长期结实和不结实红松针叶光合生理参数的差异

doi:10.12302/j.issn.1000-2006.202203032

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南京林业大学学报（自然科学版） ›› 2023, Vol. 47 ›› Issue (3): 137-146.doi: 10.12302/j.issn.1000-2006.202203032

长期结实和不结实红松针叶光合生理参数的差异

林强¹(), 陆天宇¹^,², 沈海龙¹^,^*(), 王元兴³, 张鹏¹

1.林木遗传育种国家重点实验室,东北林业大学林学院,黑龙江哈尔滨 150040
2.辽宁省农业科学院,辽宁沈阳 110161
3.吉林省林业科学研究院,吉林长春 130031

收稿日期:2022-03-15 修回日期:2022-11-12 出版日期:2023-05-30 发布日期:2023-05-25
通讯作者: 沈海龙
基金资助:
国家重点研发计划(2017YFD0600601)

Analysis of needle photosynthetic index characteristics for long period seed setting and non-setting trees of Pinus koraiensis

LIN Qiang¹(), LU Tianyu¹^,², SHEN Hailong¹^,^*(), WANG Yuanxing³, ZHANG Peng¹

1. State Key Laboratory of Forest Genetics and Tree Breeding,School of Forestry,Northeast Forestry University, Harbin 150040,China
2. Liaoning Academy of Agricultural Sciences,Shenyang 110161,China
3. Jilin Province Academy of Forestry Science,Changchun 130031, China

Received:2022-03-15 Revised:2022-11-12 Online:2023-05-30 Published:2023-05-25
Contact: SHEN Hailong

摘要/Abstract

摘要：

【目的】选择营养生长基本一致而结实状况差异很大的红松(Pinus koraiensis)个体,分析其光合生理指标差异情况,为红松果材兼用培育提供科学依据。【方法】以吉林省露水河林业局种子园内30年生红松为研究对象,通过对不同结实特性及环境差异下红松个体光合指标进行连续观测,分析其生长季内叶绿素含量、比叶面积(SLA)、光响应参数、非结构性碳水化合物(NSC)含量等指标的变化情况。【结果】①不同结实状况红松叶绿素a(Chl a)、叶绿素b(Chl b)、类胡萝卜素(Car)和总叶绿素(Chl T)含量存在显著差异。在生长季期间,不结实红松中的Chl a、Chl b、Car和Chl T含量显著高于结实红松,而叶绿素a与b质量比(Chl a/b)相反,除7月外SLA均表现为结实红松>不结实红松。两种结实特性红松各指标在不同月份之间差异显著。两种结实特性红松叶绿素含量和比叶面积变化趋势一致,Chl a、Chl b、Car和Chl T含量呈下降趋势,而SLA则上升,Chl a/b先下降后上升;②两种结实特性的红松只有光饱和点(LSP)存在显著差异,表观量子效率(AQY)、光补偿点(LCP)、最大净光合速率(P_n,max)、暗呼吸速率(R_d)等均不显著。在整个生长季,除LCP外,两种结实特性红松各指标变化趋势基本一致,AQY和R_d表现为上升趋势,P_n,max、LCP、LSP则表现为下降趋势。除5月外,其他月份两种结实特性红松针叶AQY均为结实红松<不结实红松,R_d则相反,P_n,max和LSP表现为结实红松>不结实红松,LCP在5、7月份中为结实红松<不结实红松,6、8月份为结实红松>不结实红松,9月份趋于持平。同时,LCP和LSP随冠层升高而逐渐增大,且各指标在月份之间差异显著;③两种结实特性的红松个体针叶中可溶性糖、淀粉和NSC含量之间差异不显著,在生长季均表现为下降趋势且月份之间差异显著,可溶性糖和淀粉含量在不同生境下差异显著,针叶可溶性糖和NSC含量除4、8月外,均表现为结实红松>不结实红松,淀粉含量除6月外均表现为结实红松<不结实红松。【结论】吉林省红松林区的结实红松具有更高的光合潜力和物质消耗能力,因此在光照条件下会同化更多的碳,并提高了能量转换效率,以便在生长季存储更多的淀粉和NSC为果实发育提供营养支撑。

关键词: 红松, 光响应参数, 非结构性碳水化合物, 季节动态

Abstract:

【Objective】Photosynthetic products form the basis for forest growth and fruit development. This study analyzed the differences in photosynthetic physiological indexes of Korean pine (Pinus koraiensis) individuals that were largely similar in terms of their vegetative growth patterns, but significantly different with regards to seed setting. This study was conducted to provide a scientific basis for the optimum cultivation of the fruit and wood of P. koraiensis. 【Method】Thirty-year-old Pinus koraiensis in the seed orchard of the Lushuihe Forestry Bureau was selected as the research site, and the dynamic changes in chlorophyll, specific leaf area (SLA), light response parameters, and non-structural carbohydrates (NSC) during the growing season were assessed. 【Result】(1) There were significant differences in chlorophyll a (Chl a), chlorophyll b (Chl b), chlorophyll carotenoids, and total chlorophyll (Chl T) under different seed setting conditions. Specifically, Chl a, Chl b, Car and Chl T of non-fruiting P. koraiensis were significantly higher than those of fruiting Pinus koraiensis during the growing season, but chlorophyll a/b (Chl a/b) showed the opposite trend. However, the indices of the two seed-setting properties of P. koraiensis showed significant differences across different months. Chl a, Chl b, Car and Chl T decreased, SLA increased, and Chl a/b decreased at first and then increased. (2) The light saturation point (LSP) was significantly different between the two seed-setting properties of P. koraiensis; apparent quantum efficiency (AQY), light compensation point (LCP), maximum net photosynthetic rate (P_n,max) and dark respiration rate (R_d) were not significantly different. Except for the LCP, AQY and R_d of the two seed-setting properties of P. koraiensis showed an increased during the entire growing season. On the other hand, P_n,max, LCP, and LSP decreased. Two types of seed-setting characteristics of P. koraiensis needles were noted. Specifically, AQY showed less seed-setting than non-seed-setting in all months except May, but R_d, P_n,max and LSP showed the opposite trend. In addition, LCP in May and July is less seed-setting than non-seed-setting, and in June and August is more seed-setting than non-seed-setting. At the same time, LCP and LSP increased gradually with an increase in canopy, and the difference in each index month was significant. (3) There was no significant difference between soluble sugar, starch and NSC in the needles of the two seed-setting P. koraiensis individuals. However, the concentrations of soluble sugars and NSC in conifers were higher than those in non-conifers, except in April and August. In addition, the starch concentration was stable except for in June. 【Conclusion】P. koraiensis has higher photosynthetic potential and material consumption capacity, so it assimilates more carbon under light conditions and improves energy conversion efficiency; this results in it storing more starch and NSC during the growing season, providing nutritional support for fruit development.

Key words: Pinus koraiensis (Korean pine), photosynthetic parameters, non-structural carbohydrates, seasonal dynamics

中图分类号:

S791.247

林强,陆天宇,沈海龙,等. 长期结实和不结实红松针叶光合生理参数的差异[J]. 南京林业大学学报（自然科学版）, 2023, 47(3): 137-146.

LIN Qiang, LU Tianyu, SHEN Hailong, WANG Yuanxing, ZHANG Peng. Analysis of needle photosynthetic index characteristics for long period seed setting and non-setting trees of Pinus koraiensis[J].Journal of Nanjing Forestry University (Natural Science Edition), 2023, 47(3): 137-146.DOI: 10.12302/j.issn.1000-2006.202203032.

图/表 8

表1

表2

表3

图1

表4

表5

不同结实特性红松光合指标间差异"

月份 month	结实特性 fruiting characteristic	生境 habitat	冠层部位 position	表观量子效率/ (mol·mol^-1) AQY	参数/(μmol·m^-2·s^-1) indices
月份 month	结实特性 fruiting characteristic	生境 habitat	冠层部位 position	表观量子效率/ (mol·mol^-1) AQY	P_n,max	R_d	LSP	LCP
5	不结实NF		上部H	0.006 1 a	7.829 a	1.428 a	1 515.924 a	31.888 a
		林缘E	中部M	0.006 9 a	8.951 a	0.471 a	1 394.492 a	22.189 a
			下部L	0.007 7 a	9.143 a	0.577 a	1 294.542 a	26.885 a
			上部H	0.005 5 a	7.294 a	1.300 a	1 560.047 a	16.859 a
		林内I	中部M	0.004 5 a	5.471 a	0.696 a	1 385.010 a	10.846 a
			下部L	0.006 1 a	5.909 a	1.442 a	1 203.548 a	7.704 a
	结实F		上部H	0.005 7 a	9.005 a	0.736 a	1 709.400 a	10.825 a
		林缘E	中部M	0.006 8 a	8.830 a	0.811 a	1 459.760 a	19.755 a
			下部L	0.007 6 a	7.859 a	0.741 a	1 150.131 a	4.720 a
			上部H	0.005 3 a	7.053 a	1.158 a	1 583.686 a	17.813 a
		林内I	中部M	0.006 8 a	6.051 a	1.429 a	1 258.529 a	12.476 a
			下部L	0.004 7 a	5.844 a	0.899 a	1 429.791 a	9.562 a
6	不结实NF		上部H	0.006 4 a	8.138 a	0.755 a	1 386.456 a	10.476 a
		林缘E	中部M	0.006 6 a	7.130 a	0.965 a	1 236.517 a	13.029 a
			下部L	0.005 5 a	6.247 a	0.503 a	1 225.014 a	17.576 a
			上部H	0.005 9 a	8.003 a	0.822 a	1 498.790 a	13.471 a
		林内I	中部M	0.006 1 a	6.968 a	0.945 a	1 296.181 a	17.901 a
			下部L	0.008 4 a	7.255 a	2.174 a	1 137.886 a	25.477 a
	结实F		上部H	0.005 7 a	6.376 a	1.489 a	1 370.776 a	17.820 a
		林缘E	中部M	0.006 1 a	7.224 a	0.810 a	1 329.782 a	15.230 a
			下部L	0.008 2 b	7.926 a	1.080 a	1 108.502 a	13.471 a
			上部H	0.006 4 a	8.860 a	1.300 a	1 633.439 a	20.468 a
		林内I	中部M	0.004 8 a	6.878 a	1.106 a	1 706.517 b	21.993 a
			下部L	0.005 8 a	6.619 a	0.805 a	1 283.541 a	12.715 a
7	不结实NF		上部H	0.004 4 a	5.235 a	1.371 a	1 494.411 a	12.154 a
		林缘E	中部M	0.006 7 a	7.302 a	1.686 a	1 345.247 a	20.495 a
			下部L	0.009 4 a	8.474 a	1.312 a	1 036.955 a	12.154 a
			上部H	0.006 1 a	6.971 a	1.334 a	1 351.682 a	15.164 a
		林内I	中部M	0.006 8 a	7.114 a	1.421 a	1 265.974 a	12.860 a
			下部L	0.007 2 a	7.044 a	0.357 a	1 034.286 a	11.761 a
	结实F		上部H	0.006 9 b	8.333 a	1.411 a	1 481.770 a	20.327 a
		林缘E	中部M	0.006 2 a	7.184 a	1.665 a	1 575.536 a	13.945 a
			下部L	0.006 4 a	6.880 a	1.558 a	1 366.255 a	11.013 a
			上部H	0.007 2 a	5.784 a	1.042 a	1 169.053 a	14.363 a
		林内I	中部M	0.006 0 a	7.208 a	0.989 a	1 392.336 a	13.222 a
			下部L	0.066 1 a	7.453 a	0.895 a	1 140.826 a	11.442 a
8	不结实NF		上部H	0.006 7 a	8.414 a	1.554 a	1 527.547 a	17.009 a
		林缘E	中部M	0.006 7 a	7.091 a	1.417 a	1 326.158 a	13.702 a
			下部L	0.005 9 a	5.754 a	1.388 a	1 235.839 a	5.222 a
			上部H	0.008 1 a	8.226 a	1.261 a	1 211.409 a	22.293 a
		林内I	中部M	0.006 7 a	6.724 a	0.855 a	1 214.098 a	9.285 a
			下部L	0.008 2 a	4.969 a	0.627 a	668.076 a	8.366 a
	结实F		上部H	0.007 6 a	10.576 a	1.747 a	1 660.361 a	22.735 a
		林缘E	中部M	0.006 3 a	7.805 a	1.455 a	1 508.528 a	11.750 a
			下部L	0.005 4 a	5.9670 a	1.475 a	1 436.135 a	10.558 a
			上部H	0.006 3 a	7.530 a	0.947 a	1 477.218 a	12.959 a
		林内I	中部M	0.007 8 a	6.255 a	0.984 a	1 110.388 a	8.265 a
			下部L	0.008 6 a	4.013 a	0.965 a	577.589 a	4.773 a
9	不结实NF		上部H	0.005 5 a	6.097 a	1.084 a	1 312.425 a	17.509 a
		林缘E	中部M	0.006 7 a	6.819 a	1.290 a	1 275.987 a	13.702 a
			下部L	0.006 2 a	4.846 a	0.689 a	956.368 a	5.222 a
			上部H	0.007 7 a	8.066 a	1.683 a	1 264.802 a	22.793 a
		林内I	中部M	0.008 7 a	7.311 a	1.054 a	966.799 a	9.285 a
			下部L	0.007 7 a	6.058 a	0.991 a	1 008.773 a	8.366 a
	结实F		上部H	0.006 5 a	7.266 a	1.502 a	1 327.393 a	22.235 a
		林缘E	中部M	0.005 8 a	6.469 a	1.342 a	1 359.732 a	11.750 a
			下部L	0.006 4 a	5.782 a	1.268 a	1 323.383 a	10.558 a
			上部H	0.006 0 a	5.581 a	1.440 a	1 178.921 a	12.459 a
		林内I	中部M	0.008 7 a	6.217 a	0.980 a	898.010 a	8.265 a
			下部L	0.008 6 a	8.119 a	0.887 a	1 132.021 a	4.772 a

表5

图2

图3

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生境 habitat	样木分组 tree group	林龄/a stand age	样木编号 tree No.	树高/ m tree height	胸径/ cm DBH	平均冠幅/m average crown width
林缘(E) forest edge	E1	29	不结实NF1	9.9	22.6	7.2
		29	结实F2	10.3	26.5	8.5
	E2	29	不结实NF3	11.0	32.1	7.9
		29	结实F4	10.8	28.1	9.3
	E3	29	不结实NF5	10.3	27.1	8.6
		29	结实F6	10.8	31.5	9.0
林内(I) inside stand	I4	30	不结实NF7	11.3	27.3	6.4
		30	结实F8	11.9	24.6	5.7
	I5	31	不结实NF9	12.3	26.8	7.3
		31	结实F10	11.8	29.1	9.8
	I6	29	不结实NF11	10.4	26.6	7.7
		29	结实F12	11.1	29.8	8.7

参数 parameters	结实特性 (FC) fruiting characteristic	月份 (M) month	生境 (H) habitat	FC× M	FC × H
叶绿素a含量Chl a content	9.976^***	42.336^***	3.436	0.158	1.619
叶绿素b含量Chl b content	10.017^***	20.301^***	2.133	0.219	1.977
类胡萝卜素含量Car content	8.937^***	25.135^***	4.020^*	0.426	0.709
总叶绿素含量Chl T content	10.237^***	37.467^***	3.196	0.166	1.746
叶绿素a/b Chla/b	1.494	9.861^***	0.288	0.468	0.764
比叶面积 SLA	0.549	38.755^***	0.864	0.246	3.663
可溶性糖含量 soluble sugar content	2.974	22.187^***	5.545^***	1.105	1.189
淀粉含量starch content	0.036	276.690^***	6.964^***	0.354	1.999
非结构性碳水化合物含量NSC content	1.273	205.219^***	0.248	0.512	0.133

月份 month	结实特性 fruiting characteristic	生境 habitat	含量/(mg·g^-1) content				Chl a/b	比叶面积/ (cm²·g^-1) SLA
月份 month	结实特性 fruiting characteristic	生境 habitat	Chl a	Chl b	Car	Chl T	Chl a/b	比叶面积/ (cm²·g^-1) SLA
5	不结实NF	林缘E	1.53 A	0.36 A	0.54 A	1.89 A	4.23 A	38.73 A
		林内I	1.27 a	0.31 a	0.45 a	1.58 a	4.12 a	38.16 a
	结实F	林缘E	1.40 A	0.34 A	0.50 A	1.74 A	4.17 A	38.89 A
		林内I	1.22 a	0.29 a	0.44 a	1.51 a	4.20 a	38.91 a
6	不结实NF	林缘E	1.40 A	0.35 A	0.50 A	1.75 A	4.03 A	41.42 A
		林内I	1.24 a	0.30 a	0.44 a	1.55 a	4.12 a	43.51 a
	结实F	林缘E	1.30 A	0.31 A	0.46 A	1.61 A	4.23 B	41.87 A
		林内I	1.23 a	0.31 a	0.44 a	1.54 a	4.07 a	46.01 a
7	不结实NF	林缘E	1.25 A	0.33 A	0.44 A	1.57 A	3.89 A	49.39 A
		林内I	1.23 a	0.31 a	0.45 a	1.54 a	3.98 a	51.84 a
	结实F	林缘E	1.17 A	0.30 A	0.44 A	1.47 A	3.96 A	49.50 A
		林内I	1.10 a	0.28 a	0.40 a	1.38 a	3.91 a	49.62 a
8	不结实NF	林缘E	1.05 A	0.29 A	0.39 A	1.34 A	3.68 A	55.59 A
		林内I	1.01 a	0.26 a	0.36 a	1.26 a	3.95 a	49.07 a
	结实F	林缘E	0.88 B	0.24 A	0.32 B	1.12 B	3.77 B	49.54 A
		林内I	1.03 a	0.27 a	0.37 a	1.29 a	3.89 a	57.53 b
9	不结实NF	林缘E	0.92 A	0.23 A	0.37 A	1.16 A	3.98 A	55.15 A
		林内I	0.97 a	0.26 a	0.39 a	1.23 a	3.71 a	52.88 a
	结实F	林缘E	0.78 B	0.19 B	0.32 B	0.97 B	4.11 A	54.74 A
		林内I	0.88 a	0.23 a	0.33 b	1.11 a	3.88 b	56.36 a

参数 parameters	结实特性 (FC) fruiting characteristic	月份 (M) month	冠层 (C) canopy	FC×M	FC×C
表观量子效率AQY	0.223	1.331	2.262	0.058	0.113
最大净光合速率P_n,max	0.045	0.872	2.631	0.008	0.063
暗呼吸速率R_d	0.291	1.348	2.328	0.067	0.036
光饱和点LSP	4.185^*	4.275^**	17.978^***	0.094	0.234
光补偿点LCP	0.429	2.000	10.848^***	0.986	0.625

长期结实和不结实红松针叶光合生理参数的差异

Analysis of needle photosynthetic index characteristics for long period seed setting and non-setting trees of Pinus koraiensis

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