多倍化对青钱柳叶形态、光合性能和次生代谢产物积累的影响

doi:10.11707/j.1001-7488.LYKX20230322

摘要/Abstract

摘要：

目的: 比较分析青钱柳二倍体和四倍体的叶形态、光合性能和次生代谢物积累的差异，以期为青钱柳叶用林良种选育提供理论参考。方法: 以鹤峰和清凉峰2个种源的成年植株为试验材料，分别测定二倍体和四倍体的叶形态指标、解剖结构、气孔参数，比较光合生理的相关参数，监测叶次生代谢产物含量的季节动态及差异。结果: 1）四倍体叶长、叶宽、叶面积分别为10.94 cm、4.57 cm、118.84 cm²，比二倍体分别高出19.5%、61.4%、94.2%，而比叶质量无显著差异；四倍体叶解剖结构各部分、气孔形态均显著大于二倍体，气孔密度为158.36 个·mm^?2，比二倍体少43.8%。2）四倍体叶类胡萝卜素含量显著高于二倍体，叶绿素a、叶绿素b无显著差异；通过对光响应曲线拟合得到二倍体和四倍体的最大净光合速率（P_nmax）、暗呼吸速率（R_d）、光补偿点（LCP）、光饱和点（LSP）分别为8.367、0.997、25.905、1 527.96 μmol·m^?2s^?1和13.797、1.573、32.905、1 311.36 μmol·m^?2s^?1；瞬时光合测定发现四倍体净光合速率、胞间CO₂、蒸腾速率均显著大于二倍体；气孔导度、蒸气压亏缺则差异不显著。3）二种倍性植株叶内主要活性物质的积累高峰均出现在5月和9月；三萜化合物和黄酮含量在主要的收获季节（5月和9月）倍性间均无显著差异；四倍体多酚含量（13.64 mg·g^?1）仅于5月显著高于二倍体（13.29 mg·g^?1），但四倍体多糖含量在整个生长季节均显著高于二倍体，5月和9月的含量分别达6.80和8.58 mg·g^?1。结论: 青钱柳四倍体叶形态、叶解剖结构和光合生理特性均显著优于二倍体，促进了四倍体叶生物量积累；四倍体在5月份的多酚含量和叶多糖含量全生长季显著高于二倍体，黄酮、三萜含量倍性间在主要的收获季节均无显著差异。综合评价认为，四倍体是低海拔地区叶用青钱柳人工林生产的主要栽培材料，可获得更高的生物活性物质产量。

关键词: 四倍体, 叶形态, 叶解剖结构, 光合生理, 次生代谢产物

Abstract:

Objective: Cyclocarya paliurus is a unique medicinal plant in China, and its diploid and autotetraploid individuals coexist in natural populations. This study aims to compare and analyze the differences in leaf morphology, photosynthetic performance and secondary metabolite accumulation between diploidy and tetraploidy, in order to provide a basis for elite genotype selection for leaf-use plantations. Method: The leaf morphology, anatomical structure, and stomatal parameters in diploid and tetraploid adult plants from Hefeng, Hubei Province and Qingliangfeng, Anhui Province were measured. Their photosynthetic physiological parameters were also determined. The seasonal dynamics and differences of secondary metabolite profiles between two ploidies were compared. Result: 1) The length, width, and area of tetraploid leaves were 10.94 cm, 4.57 cm, and 118.84 cm², respectively, which were 19.5%, 61.4%, and 94.2% greater than those of diploid leaves. However, there was no significant difference in specific leaf mass. In similar, various tissues in leaf anatomical structure and stomatal morphology in tetraploidy were also significantly larger than that in diploidy. However, the stomatal density in tetraploidy was 158.36 stomata·mm^?2, which was 43.8% less than that of diploidy. 2) The carotenoid content in tetraploidy was significantly higher than that in diploidy, and there were no significant differences in chlorophyll a and chlorophyll b between the two ploidies. The parameters obtained from fitted curves of light response showed that maximum net photosynthetic rate (P_nmax), dark respiration rate (R_d), light compensation point (LCP), and light saturation point (LSP) in diploid were 8.367, 0.997, 25.905, and 1 527.96 μmol·m^?2s^?1, while were 13.797, 1.573, 32.905, and 1 311.36 μmol·m^?2s^?1 in tetraploidy, respectively. Measurement for instantaneous photosynthesis revealed that net photosynthetic rate, intercellular CO₂ concentration, and transpiration rate in tetraploidy were significantly higher than those in diploidy. There were no significant differences in stomatal conductance and vapor pressure deficit. 3) The accumulation peaks of major bioactive substances in leaves of two ploidies all occurred in May and September, respectively. There were no significant differences in the content of triterpenoids and flavonoids in two harvest seasons (May and September) between ploidies. The polyphenol content of tetraploidy (13.64 mg·g^?1) was significantly higher than that of diploidy (13.29 mg·g^?1) only in May, while the polysaccharide content of tetraploidy remained consistently higher than that of diploidy throughout the entire growing season, reaching 6.80 mg·g^?1 and 8.58 mg·g^?1 in May and September, respectively. Conclusion: The leaf morphology, anatomical structure and photosynthetic physiological characteristics in tetraploidy are significantly better than those of diploidy, thus promoting its leaf biomass production. The polyphenol content in May and polysaccharide content in whole growth seasons in tetraploidy are significantly higher than those in diploidy, but there is no significant difference in flavonoids and triterpenes contents in the main harvest seasons between two ploides. Comprehensively, the tetraploid is the candidate material for leaf-use plantations in low-altitude areas, and can obtain higher yields of bioactive substances in practice.

Key words: tetraploid, leaf morphology, leaf anatomy, photosynthetic physiology, secondary metabolite

中图分类号:

S722.3+5

王舒扬,田力,周顺陶,储月娥,梅迪,袁佳秋,余延浩,洑香香. 多倍化对青钱柳叶形态、光合性能和次生代谢产物积累的影响[J]. 林业科学, 2024, 60(8): 120-131.

Shuyang Wang,Li Tian,Shuntao Zhou,Yuee Chu,Di Mei,Jiaqiu Yuan,Yanhao Yu,Xiangxiang Fu. Effects of Polyploidization on Leaf Morphology, Photosynthetic Performance and Accumulation of Secondary Metabolites in Cyclocarya paliurus[J]. Scientia Silvae Sinicae, 2024, 60(8): 120-131.

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种源Provenance	二倍体Diploidy (2n)			四倍体Tetraploidy (4n)
种源Provenance	家系Family	样株编号 Sample No	海拔 Altitude/m	家系Family	编号 Number	海拔 Altitude/m
湖北鹤峰 Hefeng, Hubei (HF)	HF5#	NW19-8	1 400~1 600	HF3#	NW16-2	830~1 400
	HF5#	NW19-9		HF3#	NW16-6
	HF11#	NW10-1		HF8#	NW19-6
	HF11#	NW10-2		HF8#	NW20-5
	HF2#	NW3-15		HF10#	NW3-1**
		NW3-17**			NW3-2**
		NW3-16**			NW3-5**
		NW4-15**			NW4-1*
		NW4-16*			NW4-3
安徽清凉峰 Qingliangfeng, Anhui (QLF)	QLF2#	NW9-7*	1 100~1 450	QLF4#	NW19-11	650~1 100
		NW9-8*			NW19-12
		NW9-10			NW20-10*
		NW9-11			NW20-11*
		NW10-10			NW21-12
		NW10-11*			NW21-13
		NW10-12			NW 22-11*
		NW10-13			NW22-12*
		NW10-14*			NW22-13

种源Provenance	倍性 Ploidy	叶绿素a Chlorophyll a/(mg·g^?1)	叶绿素b Chlorophyll b/(mg·g^?1)	总叶绿素 Total chlorophyll/(mg·g^?1)	类胡萝卜素 Carotenoid/(mg·g^?1)
HF	2n	1.71±0.22Aa	0.82±0.13Aa	2.53±0.34Aa	0.30±0.04Ba
HF	4n	1.89±0.16Aa	0.84±0.06Aa	2.73±0.22Aa	0.36±0.02Aa
QLF	2n	1.82±0.20Aa	0.83±0.14Aa	2.65±0.33Aa	0.32±0.03Aa
QLF	4n	2.01±0.26Aa	0.88±0.11Aa	2.89±0.36Aa	0.36±0.02Aa
Total	2n	1.76±0.22Aa	0.82±0.14Aa	2.59±0.34Aa	0.31±0.04Ba
Total	4n	1.95±0.23Aa	0.86±0.09Aa	2.81±0.31Aa	0.36±0.02Aa

种源 Provenance	倍性 Ploidy	净光合速率 P_n/(μmol·m^?2s^?1)	胞间CO₂浓度 C_i/(μmol·mol^?1)	蒸腾速率 Tr/(mmol·m^?2s^?1)	气孔导度 G_s/(mol·m^?2s^?1)	蒸气压亏缺 VPD/kPa
HF	2n	8.13±0.20Aa	217.67±19.78Ba	1.47±0.07Ba	143.00±12.48Aa	18.43±3.00Aa
HF	4n	10.83±0.85Aa	247.92±3.59Aa	1.88±0.06Aa	124.50±9.85Ab	18.08±2.62Ab
QLF	2n	8.08±0.24Ba	191.00±2.11Ba	1.52±0.04Ba	135.75±14.39Aa	18.15±0.90Ba
QLF	4n	10.92±0.27Aa	238.42±5.26Ab	1.77±0.05Ab	140.50±3.99Aa	24.47±1.72Aa
Total	2n	8.11±0.22Ba	204.33±20.15Ba	1.49±0.06Ba	139.38±13.95Aa	18.29±2.22Aa
Total	4n	10.88±0.63Aa	243.17±6.55Aab	1.82±0.08Aab	132.50±10.98Aab	21.28±3.88Aab

种类Species	5月 May	6月 June	7月 July	8月 August	9月 September
2n HF	12.88±1.12Ab	12.41±0.94Abc	11.73±1.02Ac	12.64±0.93ABbc	13.93±0.93ABa
4n HF	13.70±0.73Aa	12.95±0.82Aab	12.22±0.71Ab	12.65±0.65ABb	13.46±0.93Ba
2n QLF	12.73±1.54Aab	12.08±1.48Abc	11.38±1.03Ac	11.84±1.12Bbc	13.70±0.94ABa
4n QLF	13.57±1.46Aab	12.76±1.15Abc	12.05±0.73Ac	12.89±0.90Abc	14.47±0.86Aa

种类Species	5月 May	6月 June	7月 July	8月 August	9月 September
2n HF	16.26±1.58Ab	15.11±1.65Abc	14.34±1.10Ac	16.08±1.10ABb	18.35±1.28Aa
4n HF	15.80±1.24Ab	14.38±1.57Ab	14.41±1.53Ab	15.08±1.20ABb	17.21±1.55ABa
2n QLF	15.05±1.84Aa	14.51±1.98Aa	12.51±1.29Bb	14.79±1.78Ba	16.08±1.54Ba
4n QLF	15.58±0.81Abc	15.68±0.83Abc	14.64±1.63Ac	16.47±1.38Ab	17.85±0.80Aa