小叶杨×欧洲黑杨杂交F1代生长及叶片解剖结构杂种优势分析与抗旱性评价

doi:10.12302/j.issn.1000-2006.202310036

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南京林业大学学报（自然科学版） ›› 2025, Vol. 49 ›› Issue (1): 46-58.doi: 10.12302/j.issn.1000-2006.202310036

小叶杨×欧洲黑杨杂交F₁代生长及叶片解剖结构杂种优势分析与抗旱性评价

张伟溪¹(), 丁密¹, 苏晓华¹, 李爱平², 王小江², 余金金¹, 李政宏¹, 黄秦军¹, 丁昌俊¹^,^*()

1.林木遗传育种全国重点实验室,国家林业和草原局林木培育重点实验室,中国林业科学研究院林业研究所,北京 100091
2.内蒙古自治区林业科学研究院,内蒙古呼和浩特 010010

收稿日期:2023-10-31 修回日期:2023-12-15 出版日期:2025-01-30 发布日期:2025-01-21
通讯作者: * 丁昌俊(changjunding@caf.ac.cn),研究员。
作者简介:
张伟溪(weixizhang@caf.ac.cn),副研究员。
基金资助:
科技创新2030-重大项目(2023ZD0405603);国家自然科学基金面上项目(31870662);中国林业科学研究院基本科研业务费专项资金项目(CAFYBB2020SZ002)

Heterosis and drought resistance assessment of Populus simonii × P. nigra F₁ hybrids based on growth traits and leaf anatomical structures

ZHANG Weixi¹(), DING Mi¹, SU Xiaohua¹, LI Aiping², WANG Xiaojiang², YU Jinjin¹, LI Zhenghong¹, HUANG Qinjun¹, DING Changjun¹^,^*()

1. State Key Laboratory of Tree Genetics and Breeding, Key Laboratory of Tree Breeding Cultivation of National Forestry and Grassland Administration, Research Institute of Forestry, Chinese Academy of Forestry, Beijing 100091, China
2. Inner Mongolia Academy of Forestry Sciences, Hohhot 010010, China

Received:2023-10-31 Revised:2023-12-15 Online:2025-01-30 Published:2025-01-21

摘要/Abstract

摘要：

【目的】以内蒙古通辽天然小叶杨(Populus simonii)与荷兰北部欧洲黑杨(P. nigra)杂交F₁代的30个无性系为试验材料,对其自然半干旱条件下的F₁代生长性状及叶片解剖结构性状进行综合评价研究,为选育抗旱适应性强的杨树品种及亲本资源利用提供依据。【方法】对F₁代30个无性系的不同林龄(4、5、6 a)生长性状及9个叶片解剖结构性状(6年生)进行差异比较、遗传变异分析和杂种优势研究,采用相关性分析评判各指标的关联程度,采用主成分分析筛选典型叶片解剖结构性状,最后采用隶属函数法对30个杨树无性系6年生典型叶片解剖结构与生长性状进行抗旱性综合评价。【结果】不同无性系间9个叶片解剖结构及生长性状均存在极显著差异;树高和胸径生长性状变异系数变化范围分别为17.28%~19.24%和28.22%~29.87%,不同树龄树高和胸径均明显高于双亲,高亲优势率为29.27%~36.83%,部分杂交无性系的树高和胸径生长性状已经形成明显的超亲优势;各叶片解剖结构性状变异系数范围为5.78%~18.82%,重复力变化范围为0.91~0.97;F₁代的下表皮厚度、栅海比和叶片组织紧密度、栅栏组织厚度的正向超中亲优势明显,且角质层厚度、下表皮厚度、叶片组织紧密度出现明显正向超高亲优势,超高亲优势率为1.34%~1.77%。相关性分析结果表明,各指标之间具有较高的相关性,最终筛选出角质层厚度、叶片组织疏松度、叶片组织紧密度、栅海比和海绵组织厚度5个指标为小叶杨×欧洲黑杨杂交子代抗旱性评价的叶片解剖结构指标。通过隶属函数分析,最终筛选出02-06、02-01、02-05、02-24、02-03、02-13等6个最具有生长潜力和抗旱能力的无性系。【结论】小叶杨×欧洲黑杨杂交F₁代的生长和叶片解剖结构性状变异丰富,具有较大的选择潜力和杂种优势,初步筛选出6个最具有生长和抗旱潜力的无性系,为干旱地区选育高产高抗杨树新品种及育种亲本选配提供重要依据。

关键词: 小叶杨×欧洲黑杨, 生长, 叶片解剖结构, 遗传变异, 杂种优势

Abstract:

【Objective】30 F₁ hybrids of Populus simonii (from Tongliao, Inner Mongolia) × P. nigra (from northern Netherland) were used as experimental materials in this study. Aimed to comprehensive evaluate the growth and leaf anatomical structure traits of the F₁ hybrids under natural semi-arid conditions, which will provide a basis for selecting poplar varieties with strong drought tolerance adaptability and utilizing parental resources. 【Method】 Differences, genetic variation analysis, and heterosis analysis were conducted on the growth traits of F₁ hybrids at four, five, six years and nine leaf anatomical structure traits at six years. Correlation analysis was used to evaluate the correlation of each parameter. The principal component analysis was used to screen typical leaf anatomical structure traits. The membership function method was used to comprehensively evaluate the drought tolerance. 【Result】Growth traits and leaf anatomical structures were significant different among F₁ hybrids. The coefficient of variative (CV) of tree height (H) and diameter at breast height (DBH) is 17.28%-19.24% and 28.22%-29.87%, respectively. The H and DBH of F₁ hybrids at different ages are significantly higher than those of their parents, with a high-parent heterosis rate (R_Hb) 29.27%-36.83%. Partial hybrid clones have exhibited high-parent heterosis (H_b) at H and DBH. The CV of leaf anatomical structure traits is 5.78%-18.82%, with repeatability of which is 0.91-0.97. F₁ hybrids showed a significant positive mid-parent heterosis in lower epidermis thickness, thickness ratio of palisade to spongy, compaction of leaf tissue, thickness of palisade tissue, and a significant positive H_b in cuticle thickness, lower epidermis thickness and compaction of leaf tissue, with R_Hb of 1.34%-1.77%. There is a high correlation between various indicators, and compaction of leaf tissue, cuticle thickness, compaction of leaf tissue, palisade/spongy and thickness of spongy tissue were ultimately selected as the main component indicators of leaf anatomical structures for evaluating drought resistance of F₁ hybrids. Six clones (02-06, 02-01, 02-05, 02-24, 02-03, 02-13) with potential of growth and drought tolerance were ultimately selected. 【Conclusion】 Growth traits and leaf anatomical structure varied greatly in F₁ hybrids of P. simonii× P. nigra, which have great potential of selection and heterosis. Anatomical structure of leaves could be used to evaluate drought resistance of F₁ hybrids. Six clones (02-06, 02-01, 02-05, 02-24, 02-03, 02-13) with potential of fast growth and drought toterlance were initially selected. These results can provide candidate clones and important information of selecting breeding parents for breeding new high-yield and high resistance poplar varieties in arid areas.

Key words: Populus simonii ×, P. nigra, growth, leaf anatomical structures, genetic variation, heterosis

中图分类号:

S722.3

张伟溪,丁密,苏晓华,等. 小叶杨×欧洲黑杨杂交F₁代生长及叶片解剖结构杂种优势分析与抗旱性评价[J]. 南京林业大学学报（自然科学版）, 2025, 49(1): 46-58.

ZHANG Weixi, DING Mi, SU Xiaohua, LI Aiping, WANG Xiaojiang, YU Jinjin, LI Zhenghong, HUANG Qinjun, DING Changjun. Heterosis and drought resistance assessment of Populus simonii × P. nigra F₁ hybrids based on growth traits and leaf anatomical structures[J].Journal of Nanjing Forestry University (Natural Science Edition), 2025, 49(1): 46-58.DOI: 10.12302/j.issn.1000-2006.202310036.

图/表 12

表1

F1代无性系4~6年生长性状均值和标准差"

无性系 clone	4年生 the 4th growth year		5年生 the 5th growth year		6年生 the 6th growth year
无性系 clone	H/m	D_BH/cm	H/m	D_BH/cm	H/m	D_BH/cm
02-01	3.94±0.24 cdefghi	3.55±0.15 ab	5.13±0.13 defg	4.83±0.20 abcdef	6.78±0.20 cde	6.53±0.33 bcd
02-02	2.85±0.14 k	2.23±0.67 e	3.83±0.44 h	2.75±0.62 j	4.33±0.41 kl	3.23±0.38 hi
02-03	4.27±0.27 abcde	3.87±0.72 a	5.67±0.55 abcde	5.58±1.16 abc	7.17±0.53 abcd	7.70±1.26 abc
02-04	3.02±0.29 jk	1.93±0.43 e	3.73±0.15 h	2.49±0.46 j	4.03±0.16 kl	2.87±0.42 i
02-05	4.57±0.24 abcd	3.95±0.30 a	6.09±0.14 abc	5.88±0.41 ab	7.34±0.05 abc	8.19±0.50 a
02-06	4.45±0.29 abcde	4.05±0.47 a	5.96±0.05 abcd	5.66±0.55 abc	7.86±0.23 ab	7.71±1.01 abc
02-08	1.92±0.19 e	0.88±0.22 f	2.26±0.14 i	1.11±0.11 k	2.36±0.22 m	1.33±0.21 j
02-09	3.73±0.24 efghij	3.37±0.21 abc	5.23±0.16 def	4.53±0.42 bcdefg	6.66±0.35 cdef	5.84±0.44 def
02-10	4.12±0.32 abcdefgh	3.58±0.37 ab	5.32±0.35 cdef	4.46±0.49 cdefgh	6.50±0.41 cdef	5.47±0.52 defg
02-11	4.64±0.58 abc	4.16±1.05 a	6.20±0.82 ab	5.73±1.34 abc	7.87±0.85 ab	8.02±1.91 ab
02-12	4.60±0.16 abcd	3.67±0.22 ab	5.94±0.11 abcd	4.80±0.68 abcdef	7.34±0.14 abc	6.32±0.87 cd
02-13	4.22±0.58 abcdef	3.22±0.93 abcd	5.55±0.40 bcde	4.91±1.21 abcde	7.21±0.58 abcd	6.78±1.32 abcd
02-14	3.20±0.66 ijk	2.37±0.95 de	4.41±0.75 gh	3.46±1.23 ghij	5.39±0.54 hij	4.62±1.01 fgh
02-15	4.80±0.39 a	4.03±0.35 a	6.38±0.56 a	5.92±0.72 a	7.97±0.35 a	7.73±0.86 abc
02-16	4.48±0.72 abcde	3.86±0.67 a	5.93±0.71 abcd	5.80±1.29 abc	7.72±0.59 ab	7.71±1.12 abc
02-18	4.15±0.36 abcdefgh	3.88±0.36 a	5.58±0.24 abcde	5.32±0.54 abcd	7.38±0.21 abc	7.56±0.79 abc
02-19	3.43±0.78 hijk	2.22±0.94 e	4.64±0.95 fg	3.22±1.26 hij	5.52±1.33 ghij	4.29±1.50 ghi
02-20	4.70±0.30 ab	3.34±0.24 abcd	5.91±0.48 abcd	4.62±0.55 abcdefg	7.18±0.58 abcd	5.90±0.89 def
02-21	4.17±0.20 abcdefgh	3.53±0.05 ab	5.63±0.31 abcde	4.96±0.29 abcde	6.59±0.33 cdef	6.23±0.54 cde
02-22	4.61±0.39 abcd	3.68±0.58 a	5.64±0.37 abcde	4.96±0.63 abcde	6.83±0.36 cde	6.27±0.56 cde
02-23	4.48±0.27 abcde	3.60±0.22 ab	5.82±0.37 abcd	5.04±0.23 abcde	7.05±0.24 bcd	6.24±0.17 cde
02-24	4.20±0.14 abcdefg	3.52±0.27 ab	5.66±0.24 abcde	4.92±0.42 abcde	6.72±0.30 cdef	5.94±0.68 def
02-25	4.73±0.28 ab	3.77±0.10 a	5.86±0.37 abcd	5.32±0.25 abcd	7.21±0.33 abcd	6.75±0.55 abcd
02-26	4.00±0.36 bcdefgh	3.25±0.58 abcd	5.21±0.19 def	4.55±0.66 bcdefg	6.35±0.35 defg	5.64±0.71 defg
02-27	3.78±0.18 efghi	3.33±0.20 abcd	5.14±0.14 defg	4.92±0.17 abcde	6.36±0.19 defg	6.30±0.34 cde
02-28	3.25±0.33 ijk	2.37±0.19 cde	4.62±0.27 fg	3.57±0.17 fghij	5.04±0.33 ijk	3.86±0.52 hi
02-29	3.47±0.58 ghijk	2.25±0.79 e	5.14±0.58 defg	3.76±0.73 efghij	5.96±0.62 efgh	4.72±0.92 efgh
02-30	3.50±0.16 fghijk	3.28±0.33 abcd	4.90±0.08 efg	4.68±0.34 abcdefg	5.85±0.26 fghi	5.74±0.60 defg
02-31	3.86±0.42 defghi	2.68±0.60 bcde	5.19±0.32 defg	4.15±0.70 defghi	6.12±0.53 efgh	5.47±0.93 defg
02-32	3.45±0.18 hijk	2.22±0.31 e	4.58±0.20 fg	2.98±0.23 ij	4.98±0.08 jk	3.67±0.33 hi
均值mean	3.94±0.74	3.17±0.90	5.22±0.90	4.48±1.26	6.38±1.25	5.78±1.73

表1

表2

表3

表4

表5

F1代无性系叶片解剖结构性状均值及多重比较"

无性系 clone	叶片厚度/μm leaf thickness	角质层厚度/μm cuticle thickness	上表皮厚度/μm upper epidermis thickness	下表皮厚度/ μm lower epidermis thickness	栅栏组织厚度/μm thickness of palisade tissue	海绵组织厚度/μm thickness of spongy tissue	栅海比 palisade to spongy ratio	组织紧密度/% compaction of leaf tissue	组织疏松度/% porosity of leaf tissue
02-01	266.45±10.34 efghi	6.46±1.04 a	11.90±0.90 a	12.44±0.97 a	116.69±5.89 bcd	117.14±6.86 ij	1.00±0.04 abc	43.81±1.78 ab	43.94±1.24 fghijkl
02-02	271.96±18.98 defg	3.99±0.69 no	11.89±0.98 a	9.42±1.01 cd	115.21±7.89 bcdef	125.69±10.57 defgh	0.92±0.05 efghi	42.39±1.43 bcdef	46.19±1.52 bcd
02-03	264.38±7.61 ghi	5.28±0.98 cde	12.24±1.00 a	10.58±1.56 b	109.54±7.18 ghijk	116.84±9.42 ij	0.94±0.07 cdefgh	41.41±2.09 fghi	44.15±2.73 fghijk
02-04	260.13±10.63 hij	4.07±0.39 mno	10.10±0.88 cdefghi	9.92±0.99 bc	99.94±4.59 no	127.25±8.59 cde	0.79±0.05 m	38.44±1.42 m	48.89±1.92 a
02-05	268.22±13.87 efgh	6.65±0.67 a	10.04±0.95 cdefghi	10.47±1.30 b	112.29±5.87 defghi	120.26±9.78 fghi	0.94±0.06 defgh	41.89±1.48 defgh	44.79±1.81 defghi
02-06	269.28±13.04 defgh	5.70±0.59 bc	10.55±1.43 cde	10.40±1.11 b	118.39±5.61 bc	117.99±6.54 ij	1.01±0.06 ab	44.00±1.82 a	43.82±1.36 ghijkl
02-08	232.29±5.14 mn	3.71±0.36 o	8.54±1.13 lm	7.77±1.03 ijklm	92.86±6.01 p	108.94±6.71 k	0.86±0.10 jkl	39.97±2.32 ijkl	46.92±2.99 bc
02-09	266.33±18.30 efghi	4.27±0.56 klmn	10.35±1.40 cdef	9.34±1.08 cd	114.73±8.11 bcdefg	115.92±9.57 ij	0.99±0.07 abcd	43.10±1.68 abcd	43.52±1.83 hijkl
02-10	266.68±4.16 efghi	4.02±0.54 no	10.66±1.04 cd	8.94±1.07 defg	115.39±2.71 bcdef	119.64±4.00 fghi	0.97±0.03 bcde	43.28±1.08 abcd	44.86±1.06 defghi
02-11	265.07±6.57 fghi	4.88±0.73 efghi	8.82±0.99 klm	7.57±0.95 jklm	106.47±7.92 jklm	119.92±6.03 fghi	0.89±0.09 hij	40.14±2.40 ijkl	45.25±2.30 defg
02-12	256.94±9.80 ij	4.90±0.70 defghi	8.38±1.23 m	7.19±0.84 m	100.65±8.12 no	121.65±8.37 efghi	0.83±0.10 klm	39.16±2.61 klm	47.35±2.71 b
02-13	237.26±4.73 m	4.72±0.75 ghijk	8.56±1.06 lm	7.43±0.55 lm	101.93±3.73 mno	101.40±3.14 l	1.01±0.05 ab	42.97±1.66 abcde	42.73±0.71 kl
02-14	270.45±5.47 defg	4.22±0.53 lmn	9.20±0.96 ijklm	7.73±1.11 ijklm	107.44±6.86 ijkl	132.09±8.04 bc	0.82±0.08 lm	39.72±2.25 jklm	48.84±2.84 a
02-15	264.38±13.63 ghi	5.88±0.56 b	9.55±1.25 fghijk	8.00±1.19 hijklm	108.68±5.78 hijkl	117.43±11.02 ij	0.93±0.07 efgh	41.13±1.61 fghij	44.34±2.28 efghijk
02-16	239.38±16.41 lm	4.77±0.55 fghij	9.49±0.91 fghijk	7.34±0.64 m	93.89±9.96 p	107.07±10.69 k	0.88±0.11 hijk	39.19±2.63 klm	44.69±2.78 defghij
02-18	278.77±12.80 cd	5.70±0.83 bc	10.17±1.55 cdefgh	8.38±0.79 fghij	115.32±6.96 bcdef	125.84±8.85 defg	0.92±0.07 efghi	41.37±1.56 fghi	45.13±2.00 defgh
02-19	287.52±8.01 b	4.64±0.34 ghijkl	9.36±1.35 hijkl	8.37±1.11 fghij	115.82±7.28 bcde	134.72±5.26 b	0.86±0.08 ijkl	40.27±2.01 ijkl	46.89±2.30 bc
02-20	266.05±18.62 efghi	5.29±0.38 cde	10.33±1.31 cdef	8.66±1.27 defgh	110.13±5.73 fghij	121.61±17.00 efghi	0.92±0.12 efghi	41.52±2.72 efghi	45.52±3.32 cdef
02-21	272.90±16.09 defg	4.51±0.39 hijklm	10.79±1.10 bc	9.20±1.07 cde	115.68±12.10 bcde	120.43±6.94 fghi	0.96±0.10 bcdef	42.31±2.51 bcdefg	44.19±2.21 fghijk
02-22	246.67±12.90 kl	5.07±0.74 defg	9.85±1.36 defghij	7.44±0.68 lm	103.64±7.06 lmno	108.03±8.22 k	0.96±0.09 bcde	42.06±2.64 defgh	43.78±2.06 ghijkl
02-23	275.14±6.04 de	4.40±0.50 jklmn	11.52±1.40 ab	9.14±1.46 def	118.52±3.69 bc	119.56±4.72 ghi	0.99±0.06 abcd	43.09±1.37 abcd	43.46±1.47 ijkl
02-24	265.43±5.50 efghi	4.96±0.47 defgh	10.64±0.87 cd	8.50±1.30 efghi	115.81±8.09 bcde	112.50±4.65 jk	1.03±0.11 a	43.62±2.79 abc	42.40±1.98 l
02-25	264.75±13.39 ghi	5.90±0.68 b	9.69±1.23 efghijk	7.53±1.08 klm	111.18±3.95 efghij	116.43±8.97 ij	0.96±0.07 bcdefg	42.05±1.54 defgh	43.94±1.79 fghijkl
02-26	284.88±11.78 bc	4.86±0.45 efghi	11.53±1.14 ab	8.31±1.12 ghijk	116.66±8.64 bcd	129.59±7.36 bcd	0.90±0.08 fghij	40.93±2.13 fghij	45.50±1.97 cdef
02-27	271.89±12.04 defg	5.17±0.47 def	10.30±0.84 cdefg	8.17±0.79 ghijkl	110.31±3.07 fghij	125.03±7.85 defgh	0.89±0.06 hijk	40.62±1.43 hijk	45.97±1.70 bcde
02-28	251.15±10.44 jk	4.51±0.28 ijklm	10.68±1.30 cd	8.46±1.15 efghi	104.35±7.64 klmn	108.56±6.25 k	0.96±0.09 bcde	41.52±1.83 efghi	43.26±2.53 ijkl
02-29	266.10±14.94 efghi	4.11±0.27 mno	9.48±0.91 fghijk	7.95±0.89 hijklm	108.40±6.38 hijkl	120.47±12.42 fghi	0.91±0.10 efghij	40.79±2.21 ghij	45.20±2.89 defg
02-30	274.08±17.73 defg	4.51±0.56 ijklm	9.41±1.12 ghijk	8.19±0.93 ghijkl	113.27±6.69 cdefgh	126.27±11.28 cdef	0.90±0.06 ghij	41.36±1.22 fghi	46.03±1.94 bcd
02-31	266.09±13.20 efghi	4.68±0.46 ghijk	9.69±1.00 efghijk	8.25±0.82 ghijkl	107.99±5.45 hijkl	121.12±7.65 efghi	0.89±0.05 hij	40.61±1.39 hijk	45.50±1.28 cdef
02-32	274.81±11.70 def	4.61±0.54 hijkl	9.92±0.87 cdefghij	8.22±1.11 ghijkl	119.02±5.58 b	119.08±8.39 hi	1.00±0.07 ab	43.32±1.31 abcd	43.31±1.93 ijkl

表5

图1

表6

表7

表8

表9

表10

表11

30个杨树无性系抗旱性的综合评价结果"

无性系 clone	隶属函数值 membership function value							均值 mean	抗旱能力排序 sort drought tolerance
无性系 clone	年均高生长量 average annual growth of heigh	年均胸径生长量 average annual growth of DBH	角质层厚度 cuticle thickness	叶片组织疏松度 porosity of leaf tissue	海绵组织厚度 thickness of spongy tissue	栅海比 palisade to spongy ratio	叶片组织紧密度 compaction of leaf tissue	均值 mean	抗旱能力排序 sort drought tolerance
02-06	1.060 71	0.824 24	0.677 37	0.780 91	0.502 16	0.888 40	1.000 00	0.819 11	1
02-01	0.857 14	0.618 18	0.935 23	0.762 39	0.527 51	0.854 64	0.964 65	0.788 53	2
02-05	0.832 14	1.000 00	1.000 00	0.631 65	0.433 94	0.609 65	0.620 06	0.732 49	3
02-24	0.635 71	0.500 00	0.464 77	0.787 90	0.667 03	1.000 00	0.931 94	0.712 48	4
02-03	0.878 57	0.875 76	0.535 67	0.729 98	0.536 64	0.624 55	0.534 87	0.673 72	5
02-13	0.821 43	0.518 18	0.406 15	0.237 50	1.000 00	0.890 88	0.815 29	0.669 92	6
02-16	0.975 00	0.836 36	0.740 11	0.700 64	0.830 02	0.387 32	0.135 29	0.657 82	7
02-20	0.996 43	0.830 30	0.676 14	0.579 88	0.393 38	0.538 12	0.553 97	0.652 60	8
02-22	0.728 57	0.490 91	0.536 97	0.518 90	0.801 22	0.720 43	0.650 75	0.635 39	9
02-10	0.889 29	0.463 64	0.192 12	0.828 21	0.452 48	0.724 07	0.869 86	0.631 38	10
02-23	0.707 14	0.533 33	0.273 14	0.724 51	0.454 94	0.838 42	0.835 92	0.623 91	11
02-25	0.760 71	0.515 15	0.233 49	0.837 25	0.549 04	0.699 15	0.648 87	0.606 24	12
02-18	1.000 00	0.881 82	0.361 30	0.646 47	0.266 32	0.536 55	0.526 28	0.602 68	13
02-28	0.682 14	0.439 39	0.393 15	0.522 62	0.785 31	0.720 31	0.553 60	0.585 22	14
02-32	0.682 14	0.460 61	0.270 85	0.440 94	0.469 33	0.877 67	0.877 44	0.582 71	15
02-27	0.728 57	0.618 18	0.746 92	0.762 85	0.290 66	0.397 40	0.391 91	0.562 36	16
02-26	0.742 86	0.448 48	0.427 31	1.000 00	0.153 84	0.469 50	0.448 24	0.527 18	17
02-29	0.764 29	0.615 15	0.497 78	0.449 25	0.427 69	0.491 32	0.422 26	0.523 96	18
02-12	0.996 43	0.884 85	0.399 80	0.559 91	0.392 21	0.181 84	0.130 27	0.506 47	19
02-14	0.910 71	0.793 94	0.343 19	0.948 63	0.078 77	0.119 82	0.230 27	0.489 33	20
02-08	0.650 00	0.560 61	0.331 72	0.521 83	0.773 76	0.285 09	0.275 77	0.485 54	21
02-21	0.589 29	0.342 42	0.314 95	0.307 68	0.428 77	0.709 70	0.696 09	0.484 13	22
02-31	0.732 14	0.463 64	0.135 27	0.567 99	0.408 24	0.431 16	0.389 85	0.446 90	23
02-30	0.482 14	0.166 67	0.271 93	0.867 27	0.253 46	0.460 52	0.525 16	0.432 45	24
02-11	0.692 86	0.287 88	0.105 85	0.621 08	0.444 24	0.421 15	0.306 02	0.411 30	25
02-15	0.625 00	0.396 97	0.173 53	0.006 49	0.518 84	0.582 93	0.484 87	0.398 38	26
02-02	0.371 43	0.018 18	0.094 59	0.415 98	0.270 99	0.536 16	0.709 68	0.345 29	27
02-09	0.000 00	-0.148 48	0.000 00	0.303 63	0.564 27	0.838 08	0.838 21	0.342 24	28
02-19	0.389 29	0.154 55	0.307 59	0.860 36	0.000 00	0.303 46	0.329 00	0.334 89	29
02-04	0.203 57	0.000 00	0.124 02	0.000 00	0.224 06	0.000 00	0.000 00	0.078 81	30

表11

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树龄 tree age	性状 trait	平方和 sum of square	自由度 degrees of freedom	均方 mean square	F	P
4年生 the 4th growth year	H	66.11	29	2.28	14.92	<0.001
	D_BH	91.32	29	3.15	11.28	<0.001
5年生 the 5th growth year	H	105.98	29	3.65	20.33	<0.001
	D_BH	187.81	29	6.48	13.30	<0.001
6年生 the 6th growth year	H	219.77	29	7.58	34.35	<0.001
	D_BH	382.07	29	13.17	19.26	<0.001

树龄 tree age	性状 trait	母本 female	父本 male	中亲值 MPV	中亲优势 H_m	高亲值 BPV	超亲优势 H_b	中亲优势率/% R_Hm	超亲优势率/% R_Hb
4年生 the 4th growth year	H/m	2.94	2.67	2.81	1.15^**	2.94	1.01^**	40.94	34.33
	D_BH/cm	2.33	1.08	1.71	1.49^**	2.33	0.86^**	87.07	36.83
5年生 the 5th growth year	H/m	3.98	3.88	3.93	1.31^**	3.98	1.26^**	33.30	31.71
	D_BH/cm	3.45	1.85	2.65	1.85^**	3.45	1.05^**	69.64	30.46
6年生 the 6th growth year	H/m	4.94	3.74	4.34	2.05^**	4.94	1.45^**	47.13	29.27
	D_BH/cm	4.30	1.89	3.10	2.72^**	4.30	1.52^**	87.90	35.25

性状 trait	自由度 df	均方 MS	F	P
叶片厚度 leaf thickness	29	2 925.602	19.447	<0.001
角质层厚度 cuticle thickness	29	9.466	26.750	<0.001
上表皮厚度 upper epidermis thickness	29	18.342	14.288	<0.001
下表皮厚度 lower epidermis thickness	29	25.260	22.610	<0.001
栅栏组织厚度 thickness of palisade tissue	29	896.639	19.479	<0.001
海绵组织厚度 thickness of spongy tissue	29	1 027.543	13.826	<0.001
栅海比 palisade to spongy ratio	29	666.859	11.129	<0.001
叶片组织紧密度 compaction of leaf tissue	29	39.530	10.272	<0.001
叶片组织疏松度 porosity of leaf tissue	29	47.405	10.376	<0.001

性状 trait	叶片厚度/μm leaf thickness	角质层厚度/μm cuticle thickness	上表皮厚度/μm upper epidermis thickness	下表皮厚度/μm lower epidermis thickness	栅栏组织厚度/μm thickness of palisade tissue	海绵组织厚度/μm thickness of spongy tissue	栅海比 palisade to spongy ratio	叶片组织紧密度/% compaction of leaf tissue	叶片组织疏松度/% porosity of leaf tissue
平均值average	264.85±17.31	4.88±0.92	10.12±1.48	8.64±1.55	110.01±9.58	119.28±11.21	92.78±9.62	41.53±2.40	45.01±2.62
变幅variable amplitude	217.36~305.74	2.95~8.39	6.50~14.20	5.91~13.66	79.14~138.21	89.23~150.02	65.30~121.37	34.18~48.01	38.02~54.73
变异系数/% CV	6.54	18.82	14.66	17.99	8.71	9.40	10.37	5.78	5.82
重复力repeatability	0.95	0.96	0.93	0.96	0.95	0.93	0.91	0.90	0.90

项目 item	叶片厚度/μm leaf thickness	角质层厚度/μm cuticle thickness	上表皮厚度/μm upper epidermis thickness	下表皮厚度/μm lower epidermis thickness	栅栏组织厚度/μm thickness of palisade tissue	海绵组织厚度/μm thickness of spongy tissue	栅海比/% palisade to spongy ratio	叶片组织紧密度/% compaction of leaf tissue	叶片组织疏松度/% porosity of leaf tissue
母本female	309.95±18.90	4.80±0.49	9.97±1.33	8.49±0.64	126.43±11.38	145.72±7.65	87.76±6.31	40.76±2.10	47.07±1.88
父本male	225.07±11.91	3.96±0.43	10.64±1.19	8.47±1.09	92.14±4.18	97.12±8.13	95.33±7.12	40.98±1.62	43.10±1.79
中亲值MPV	267.51	4.38	10.31	8.48	109.28	121.42	91.04	40.87	45.08
中亲优势H_m	-3.01	0.52^**	-0.19	0.15	0.37	-2.14^**	1.74	0.58^*	-0.01
高亲值BPV	309.95	4.80	10.64	8.49	126.43	145.72	95.33	40.98	47.07
超高亲优势H_b	-45.10^**	0.08	-0.52^**	0.15	-16.42^**	-26.44^**	-2.55^*	0.55	-2.06^**
中亲优势率/% R_Hm	-1.12	11.86	-1.80	1.75	0.33	-1.76	1.91	1.42	-0.03
超高亲优势率/% R_Hb	-14.55	1.67	-4.89	1.77	-12.99	-18.14	-2.67	1.34	-4.38