲线和叶绿素荧光参数的变化。结果表明：1）遮阴后净光合速率（Pn）、气孔导度（Gs）、蒸腾速率（Tr）、最大净光合速率（Pmax）、光饱和点（LSP）、光补偿点（LCP）较CK显著下降；‘中薯20’的Pn、Pmax、LSP较高，LCP和暗呼吸速率（Rd）较低。2）不同基因型CO2响应参数无显著差异，但‘中薯20’的最大羧化速率（Vcmax）和最大电子传递速率（Jmax）较高，CO2补偿点（CCP）较低。3）高光诱导过程中，‘中薯20’反应较快，光合能力较强。4）初始荧光（Fo）、最大荧光（Fm）、最大光化学量子效率（Fv/Fm）较CK显著增加，PSⅡ实际光化学量子效率（ΔF/Fm'）、表观电子传递速率（ETR）和光化学猝灭系数（qP）较CK显著下降，非光化学猝灭系数（NPQ）呈增加趋势。5）非光化学热耗散（ФNPQ）和荧光耗散途径（Фf，d）比例显著增加，光化学猝灭耗散途径（ФPSⅡ）比例显著减少，主要以增加热耗散为主。遮阴后，耐弱光基因型‘中薯20’的NPQ和ФNPQ均高于不耐弱光的‘丽薯6号’，说明‘中薯20’的光合机构保护能力更强。综合分析表明，遮阴后耐弱光基因型马铃薯具有较高的净光合速率、较低的光补偿点、较低的CO2补偿点、较快的光诱导反应速度和较高的非光化学热耗散能力。Abstract: With constant development of potato cultivation in winter agriculture fields in south China and in relay-intercropping systems in recent years, low light stress has been becoming the emerging limitation to potato development in these regions and it has increasingly gained considerable research attention. In order to explore the effects of low light stress on photosynthetic and chlorophyll fluorescent characteristics along with the distribution of absorbed light energy of low-light tolerant genotype potato, a field experiment was conducted with low-light sensitive potato cultivar 'Lishu 6' and low-light tolerant potato cultivar 'Zhongshu 20'. The experiment consisted of a shade treatment using black net that can filter 70% of sunlight (T) and then a light treatment with full natural sunlight (CK) after seed emergence. The variations in photosynthesis, light and CO2 response curves, light induced curve and chlorophyll fluorescence parameters of the different potato genotypes following 30 days of shade treatment were measured. The results following the shade treatment were as follows: 1) the net photosynthetic rate (Pn), stomatal conductance (Gs), transpiration rate (Tr), maximum net photosynthetic rate (Pmax), light saturation point (LSP) and light compensation point (LCP) under T were significantly lower than those under CK. While Pn, Pmax and LSP of 'Zhongshu 20' potato cultivar were relatively higher than those of 'Lishu 6' potato cultivar, LCP and dark respiration rate (Rd) of 'Zhongshu 20' potato cultivar were lower than those of 'Lishu 6' potato cultivar. 2) The parameters of CO2 response curve were not significantly different between the shade and control treatments, but 'Zhongshu 20' potato cultivar had higher maximum carboxylation rate (Vcmax), higher maximum electron transportation rate (Jmax) and lower CO2 compensation point (CCP) than 'Lishu 6' potato cultivar. 3) Potato cultivar 'Zhongshu 20' reacted quicker to high introduction rate and performed better in terms of photosynthetic capacity than potato cultivar 'Lishu 6'. 4) Compared with the control (CK), the increases of initial fluorescence (Fo), maximum fluorescence (Fm) and maximum photochemical quantum efficiency (Fv/Fm) were significantly higher in T treatments. Also reductions in actual photochemical quantum efficiency of PSⅡ(ΔF/Fm'), apparent electron transfer rate (ETR) and photochemical quenching coefficient (qP) were significantly higher in T treatments. Furthermore, there was an increasing trend in non photochemical quenching coefficient (NPQ). 5) Compared with the control, the ratios of non-photochemical dissipation (ФNPQ) and fluorescence dissipation (Фf, d) pathways increased remarkably under shade treatment. The ratio of photochemical quenching pathway (ФPSⅡ) decreased significantly, making ФNPQ as the main pathway of dissipation. After shade treatment, the low-light tolerant potato genotype 'Zhongshu 20' overall performance for NPQ and ФNPQ was higher than the low-light sensitive potato genotype 'Lishu 6', suggesting that 'Zhongshu 20' cultivar had more stronger photosynthetic apparatus protective capability. The above comprehensive analysis indicated that low-light tolerance genotype potatoes had relatively higher Pn, lower LCP and CCP, faster light induced reaction rate and higher non-photochemical dissipation capacity under shade conditions compared with that of low-light sensitive potato cultivar.
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表 1 遮光对马铃薯不同弱光耐受性品种的光合特征参数的影响
Table 1 Effect of shading on photosynthetic characteristics of potato cultivars with different low-light tolerance
光合特征参数
Photosynthetic parameter丽薯6号Lishu 6 中薯20 Zhongshu 20 CK T CK T 净光合速率(Pn)
Net photosynthetic rate (μmol·m-2·s-1)21.45±2.10a 12.26±0.80b 24.20±1.75a 17.52±0.24b 气孔导度(Gs)
Stomatal conductance (mol·m-2·s-1)0.40±0.08a 0.18±0.05b 0.57±0.11a 0.45±0.09b 胞间CO2浓度(Ci)
Intercellular carbon dioxide concentration (μmol·mol-1)275.96±6.81a 272.17±18.15a 286.80±10.22b 307.30±13.45a 蒸腾速率(Tr)
Transpiration rate (mmol·m-2·s-1)4.39±0.40a 2.68±0.47b 5.81±0.48a 4.64±0.41b 水分利用效率(WUE)
Water use efficiency (μmol·mmol-1)54.75±6.17b 69.49±12.22a 43.45±7.05a 41.27±9.78a 最大净光合速率(Pmax)
Maximum net photosynthetic rate (μmol·m-2·s-1)20.91±0.90a 16.31±2.05b 25.56±2.08a 20.16±0.23b 光饱和点(LSP)
Light saturation point (μmol·m-2·s-1)1 799.16±361.67a 1 163.51±148.58b 2 293.35±61.18a 1 557.37±101.35b 光补偿点(LCP)
Light compensation point (μmol·m-2·s-1)45.83±13.12a 19.05±2.52b 27.31±2.90a 17.75±1.52b 暗呼吸速率(Rd)
Dark respiration rate (μmol·m-2·s-1)-2.64±0.18a -1.30±0.13b -1.47±0.31a -1.11±0.19a 表观量子产额(AQY)
Apparent quantum yield (μmol·mol-1)0.06±0.01a 0.07±0.00a 0.05±0.01a 0.06±0.00a 羧化效率(CE)
Carboxylation efficiency (mol·mol-1)0.06±0.03a 0.09±0.00a 0.10±0.01a 0.09±0.00a 光呼吸(RL)
Photorespiration (μmol·m-2·s-1)-4.21±1.71a -5.09±0.77a -5.74±0.82a -5.31±0.50a CO2补偿点(CCP)
Carbon dioxide compensation point (μmol·mol-1)67.80±7.01a 59.92±9.13a 57.34±7.08a 57.42±5.19a 最大羧化速率(Vcmax)
Maximum carboxylation rate (μmol·m-2·s-1)67.17±37.40a 91.86±6.14a 130.78±16.74a 104.65±3.78a 最大电子传递速率(Jmax)
Maximum electron transportation rate (μmol·m-2·s-1)190.57±98.16a 255.72±28.24a 399.83±61.21a 294.64±16.23b CK:自然光照处理; T:遮光70%处理。数据表示为平均值±标准差。不同小写字母表示5%水平差异显著。CK: normal light; T: light decreased by 70%. ‘Lishu 6’ is low-light sensitive cultivar, ‘Zhongshu 20’ is low-light tolerant cultivar. The data is expressed as mean ± standard deviation. Values followed by different lowercase letters are significantly different at 5% level. 
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表 2 遮阴对马铃薯不同弱光耐受性品种光合诱导参数的影响
Table 2 Effect of shading on photoinduction parameters of potato cultivars with different low-light tolerance
光合诱导参数
Photoinduction parameter丽薯6号Lishu 6 中薯20Zhongshu 20 CK T CK T T50%P(s) 361.76±205.93a 275.30±129.46a 123.35±22.99a 156.98±29.61a T90%P(s) 1 354.08±780.29a 879.27±455.32a 381.88±83.01a 469.29±116.88a IS1 min(%) 5.17±0.61a 2.43±1.17b 6.93±0.65a 3.12±0.26b IS5 min(%) 24.29±7.42a 19.69±1.73a 29.06±4.70a 20.18±1.68b IS10 min(%) 26.98±8.42a 18.30±2.04a 26.71±3.07a 17.74±1.39b T50%P、T90%P:暗适应后达到最大净光合速率50%、90%所需的时间; IS1 min、IS5 min、IS10 min:暗适应后1 min、5 min、10 min所达到的最大净光合速率百分比。CK:自然光照处理; T:遮光70%处理。数据表示为平均值±标准差。不同小写字母表示5%水平差异显著。T50%P, T90%P: the time to reach 50% and 90% of maximum photosynthetic rate of dark adaptation; IS1 min, IS5 min, IS10 min: the maximum photosynthetic rate within 1 min, 5 min and 10 min, respectively, after dark adaptation. CK: normal light; T: light decreased by 70%. ‘Lishu 6’ is low-light sensitive cultivar, ‘Zhongshu 20’ is low-light tolerant cultivar. The data is expressed as mean ± standard deviation. Values followed by different lowercase letters are significantly different at 5% level.
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表 3 遮阴对马铃薯不同弱光耐受性品种叶绿素荧光参数的影响
Table 3 Effect of shading on chlorophyll fluorescence parameters of potato cultivars with different low-light tolerance
叶绿素荧光参数Chlorophyll fluorescence parameter 丽薯6号Lishu 6 中薯20Zhongshu 20 CK T CK T Fo 171.79±2.24b 174.58±1.42a 177.15±3.26b 182.30±3.53a Fm 1 041.34±29.43b 1 109.82±20.99a 1 042.45±45.71b 1 130.21±31.61a Fv/Fm 0.83±0.00b 0.84±0.00a 0.83±0.01b 0.84±0.00a Fv′/Fm′ 0.57±0.04a 0.57±0.01a 0.57±0.02a 0.55±0.01b ∆F/Fm′ 0.29±0.04a 0.20±0.01b 0.32±0.03a 0.20±0.02b ETR 182.70±22.92a 128.33±8.76b 199.92±21.71a 125.76±14.42b NPQ 1.74±0.31a 1.75±0.09a 1.74±0.14a 1.84±0.11a qP 0.51±0.04a 0.36±0.02b 0.56±0.05a 0.36±0.04b Fo:暗适应下的初始荧光; Fm:暗适应下最大荧光; Fv/Fm:暗适应下最大光化学量子效率; Fv′/Fm′:光适应下PSⅡ最大光化学量子效率; ∆F/Fm′: PSⅡ实际光化学量子效率; ETR:表观电子传递速率; NPQ:非光化学猝灭系数; qP:光化学猝灭系数。CK:自然光照处理; T:遮光70%处理。数据表示为平均值±标准差。相同品种同一处理之间不同小写字母表示5%水平差异显著。Fo: initial fluorescence under dark adaptation; Fm: maximum fluorescence under dark adaptation; Fv/Fm: maximal photochemical quantum efficiency under dark adaptation; Fv′/Fm′: maximum photochemical efficiency of PSⅡ under light adaptation; ∆F/Fm′: actual photochemical quantum efficiency of PSII under light adaptation; ETR: apparent electron transfer rate; NPQ: non-photochemical quenching; qP: photochemical quenching. CK: normal light; T: light decreased by 70%. ‘Lishu 6’ is low-light sensitive cultivar, ‘Zhongshu 20’ is low-light tolerant cultivar. The data is expressed as mean ± standard deviation. Values followed by different lowercase letters are significantly different at 5% level.
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表 4 遮阴对马铃薯不同弱光耐受性品种光能分配的影响
Table 4 Effect of shading on energy distribution of potato cultivars with different low-light tolerance
光能分配参数
energy distribution parameter丽薯6号Lishu 6 中薯20Zhongshu 20 CK T CK T ФPSII 0.29±0.04a 0.20±0.01b 0.32±0.03a 0.20±0.02b ФNPQ 0.45±0.05b 0.51±0.02a 0.43±0.03b 0.52±0.02a Фf, d 0.26±0.02b 0.29±0.01a 0.25±0.01b 0.28±0.28a ФPSⅡ:光化学耗散比例; ФNPQ:非光化学猝灭耗散比例; Фf, d:荧光耗散比例。CK:自然光照处理; T:遮光70%处理。数据表示为平均值±标准差。不同小写字母表示5%水平差异显著。ФPSⅡ: quantum yield of photochemical dissipation; ФNPQ: quantum yield of non-photochemical quenching dissipation; Фf, d: quantum yield of fluorescence quenching dissipation. CK: normal light; T: light decreased by 70%. ‘Lishu 6’ is low-light sensitive cultivar, ‘Zhongshu 20’ is low-light tolerant cultivar. The data is expressed as mean ± standard deviation. Values followed by different lowercase letters are significantly different at 5% level.
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