美洲黑杨无性系苗木对不同光周期处理的生长与生理响应

doi:10.11707/j.1001-7488.LYKX20220634

摘要/Abstract

摘要：

目的: 研究不同光周期处理下美洲黑杨无性系苗木生长性状表现，筛选高效评价美洲黑杨光周期敏感性的性状指标，明确光周期对不同敏感度美洲黑杨生理状态的影响。方法: 以26个美洲黑杨无性系为研究材料，利用人工气候室设置2种光周期处理（光照16 h/暗期8 h，光照8 h/暗期16 h），通过对不同光周期处理下美洲黑杨12个生长性状多时间点连续观测分析，利用隶属函数法和主成分分析，获取美洲黑杨生长性状光周期敏感性综合评价D值。采用回归分析和通径分析，筛选适合美洲黑杨光周期敏感性评价的直接效应指标，选择不同光周期敏感度美洲黑杨无性系进行与生长监测同步的生理指标测定分析。结果: 1）方差分析结果表明，美洲黑杨12个生长性状在不同光周期处理下存在显著差异（P<0.05），短日照下美洲黑杨的叶长、叶宽、叶面积和比叶面积均显著大于长日照下的观测值，而苗高生长净值、地径生长净值、叶片干质量、茎段鲜质量、根系鲜质量、茎段干质量、根系干质量和节间数观测值均显著小于长日照处理。2）美洲黑杨光周期敏感性综合评价D值分析结果表明，茎段鲜质量、节间数、地径生长净值、苗高生长净值等指标对光周期敏感度综合评价值的直接效应较大（0.113~0.338），可以作为评价美洲黑杨光周期敏感性的性状指标，其中苗高生长净值更适合生产实际应用。3） 12个性状的光周期敏感度综合比较发现，相同种源内不同无性系光周期敏感度不同，26个美洲黑杨无性系中光周期综合敏感度最高的为LS8、TN3、WA1，最低的为QB7、LS5、TN6，结合苗高生长表现评选出3个高生长高光周期敏感度无性系LS9、LS4、TN4，3个低生长低敏感度无性系QB7、QB2、QB4。4）高生长高敏感度无性系在长日照处理下的Rubisco活性、磷酸丙糖异构酶活性、果糖-1,6-二磷酸醛缩酶活性均随处理时间增加而增强，总叶绿素含量呈先下降后上升的变化趋势，谷氨酰胺合成酶活性随处理时间增加而逐渐降低，在短日照处理下其Rubisco活性先增强后降低，总叶绿素含量逐渐降低；低生长低敏感度无性系的Rubisco活性和总叶绿素含量随处理时间增加而逐渐降低，在短日照处理下，其果糖-1,6-二磷酸醛缩酶活性先降低后增强，Rubisco活性总体随时间增加逐渐降低；不同美洲黑杨无性系在不同光周期处理下的酶活性和总叶绿素含量变化趋势不同。结论: 美洲黑杨生长性状表现受光周期显著影响，其中苗高生长净值、地径生长净值、茎段鲜质量和节间数都表现较强光周期敏感性，可作为美洲黑杨光周期敏感性评价的性状指标，结合苗高生长净值划分出高生长高敏感度和低生长低敏感度类型无性系。不同光周期敏感度的美洲黑杨对光周期变化的生理响应有明显差异，高敏感度无性系的果糖-1,6-二磷酸醛缩酶活性、谷氨酰胺合成酶活性对长日照更敏感，不同敏感度美洲黑杨的Rubisco活性在长日照下表现相反的变化趋势，可能导致不同光周期敏感度美洲黑杨的生长差异。研究结果可为杨树及林木光周期响应机理研究提供有益参考，筛选出的不同光周期敏感度美洲黑杨可为培育广适性杨树新品种提供种质。

关键词: 美洲黑杨, 光周期敏感性, 生长, 生理指标

Abstract:

Objective: This study aims to investigate the performance of plant growth traits of Populus deltoides clones under different photoperiod treatments, and screen the traits that can effectively evaluate photoperiod sensitivity of P. deltoides, so as to clarify the effect of photoperiods on physiological status of P. deltoides clones with different sensitivity. Method: In this study, 26 clones of P. deltoides were used as the research materials, and subjected to two photoperiod treatments (light 16 h/dark period 8 h, light 8 h/dark period 16 h) in artificial climate chambers. A total of 12 growth traits of P. deltoides were continuously observed at multiple time points under different photoperiod treatments, and the subordinate function method and principal component analysis were used to obtain the comprehensive evaluation index D value of photoperiod sensitivity of growth traits of P. deltoides. Regression analysis and path analysis were used to screen the direct effect indexes suitable for the evaluation of photoperiod sensitivity of P. deltoides, and the P. deltoides clones with different photoperiod sensitivity were selected for synchronously monitoring the growth and measuring the physiological indexes. Result: 1) The results of variance analysis showed that there were significant differences in 12 growth traits of P. deltoides under different photoperiod treatments (P<0.05). The leaf length, leaf width, leaf area and specific leaf area ofP. deltoides under short photoperiod were significantly greater than those under long photoperiod, while the net growth value of plant height, net ground diameter, dry weight of leaf, fresh weight of stem segment, fresh weight of root system, dry weight of stem segment, root dry weight and the observed values of node number were significantly lower than those under long photoperiod. 2) The analysis results of D value of comprehensive evaluation indexes for photoperiod sensitivity of P. deltoides showed that four indexes, including fresh weight of stem segment, number of nodes, net growth value of ground diameter and net growth value of plant height, had a greater direct effect on the comprehensive evaluation value of photoperiod sensitivity (0.113–0.338), which could be used as the character index for evaluating photoperiod sensitivity of P. deltoides. Among them, net growth value of plant height was more suitable for practical production. 3) The comprehensive comparison of photoperiod sensitivity of 12 characters showed that different clones in the same provenance had different photoperiod sensitivity. Among the 26 clones of P. deltoides, LS8, TN3 and WA1 had the highest photoperiod sensitivity, and QB7, LS5 and TN6 had the lowest photoperiod sensitivity. Three clones with high growth and high photoperiod sensitivity, LS9, LS4 and TN4, and three clones with low growth and low sensitivity, QB7, QB2 and QB4, were selected based on the growth performance of plants. 4) The Rubisco activity, triose phosphate isomerase activity and fructose-1,6-bisphosphate aldolase (FBA) activity of clones with high growth and high sensitivity under long photoperiod treatment increased with the increase of treatment time. The total chlorophyll content decreased first and then increased. The glutamine synthetase (GS) activity decreased gradually with the increase of treatment time. Under short photoperiod treatment, the Rubisco activity increased first and then decreased, and the total chlorophyll content decreased gradually. Rubisco activity and total chlorophyll content of clones with low growth and low sensitivity decreased gradually with the increase of treatment time. Under the treatment of short photoperiod, their FBA activity first decreased and then increased, and Rubisco activity generally decreased with the increase of treatment time. The changes of enzyme activities and total chlorophyll content of different P. deltoides clones were different under different photoperiod treatments. Conclusion: The growth characters of P. deltoides are significantly affected by photoperiod, among which the net growth value of plant height, net growth value of ground diameter, fresh weight of stem segments are more sensitive to photoperiod, which can be used as the character indicators for evaluating photoperiod sensitivity of P. deltoides. Combined with the net growth value of plant height, the clones can be divided into high growth sensitivity and low growth sensitivity types. The physiological responses of P. deltoides with different photoperiod sensitivities to photoperiod changes are significantly different. The FBA activity and GS activity of clones with high sensitivity are more sensitive to long photoperiod. The Rubisco activity of P. deltoides with different photoperiod sensitivities shows a reverse change trend under long photoperiod, which may lead to the growth differences of P. deltoides with different photoperiod sensitivities. The results can provide useful reference for the study of photoperiod response mechanism of poplar and forest trees, and the screened P. deltoides with different photoperiod sensitivity can provide germplasm for cultivating new poplar varieties with wide adaptability.

Key words: Populus deltoides, photoperiod sensitivity, growth, physiological index

中图分类号:

S722.3⁺3

李政宏,丁昌俊,张伟溪,张静,沈乐,张腾倩,丁密,苏晓华,吴钟亲,方发之. 美洲黑杨无性系苗木对不同光周期处理的生长与生理响应[J]. 林业科学, 2023, 59(3): 127-144.

Zhenghong Li,Changjun Ding,Weixi Zhang,Jing Zhang,Le Shen,Tengqian Zhang,Mi Ding,Xiaohua Su,Zhongqin Wu,Fazhi Fang. Growth and Physiological Response of Seedlings of Populus deltoides Clones to Different Photoperiods[J]. Scientia Silvae Sinicae, 2023, 59(3): 127-144.

图/表 13

表1

表2

表3

表4

表5

表6

美洲黑杨生长性状与光周期敏感综合指标D值的通径分析"

性状 Traits	与D值简单相关Simple relevant with D	直接效应Direct effect	间接效应 Indirect effect
性状 Traits	与D值简单相关Simple relevant with D	直接效应Direct effect	叶面积 Leaf area	叶宽 Leaf width	叶长 Leaf length	比叶面积 Specific leaf area	根系干质量 Root dry weight	根系鲜质量 Root fresh weight	叶片干质量 Leaf dry weight	茎段干质量 Stem dry weight	节间数 Internode number	茎段鲜质量 Fresh weight of stem segment	地径净值 Net growth of ground diameter	苗高净值 Net growth of plant height
叶面积 Leaf area	0.469	0.092		0.074	0.059	?0.004	?0.006	?0.011	0.033	0.070	0.052	0.066	0.05	?0.007
叶宽 Leaf width	0.431	0.084	0.412		0.242	0.042	?0.018	?0.042	0.082	0.196	0.170	0.179	0.072	?0.015
叶长 Leaf length	0.372	0.067	0.416	0.281		0.033	0.005	?0.005	0.134	0.116	0.045	0.070	0.117	?0.005
比叶面积 Specific leaf area	0.082	?0.008	0.249	0.223	0.150		?0.139	?0.110	?0.172	?0.049	0.286	?0.023	0.011	0.005
根系干质量 Root dry weight	0.288	0.086	?0.031	?0.028	0.007	?0.040		0.249	0.147	0.352	?0.189	0.201	?0.019	?0.028
根系鲜质量 Root fresh weight	0.261	0.076	?0.066	?0.069	?0.007	?0.035	0.275		0.105	0.363	?0.152	0.214	?0.069	?0.026
叶片干质量 Leaf dry weight	0.320	0.090	0.174	0.110	0.156	?0.044	0.132	0.086		0.180	?0.167	0.136	0.088	?0.016
茎段干质量 Stem dry weight	0.913	0.335	0.098	0.093	0.047	?0.004	0.111	0.104	0.063		0.298	0.874	0.040	?0.007
节间数 Internode number	0.600	0.228	0.106	0.122	0.028	0.039	?0.091	?0.066	?0.089	0.453		0.542	0.032	0.013
茎段鲜质量 Fresh weight of stem segment	0.902	0.338	0.092	0.086	0.029	?0.002	0.064	0.062	0.048	0.885	0.361		0.033	?0.004
地径净值 Net growth of ground diameter	0.722	0.329	0.139	0.084	0.117	0.002	?0.015	?0.048	0.075	0.098	0.052	0.080		0.048
苗高净值 Net growth of plant height	0.529	0.150	?0.03	?0.061	?0.017	0.004	?0.075	?0.063	?0.047	?0.064	0.074	?0.030	0.166

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种源编码Provenance coding	无性系数量Number of clones	种源地Provenance	经纬信息Longitude and latitude information
种源编码Provenance coding	无性系数量Number of clones	种源地Provenance	西经West longitude	北纬North latitude
LS	9	美国路易斯安那州Louisiana，USA	92°47'	31°51'
TN	7	美国田纳西州Tennessee，USA	89°32'	36°30'
OW	1	美国艾奥瓦州Iowa，USA	93°50'	41°52'
WA	1	美国华盛顿州Washington, USA	119°40'	46°12'
QB	8	加拿大魁北克省Quebec，Canada	71°59'	46°34'

性状Trait	短日照Short photoperiod	长日照Long photoperiod	均方Mean square	F值F value
苗高净值Net growth of plant height /cm	9.84±5.03b	16.94±10.30a	5 790.052	88.172
地径净值Net growth of ground diameter/cm	0.097±0.04b	0.15±0.05a	464 852.454	188.209
叶长Leaf length /mm	86.00±13.33a	78.97±14.33b	5 439.111	28.403
叶宽Leaf width /mm	70.00±9.66a	65.23±9.26b	2 499.004	27.922
叶柄长Petiole length /mm	47.70±9.40a	49.91±10.27a	536.809	5.534
叶面积Leaf area/ mm²	4 449.06±1320.21a	3 672.21±1 222.45b	66 385 172.956	41.012
叶鲜质量Fresh weight of leaves/g	0.74±0.29a	0.76±0.28a	0.034	0.414
叶干质量Leaf dry weight/g	0.21±0.09b	0.27±0.11a	0.417	41.978
比叶面积Specific leaf area/(mm²·g^?1）	22 423.34±5037.26a	14 181.16±3057.38b	7 251 559 673.888	419.064
茎段鲜质量Fresh weight of stem segment/g	3.49±1.37b	5.83±2.82a	240.622	48.913
根系鲜质量Root fresh weight/g	6.69±2.97b	9.18±3.81a	272.282	23.352
茎段干质量Stem dry weight/g	1.54±0.59b	3.05±1.33a	100.687	95.206
根系干质量Root dry weight/g	1.70±0.90b	2.70±1.12a	43.880	42.312
节间长Internode length/cm	1.97±0.36a	1.91±0.36a	0.186	1.425
节间数Internode number	19.84±2.96b	24.14±6.38a	811.841	32.820

性状Traits	主成分1Component 1	主成分2Component 2	主成分3Component 3	主成分4 Component 4
叶面积Leaf area	0.913	0.096	?0.354	?0.081
叶宽Leaf width	0.851	0.063	?0.242	?0.201
叶长Leaf length	0.772	0.151	?0.474	?0.011
比叶面积Specific leaf area	0.643	?0.538	0.110	?0.337
根系干质量Root dry weight	?0.218	0.888	?0.118	?0.058
根系鲜质量Root fresh weight	?0.265	0.844	?0.008	?0.141
叶片干质量Leaf dry weight	0.166	0.661	?0.501	0.258
茎段干质量Stem dry weight	0.400	0.651	0.608	0.065
节间数Internode number	0.557	?0.134	0.712	?0.018
茎段鲜质量Fresh weight of stem segment	0.423	0.533	0.687	0.084
地径净值Net growth of ground diameter	0.377	?0.004	?0.148	0.782
苗高净值Net growth of plant height	0.041	?0.317	0.138	0.777
特征值Eigenvalue	3.507	3.090	2.075	1.476
贡献率Variance contribution(%)	29.221	25.752	17.293	12.303
累计贡献率Accumulation variance contribution (%)	29.221	54.973	72.266	84.568

无性系Clone	隶属函数值Membership function value
无性系Clone	μ 1	μ 2	μ 3	μ 4
LS8	1.000	0.430	0.680	0.650
TN3	0.330	1.000	1.000	0.370
WA1	0.910	0.360	0.460	0.750
LS9	0.680	0.220	0.870	0.570
QB6	0.570	0.740	0.410	0.080
LS4	0.530	0.300	0.510	0.620
LS7	0.680	0.230	0.630	0.110
QB8	0.500	0.410	0.390	0.480
TN4	0.130	0.410	0.850	0.650
QB5	0.470	0.180	0.680	0.550
LS3	0.440	0.310	0.100	0.460
TN2	0.460	0.340	0.090	0.430
TN5	0.430	0.280	0.120	0.400
QB3	0.490	0.220	0.100	0.500
QB2	0.300	0.790	0.050	0.530
LS1	0.460	0.160	0.860	0.270
LS2	0.640	0.240	0.250	0.350
TN1	0.370	0.060	0.940	0.330
QB1	0.310	0.340	0.020	1.000
OW1	0.140	0.450	0.590	0.340
QB4	0.690	0.320	0.000	0.000
TN7	0.390	0.290	0.230	0.260
LS6	0.380	0.000	0.410	0.410
QB7	0.000	0.270	0.500	0.440
LS5	0.280	0.000	0.410	0.460
TN6	0.170	0.180	0.350	0.290

性状Traits	D	叶面积Leafarea	叶宽Leafwidth	叶长Leaflength	比叶面积Specificleaf area	根系干质量Root dryweight	根系鲜质量Root freshweight	叶片干质量Leaf dryweight	茎段干质量Stem dryweight	节间数Internodenumber	茎段鲜质量Fresh weight ofstem segment	地径净值Net growth ofground diameter	苗高净值Net growth ofplant heigh
D	1
叶面积Leaf area	0.469*	1
叶宽Leaf width	0.431*	0.878**	1
叶长Leaf length	0.372	0.886**	0.651**	1
比叶面积Specific leaf area	0.082	0.530*	0.518*	0.403	1
根系干质量Root dry weight	0.288	?0.067	?0.064	0.019	?0.484*	1
根系鲜质量Root fresh weight	0.261	?0.140	?0.160	?0.018	?0.423	0.954**	1
叶片干质量Leaf dry weight	0.320	0.372	0.256	0.419	?0.538**	0.460*	0.329	1
茎段干质量Stem dry weight	0.913**	0.208	0.215	0.127	?0.054	0.385	0.398	0.197	1
节间数Internode number	0.600**	0.227	0.284	0.075	0.477*	?0.315	?0.253	?0.278	0.496*	1
茎段鲜质量Fresh weight ofstem segment	0.902**	0.196	0.199	0.078	?0.025	0.223	0.237	0.151	0.969**	0.601**	1
地径净值Net growth ofground diameter	0.722**	0.296	0.194	0.315	0.029	?0.051	?0.185	0.235	0.107	0.087	0.089	1
苗高净值Net growth ofplant heigh	0.529*	?0.065	?0.142	?0.047	0.044	?0.261	?0.243	?0.147	?0.070	0.124	?0.033	0.446*	1