211份板栗种质资源花序表型多样性和聚类分析

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李颖, 张树航, 郭燕, 张馨方, 王广鹏^,河北省农林科学院昌黎果树研究所,河北昌黎 066600

Catkin Phenotypic Diversity and Cluster Analysis of 211 Chinese Chestnut Germplasms

LI Ying, ZHANG ShuHang, GUO Yan, ZHANG XinFang, WANG GuangPeng^,Changli Institute of Pomology,Hebei Academy of Agricultural and Forestry Sciences,Changli 066600, Hebei

通讯作者: 王广鹏,Tel：0335-2022724;E-mail： wangguangpeng430@163.com

责任编辑: 赵伶俐
收稿日期:2020-04-2接受日期:2020-07-15网络出版日期:2020-11-16

基金资助:

河北省农林科学院创新工程项目.2019-3-4-2
河北省科技计划.16226312D
河北省农林科学院创新团队项目.F20E06001

Received:2020-04-2Accepted:2020-07-15Online:2020-11-16
作者简介 About authors
李颖,E-mail： beierdina3320@sina.com。

摘要
【目的】通过对10个省份（群体）211份中国板栗种质资源花序相关性状表型多样性和遗传特点的研究,进一步丰富中国板栗表型性状遗传多样性信息,挖掘优异基因材料,并为现有种质资源保护、利用、创新及遗传改良提供参考。【方法】采用“板栗种质资源描述规范和数据标准”中提供的方法,对河北省农林科学院昌黎果树研究所板栗种质资源圃内211份中国板栗资源的15个花序相关表型性状进行系统调查,量化赋值后,使用SPSS 20.0软件进行遗传变异分析、Shannon-weaver指数多样性分析、相关性分析和主成分分析,采用MEGA 7.0进行聚类分析。【结果】我国板栗资源花序相关性状具有丰富的表型多样性,平均变异系数和平均多样性指数分别为28.23%和1.70,其中每果枝雌花个数的变异系数最大为60.66,雄花序比例变异系数最小为7.37,序轴粗度比的多样性指数最高为1.99,每果枝两性花序个数、两性花序比例多样性指数最低为1.41。进一步分析发现不同地区板栗遗传变异和表型多样性存在差异,江苏群体变异程度最大,变异系数为35.53%;河北群体多样性水平最高,多样性指数1.86;安徽群体遗传变异和表型多样性水平均最低,变异系数和多样性指数分别为16.96%和0.95。测量性状中,除雄蕊长和序轴粗度比外,其余性状均为群体间差异显著,花序长、花轴粗、花簇密度、每果枝两性花序个数、两性花序比例、雄花序比例、每果枝雌花个数这7个性状,群体间差异极显著,不同地域间变异丰富,多样性程度高。相关性分析表明,花序形态性状内部间的相关性较明显,花序数量性状内部间的相关性较明显,但花序形态和花序数量间相关性不明显。主成分分析表明：前5个主成分反映了总信息量的84.18%,每果枝两性花序个数（0.931）、花序粗（0.912）、花序长粗比（-0.889）、花序长（0.864）、每果枝雌花个数（0.828）、雄花序比例（-0.821）、两性花序比例（0.820）、雄蕊长（0.806）8个性状的相关系数都在0.8以上,是板栗花序相关性状变异的主要因素。以第一主成分和第二主成分为标准,将211份资源分为5个类群,并筛选出9份两性花序数量多、雌雄比例高且高雌花量的板栗资源。聚类分析将211份资源分为8个类群,花相关表型性状变异相同的材料大多聚在一起,变异较大的类群和主成分分析结果相似。【结论】中国板栗资源花序相关表型性状变异丰富,多样性程度高,地域间遗传变异和多样性程度不同,群体间性状差异显著。花序形态表型性状和花序数量表型性状内部的相关性较明显,但花序形态和花序数量间相关性不明显。筛选出8个性状可作为板栗花形和花量的综合评定指标,9个两性花序数量多、雌雄比例高且高雌花量的板栗资源可作为性别调控、改善产量的亲本材料。
关键词： 板栗;种质资源;花序;表型性状;多样性;聚类分析

Abstract

【Objective】The study on the diversity and genetic rules of catkin phenotypic traits of 211 Chinese chestnut germplasms from 10 provinces was conducted, in order to provide valuable basic data and theory foundation for the protection, utilization, innovation and genetic improvement of existing germplasm resources. 【Method】According to the method described in Descriptors and Data Standard for chestnut(Castanea mollissima Bl.), the data were collected for 15 phenotypic parameters of catkin phenotypic traits from 211 Chinese chestnut germplasms from 10 provinces in Germplasm Repository of chestnut of Changli Institute of Pomology, Hebei Academy of Agricultural and Forestry Sciences. Then, the genetic variation analysis, Shannon-index, correlation andprincipal component analysis of chestnut flower phenotypic traits were carried out by using the SPSS20.0 software. The Chinese chestnut were clustered using MEGA 7.0 according to morphological data.【Result】phenotypic traits related catkin of Chinese chestnut were rich in phenotypic diversity, with mean variation coefficient (CV) of 28.23% and Shannon's diversity index(H') of 1.70. The mean variation coefficient (CV = 60.66) of female flower numbers/bearing shoot was the largest, the mean variation coefficient (CV = 7.37）of male catkin numbers/ catkin numbers was the smallest. The Shannon's diversity index (H'=1.99）of floral axis width/catkin width was the highest, The Shannon's diversity index (H'=1.41) of bisexual catkin numbers/bearing shoot and bisexual catkin numbers/catkin numbers were the lowest. Further analysis showed that the phenotypic diversity of Chinese chestnut existed differences among 10 populations, which was the highest in Hebei province. Catkin length, floral axis width, catkin density, female flower numbers/bearing shoot, bisexual catkin numbers/catkin numbers, male catkin numbers/catkin numbers, female flower numbers/bearing shoot were highly significant, which indicated that a wide range of variation existed in these two levels. The correlation analysis showed that: catkin morphology traits were highly significantly or extreme significantly related, catkin number traits were highly significantly or extreme significantly related, and the correlation between catkin morphology traits and catkin number traits were not highly significantly related. Principal component analysis showed that catkin traits in the first 5 principal components reflect the total information content 84.18%, catkin length, bisexual catkin numbers/bearing shoot, catkin width, catkin length/catkin width, female flower numbers/ bearing shoot, male catkin numbers/catkin numbers, bisexual catkin numbers/catkin numbers, stamen length height characters were the main factors causing variation in traits of Chinese chestnut. The 211 materials were divided into 8 kinds of cluster analysis. Catkin character same source with similar material part could get together, and the results were similar to principal component analysis.【Conclusion】There were abundant variation and high genetic diversity degree based on the phenotype of catkin phenotypic traits of Chinese chestnut. The degree of genetic variation and diversity was different between regions, for there were significant differences in traits between populations. The correlations between catkin morphology traits and catkin number traits were not highly significantly related. 8 traits were selected to be as the important comprehensive evaluation indexes used for Chinese chestnut germplasm. 9 chestnut germplasm with more female flowers was selected as parent materials to control sex and improve the yield.

Keywords：chestnut;germplasms;phenotypic trait;catkin;diversity;cluster analysis

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本文引用格式
李颖, 张树航, 郭燕, 张馨方, 王广鹏. 211份板栗种质资源花序表型多样性和聚类分析[J]. 中国农业科学, 2020, 53(22): 4667-4682 doi:10.3864/j.issn.0578-1752.2020.22.013
LI Ying, ZHANG ShuHang, GUO Yan, ZHANG XinFang, WANG GuangPeng. Catkin Phenotypic Diversity and Cluster Analysis of 211 Chinese Chestnut Germplasms[J]. Scientia Agricultura Sinica, 2020, 53(22): 4667-4682 doi:10.3864/j.issn.0578-1752.2020.22.013

开放科学（资源服务）标识码（OSID）：

0 引言

【研究意义】板栗（Castanea mollissima Bl.）起源于中国,植株抗旱、耐瘠薄,果实营养价值高,是世界上山地开发所依赖的重要坚果类树种^[1]。板栗在中国已有4 000多年的栽培历史,现广泛分布在境内24个省、市、自治区,各地板栗在长期的自然演化和人工选择下,形成了各具特色、遗传丰富的种质资源,挖掘利用潜力巨大^[1,2]。明确种质遗传多样性是开展创新利用的基础,其中表型性状多样性是种质整体多样性的一个重要组成部分^[3,4,5],对种质的表型性状值进行科学分析,有助于了解其遗传稳定性和育种潜力,对发掘、利用和创新种质具有重要指导意义。【前人研究进展】目前关于板栗表型遗传多样性方面的研究,主要包括叶片、刺苞、果实、品质、枝条等性状^{[6,7,8,9,10,11,12,13]},叶片和果实等性状的表型分化系数显示,板栗的表型变异主要来源于群体内部,同一群体品种间的变异更高,多样性程度高于群体间,叶片大小、叶柄的长（粗）度、果实形状、果实大小等形态性状的变异系数均高于10%,遗传多样性丰富,选择潜力巨大,中国板栗果实表型性状遗传稳定性高于品质性状,叶片性状由南向北表现出从长椭圆形逐渐变为椭圆形的规律,北方板栗果实质量小于南方板栗,枝条涉及的9个性状中,表型分化幅度较大,其主要变异来源为群体间变异。同时在分子研究方面,利用AFLP、SSR、ISSR 和RAPD等分子标记技术对栗属植物的多样性水平和起源问题也进行了研究^[14]。取得的研究结果以及通过评价鉴定出的特异或优良种质,已在育种生产及科研利用中发挥了重要作用^[15]。【本研究切入点】花序作为板栗的重要器官,其形态大小,两性花序的比例等性状是遗传变异的重要特征,同时对产量和品质影响巨大^[3,16],但至今仍缺乏关于板栗花序相关性状遗传变异多样性的系统报道。同时以现有的分子生物学信息,在花序性状相关基因发掘不明确的情况下,很难有针对性地筛选花序性状优异的相关种质。【拟解决的关键问题】本研究以我国10个板栗主产区（群体）的211份实生良种、地方品种和古树资源为材料,鉴定评价其花序相关15个表型性状的遗传多样性程度,进一步丰富中国板栗表型性状遗传多样性信息,为现有板栗种质资源的挖掘、创新和保护提供科学支撑,提高育种效率,为今后核心种质的筛选提供依据,为花序表型相关性状的基因定位奠定前期基础。

1 材料与方法

1.1 试验材料

以源自我国10个板栗主产区（群体）的211份实生良种、地方品种和古树资源为试验材料（表1）。试材保存于河北省昌黎果树研究所板栗种质资源圃,东经119°15′,北纬39°72′,属温带半湿润大陆性气候区,年平均气温11℃,无霜期186 d,年平均降水量638 mm。所有供试板栗种质均是在2004年统一嫁接于3年生‘燕山早丰’实生砧木上,每份种质嫁接5株,株行距4 m×4 m,常年树体整形修剪均采用轮替更新修剪法^[17],每m²留枝量保持在6—9条,其他管理措施一致。2017—2019年,连续3年选取每份种质生长状况基本一致的健康树体3株,分别测量各单株花序相关表型性状。

1.2 花序相关表型测定

花序长度、花序粗度和花序小花簇密度按照刘庆忠^[18]方法测定。

花轴粗度：在雄花盛花期,随机选取树冠外围1年生结果新枝中部的雄花序20条,采用游标卡尺测量花序中间花轴的直径（mm）,精确到0.1mm,取平均值。

雄蕊长度=（花序粗度-轴粗度）/2。单位为mm,精确到0.1mm。

花序长粗比=（花序长度/花序粗度）×100%

花轴长粗比：花轴长度同花序长度。花轴长粗比=（花序长度/花轴粗度）×100%

轴序粗度比=（花轴粗度/花序粗度）×100%

每果枝雄花序个数：在雄花盛花期,随机选取树冠外围1年生结果新枝20条,调查其上雄花序数量,

取平均值。

每果枝两性花序个数：在雄花盛花期,随机选取树冠外围1年生结果新枝20条,调查其上两性花序数量,取平均值。

每果枝雌花个数：在雄花盛花期,随机选取树冠外围1年生结果新枝20条,调查其上雌花数量,取平均值。

每果枝总花序个数：在雄花盛花期,随机选取树冠外围1年生结果新枝20条,调查其上花序数量,取平均值。

每两性花序上雌花个数：在雄花盛花期,随机选取树冠外围1年生结果新枝20条,调查其上两性花序数量及两性花序上着生的雌花个数。每两性花序上雌花数=雌花总数/两性花序总数。

雄花序比例=（每果枝雄花序个数/每果枝总花序个数）×100%

两性花序比例=（每果枝两性花序个数/每果枝总花序个数）×100%。

1.3 数据分析

测量性状,计算均值(X)和标准差(δ),Shannon's多样性指数(H')的计算公式为：H'=-ΣPi ln Pi,参考文献[19,20,21,22,23]方法。试验数据采用Excel 2010统计并计算各性状的平均值、标准差和变异系数等,用IBM SPSS 20软件对表型数据进行相关性分析和主成分分析,使用EGMA 7.0软件对表型性状进行聚类分析。

Table 1
表1
表1211份板栗种质资源编号、来源及名称
Table 1Name, source and number of 211 resources in chestnut

序号 No.	来源 Source	资源名称 Name	序号 No.	来源 Source	资源名称 Name
1	河北Hebei	燕宝 Yanbao	40	河北Hebei	西寨2号 Xizhai 2
2	河北Hebei	燕秋 Yanqiu	41	河北Hebei	西寨1号 Xizhai 1
3	河北Hebei	燕丽 Yanli	42	河北Hebei	邢台薄皮 Xingtaibaopi
4	河北Hebei	燕紫 Yanzi	43	河北Hebei	邢台垂枝 Xingtaichuizhi
5	河北Hebei	遵化短刺 Zunhuadanci	44	河北Hebei	后南峪垂枝 Hounayuchuizhi
6	河北Hebei	遵玉 Zunyu	45	河北Hebei	替码燕明 Timayanming
7	河北Hebei	宽城榆树沟 Kuanchengyushugou	46	河北Hebei	变异燕明 Bianyiyanming
8	河北Hebei	大桑园1号 Dasangyuan 1	47	河北Hebei	抚宁薄皮 Funingbaopi
9	河北Hebei	东冰窖1号 Dongbingjiao 1	48	河北Hebei	前3 Qian 3
10	河北Hebei	金杖子1号 Jinzhangzi 1	49	河北Hebei	大板红 Dabanhong
11	河北Hebei	金杖子2号 Jinzhangzi 2	50	河北Hebei	84--2
12	河北Hebei	桥7 Qiao 7	51	河北Hebei	小官10 Xiaoguan 10
13	河北Hebei	早3113 Zao 3113	52	河北Hebei	大官10 Da guan 10
14	河北Hebei	屈家早 Qujiazao	53	河北Hebei	燕山短枝 Yanshandanzhi
15	河北Hebei	兴隆短刺 Xinglongduanci	54	河北Hebei	早丰 Zaofeng
16	河北Hebei	大3113 Da 3113	55	河北Hebei	北峪2号 Beiyu 2
17	河北Hebei	陈中熟 Chenzhongshu	56	河北Hebei	东陵明珠 DonglingMingzhu
18	河北Hebei	白露 Bailu	57	河北Hebei	10补栽 10 Buzhai
19	河北Hebei	大青裂 Daqinglie	58	河北Hebei	达1-3 DA 1-3
20	河北Hebei	xl-001	59	河北Hebei	侯庄2号 Houzhuang 2
21	河北Hebei	xl-002	60	河北Hebei	贾庄1号 Jiazhuang 1
22	河北Hebei	迁西壮栗 Qianxizhuangli	61	河北Hebei	干2-2 Gan2-2
23	河北Hebei	崔1 Cui 1	62	河北Hebei	岔3 Cha 3
24	河北Hebei	艾峪口 Aiyukou	63	河北Hebei	沙坡峪3号 Shapoyu 3
25	河北Hebei	熊84 Xiong 84	64	河北Hebei	燕栗2号 Yanli 2
26	河北Hebei	迁西早红 Qianxizaohong	65	河北Hebei	关堂64 Guantang 64
27	河北Hebei	去暑红 Qushuhong	66	河北Hebei	龙湾1号 Longwan 1
28	河北Hebei	青龙白 Qinglongbai	67	河北Hebei	龙湾5号 Longwan 5
29	河北Hebei	燕兴 Yanxing	68	河北Hebei	石场子2-2 Shichangzi2-2
30	河北Hebei	28-11	69	河北Hebei	杨家峪13 Yanjiayu 13
31	河北Hebei	84--3	70	河北Hebei	西沟7号 Xigou 7
32	河北Hebei	919	71	河北Hebei	刑台丰收1 Xingtaifengshou 1
33	河北Hebei	X12-55	72	河北Hebei	塔54 Ta 54
34	河北Hebei	石场子1-1 Shichangzi1-1	73	河北Hebei	上庄52 Shangzhuang 52
35	河北Hebei	燕奎 Yankiui	74	河北Hebei	下庄4号 Xiazhuang 4
36	河北Hebei	早1 Zao 1	75	河北Hebei	燕金 Yanjin
37	河北Hebei	早2 Zao 2	76	河北Hebei	杨家峪5号 Yangjiayu 5
38	河北Hebei	早3 Zao 3	77	河北Hebei	浡1 Bo 1
39	河北Hebei	邢台早熟 Xingtaizaoshu	78	河北Hebei	大录洞 Daludong
79	河北Hebei	凤2 Feng 2	119	河北Hebei	燕光 Yanguang
80	河北Hebei	东沟峪39 Dongguoyu 39	120	河北Hebei	燕晶 Yanjing
81	河北Hebei	赵杖子11 Zhaozhangzi 11	121	河北Hebei	粗城大屯 Kuanchengdatun
82	河北Hebei	牛1 Niu 1	122	河北Hebei	粗城下六 Kuanchengxialiu
83	河北Hebei	杨家峪1号 Yangjiayu 1	123	河北Hebei	X19-94
84	河北Hebei	何家坟5 Hejiafeng 5	124	河北Hebei	塌山1号 Tashan 1
85	河北Hebei	M17-21	125	北京Beijing	京暑红 Jingshuhong
86	河北Hebei	波叶树 Boyeshu	126	北京Beijing	BJHJ1
87	河北Hebei	M18-16	127	北京Beijing	BJHH1
88	河北Hebei	1209	128	北京Beijing	北京8号 Beijing 8
89	河北Hebei	硕丰 Shuofeng	129	北京Beijing	黑8 Hei 8
90	河北Hebei	D5-71	130	北京Beijing	燕昌 Yanchang
91	河北Hebei	沙坡峪1号 Shapoyu 1	131	北京Beijing	燕红 Yanhong
92	河北Hebei	塔14 Ta 14	132	北京Beijing	燕丰 Yanfeng
93	河北Hebei	桑6 Sang 6	133	北京Beijing	怀丰 Huaifeng
94	河北Hebei	长南庄2号 Changnanzhuang 2	134	北京Beijing	燕平 Yanping
95	河北Hebei	杨家峪1-6 Yangjiayu 1-6	135	北京Beijing	良乡1号 Liangxiang 1
96	河北Hebei	桑1 Sang 1	136	北京Beijing	BJHW
97	河北Hebei	园门实生 Yuanmenshisheng	137	山东Shandong	红栗 Hongli
98	河北Hebei	后南峪丰收1号 Hounanyufengshou1	138	山东Shandong	徐家1号 Xujia 1
99	河北Hebei	周家峪6号 Zhoujiayu 6	139	山东Shandong	尖顶油栗 Jiandingyouli
100	河北Hebei	H7-5	140	山东Shandong	双合大红袍 Shuanghedahongpao
101	河北Hebei	H8-1	141	山东Shandong	华光 Huaguang
102	河北Hebei	燕明 Yanming	142	山东Shandong	华丰 Huafeng
103	河北Hebei	兴隆早栗子 Xinglongzaolizi	143	山东Shandong	杂35 Za 35
104	河北Hebei	迁西署红 Qianxishuhong	144	山东Shandong	野杂 Yeza
105	河北Hebei	东荒峪早 Donghuangyuzao	145	山东Shandong	黄棚 Huangpeng
106	河北Hebei	青刘1号 Qingliu 1	146	山东Shandong	沂蒙短枝 Yimengduanzhi
107	河北Hebei	迁李1号 Qianli 1	147	山东Shandong	泰栗1号 Taili 1
108	河北Hebei	南天门乡 Nantianmenxiang	148	山东Shandong	金丰 Jinfeng
109	河北Hebei	大兰口 Dalankou	149	山东Shandong	红光 Hongguang
110	河北Hebei	昌徐1号 Changxu 1	150	山东Shandong	泰安薄壳 Taianbaoke
111	河北Hebei	昌左1号 Changzu 1	151	山东Shandong	新选1号 Xinxuan 1
112	河北Hebei	邢陆1号 Xinglu 1	152	山东Shandong	石丰 Shifeng
113	河北Hebei	迁西牛1 Qianxiniu 1	153	山东Shandong	早实丰Zaoshifeng
114	河北Hebei	去暑红-1 Qushuhong-1	154	山东Shandong	大粒 Dali
115	河北Hebei	去暑红-2 Qushuhong-2	155	山东Shandong	郯城3号 Tancheng 3
116	河北Hebei	邢台短枝 Xingtaiduanzhi	156	山东Shandong	金平 Jinping
117	河北Hebei	替码珍珠 Timazhenzhu	157	山东Shandong	东岳早丰 Dongyuezaofeng
118	河北Hebei	迁早1 Qianzao 1	158	山东Shandong	岱岳早丰 Daiyuezaofeng
159	山东Shandong	黑烟青 Heiyanqing	186	江苏Jiangsu	焦扎 Jiaozha
160	山东Shandong	烟泉 Yanquan	187	江苏Jiangsu	焦杂 Jiaoza
161	山东Shandong	烟青 Yanqing	188	江苏Jiangsu	宜兴大红袍 Yixingdahongpao
162	山东Shandong	鲁栗3号 Luli 3	189	江苏Jiangsu	处暑红 Chushuhong
163	山东Shandong	鲁栗2号 Luli 2	190	江苏Jiangsu	青毛软刺 Qingmaoruanci
164	湖北Hubei	罗田乌壳栗 Luotianwukeli	191	江苏Jiangsu	重阳蒲 Chongyangpu
165	湖北Hubei	DL-1	192	江苏Jiangsu	九家种 Jiujiazhog
166	湖北Hubei	DL-2	193	浙江Zhejiang	浙早1号 Zhezao 1
167	湖北Hubei	DL-3	194	浙江Zhejiang	上光栗 Shangguangli
168	湖北Hubei	DL-4	195	浙江Zhejiang	毛板红 Maobanhong
169	湖北Hubei	六月暴 Liuyuebao	196	浙江Zhejiang	浙江魁栗 Zhejiangkuili
170	湖北Hubei	八月红 Bayuehong	197	安徽Anhui	大红袍 Dahongpao
171	湖北Hubei	玫瑰红 Meiguihong	198	安徽Anhui	软刺早 Ruancizao
172	湖北Hubei	鄂栗1号 Eli 1	199	安徽Anhui	粘底板 Niandiban
173	湖北Hubei	罗田林科所Luotianlinkesuo	200	广西Guangxi	广西14-1 Guangxi 14-1
174	湖北Hubei	桂花香 Guihuaxiang	201	广西Guangxi	广西14-2 Guangxi 14-2
175	湖南Hunan	桐优-33 Tongyou-33	202	广西Guangxi	广西14-3 Guangxi 14-3
176	湖南Hunan	双季栗 Shuangjili	203	广西Guangxi	广西14-4 Guangxi 14-4
177	湖南Hunan	安优1号 Anyou 1	204	广西Guangxi	广西14-5 Guangxi 14-5
178	湖南Hunan	步城优株 Buchengyouzhu	205	广西Guangxi	广西14-6 Guangxi 14-6
179	湖南Hunan	临湘优株1号 LinxiangYouzhu 1	206	广西Guangxi	广西油栗 Guangxiyouli
180	湖南Hunan	林场优株 Lichangyouzhu	207	广西Guangxi	广西玉林 Guangxiyulin
181	湖南Hunan	石门优株 Shimenyouzhu	208	陕西Shaanxi	长安明捡栗 Changanmingjianli
182	湖南Hunan	临湘优株2号 LinxiangYouzhu 2	209	陕西Shaanxi	金真晚栗 Jinzhenwanli
183	湖南Hunan	湖南黔栗 Hunanqianli	210	陕西Shaanxi	金真栗 Jinzhenli
184	湖南Hunan	黄花优株 Huanghuayouzhu	211	陕西Shaanxi	镇巴双法 Zhenbashuangfa
185	湖南Hunan	新田优株 Xintianyouzhu

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2 结果

2.1 板栗种质资源花序相关表型多样性分析

由表2可知,板栗15个花序表型性状的变异系数和遗传多样性指数不尽相同,呈现出较大的变异和丰富的多样性。其中雄花序比例是所有性状中唯一变异系数低于10%,极差低于均值50%的性状,可见该性状整体离散度低,遗传稳定度高,其余14个性状的变异系数均高于10%,其中每果枝雌花个数最大为60.66%,说明这些性状遗传变异相对较大,而每果枝雌花个数更能反映品种间的差异。进一步研究发现,描述花序形态的表型性状变异系数介于15.70%—26.07%,说明这些表型性状变异幅度差异较少,而描述花序数量的表型性状变异系数介于7.37%—60.66%,证明花序数量的表型性状变异幅度差异较大。序轴粗度比的多样性指数最大为1.99,两性花序比例的多样性指数最小为1.41,花序形态表型性状中多样性指数范围是1.53—1.99,均值1.77;花序数量表型中多样性指数范围是1.41—1.87,均值1.63,表明花序形态表型性状的遗传多样性程度高于花序数量表型性状。

Table 2
表2
表2211份板栗花序相关性状基本参数及多样性
Table 2Basic parameter value and diversity of traits related catkin for chestnut

表型性状 Morphological character	均值±标准差 Mean±SD	中位数 Median	众数 Mode	极差 Rang	变异系数 CV (%)	多样性指数 H'
花序长度Catkin length (cm)	17.95±3.79	17.69	17.00	19.33	15.70	1.74
花序粗度Catkin width (mm)	11.26±2.40	11.23	10.56	13.10	16.94	1.84
花序小花簇密度Density of catkin (numbers/cm)	8.25±1.97	8.08	8.00	11.96	17.97	1.53
花轴粗度Floral axis width (mm)	3.24±0.95	3.19	3.51	3.68	23.29	1.81
雄蕊长度Stamen length (mm)	4.01±1.12	4.01	3.86	6.59	21.02	1.80
花序长粗比Catkin length/ Catkin width	16.71±5.44	16.17	14.43	42.88	26.41	1.66
花轴长粗比Floral axis length / Floral axis width	59.82±21.57	56.60	56.80	84.30	26.07	1.76
序轴粗度比Floral sxis width/ Catkin width	0.30±0.09	0.28	0.25	0.93	21.91	1.99
花序形态表型均值Catkin types of phentotype mean					21.16	1.77
雄花序个数/果枝Male catkin numbers/ Bearing shoot	7.28±3.67	7.00	8.00	11.60	29.34	1.73
两性花序个数/果枝Bisexual catkin numbers/ Bearing shoot	0.89±1.06	0.81	1.00	3.51	58.19	1.41
枝雌花个数/果枝Female flower Numbers/ Bearing shoot	1.02±0.67	1.02	0.81	5.16	60.66	1.87
总花序个数/果枝Catkin numbers/ Bearing shoot	8.16±4.12	7.90	6.50	11.82	28.14	1.73
雌花个数/两性花序Female flower numbers/ Bisexual catkin	1.19±0.29	1.12	1.00	1.29	12.35	1.78
雄花序比例Male catkin numbers/ Catkin numbers	0.90±0.13	0.91	0.92	0.40	7.37	1.49
两性花序比例Bisexual catkin numbers/ Catkin numbers	0.10±0.13	0.09	0.08	0.40	58.03	1.41
花序数量表型均值Catkin quantitative phenotype mean					36.30	1.63
总均值Mean					28.23	1.70

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综上可知,花序形态表型性状离散程度低但分布频率普遍均匀,尽管变异较小,但多样性程度高,而花序数量表型性状虽然离散程度较高,但分布频率相对集中。因此,这类性状遗传变异范围较大,但多样性程度相对较低。

2.2 板栗花序表型在不同群体间的变异和多样性比较

由表3可知,板栗不同群体间15个花序表型性状的变异系数和多样性指数均存在差异。江苏群体的平均变异系数最大,为35.53;安徽群体的平均变异系数最小,为16.96。山东群体中有4个性状平均变异系数群体间最大,全部为花序形态表型性状;江苏群体有4个性状变异系数群体间最大,全部属于花序数量表型性状。安徽有5个性状变异系数群体间最小,其中5个属于花序形态性状,陕西有4个性状变异系数群体间最小,全部属于花序数量性状。可见,就整体而言,江苏群体的花序表型变异较丰富,而安徽群体的遗传更稳定。山东群体的丰富变异集中在花序形态性状上,而江苏群体的丰富变异更集中在花序数量性状上。安徽群体花序形态性状的遗传更稳定,而陕西群体花序数量性状遗传稳定性相对更高。

Table 3
表3
表3不同群体间板栗资源表型性状变异系数和多样性指数
Table 3Variation coefficient and diversity indexes of phenotypic traits of Chinese chestnut indifferent populations

性状 Trait	河北 Hebei		北京 Beijing		山东 Shandong		湖北 Hubei		湖南 Hunan		江苏 Jiangsu		浙江 Zhejiang		安徽 Anhui		广西 Guangxi		陕西 Shaanxi
性状 Trait	CV (%)	H'	CV (%)	H'	CV (%)	H'	CV (%)	H'	CV (%)	H'	CV (%)	H'	CV (%)	H'	CV (%)	H'	CV (%)	H'	CV (%)	H'
花序长度CL	14.03	2.02	9.15	1.70	19.34	1.67	13.82	1.41	10.48	1.80	18.04	1.28	10.86	1.04	9.11	0.64	20.55	1.67	24.90	1.39
花序粗度CW	22.80	2.05	20.54	1.68	20.85	1.74	23.84	1.47	28.71	1.41	11.31	1.75	4.64	1.04	16.41	1.10	38.40	1.21	24.83	1.39
花序小花簇密度 DC	17.86	1.80	10.64	1.68	11.82	1.97	29.19	1.29	9.74	1.89	28.89	0.96	5.38	1.04	18.18	0.64	15.38	1.73	7.56	1.39
花轴粗度FAW	15.40	2.02	8.39	2.02	26.61	1.93	13.07	1.67	16.35	1.50	9.05	1.55	23.21	1.04	5.15	1.10	16.73	1.21	20.53	1.04
雄蕊长度SL	19.09	2.07	15.30	1.86	34.80	1.78	17.70	1.64	15.47	1.72	11.08	1.75	30.64	1.04	7.91	1.10	14.22	1.39	20.39	1.04
花序长粗比 CL/CW	19.45	1.96	14.54	1.82	48.34	1.41	17.45	1.64	19.37	1.85	23.55	1.75	20.21	1.04	5.06	1.10	25.89	1.56	9.67	1.39
花轴长粗比 FAL/FAW	25.87	1.96	24.29	1.86	29.90	2.01	18.79	1.64	26.92	1.89	24.39	1.55	14.07	1.39	17.43	1.10	31.35	1.49	21.07	1.39
序轴粗度比 FAW/CW	21.72	2.00	22.57	1.94	26.23	1.79	23.66	1.42	17.13	1.77	11.67	1.75	19.25	1.39	14.24	1.10	23.53	1.21	14.31	1.39
雄花序个数/果枝 MCN/BS	27.65	2.05	35.36	1.42	39.13	1.89	29.70	1.77	27.45	1.55	29.11	1.55	49.58	1.04	17.06	1.10	32.58	1.56	16.69	1.39
两性花序个数/果枝 BCN/BS	50.87	1.99	69.10	1.27	65.43	1.75	61.65	1.39	42.41	1.47	98.74	1.28	35.81	1.39	36.08	1.10	48.18	1.49	31.58	1.04
雌花个数/果枝 FFN/BS	50.37	1.42	80.96	0.25	67.00	0.42	58.90	0.19	42.73	0.20	97.06	0.16	45.57	0.10	44.08	0.05	45.76	0.15	29.61	0.08
总花序个数/果枝 CN/BS	26.41	2.06	36.30	1.82	29.24	1.89	29.03	1.55	26.90	1.59	27.25	1.28	44.49	1.04	17.26	1.10	29.47	1.73	15.84	1.39
雌花个数/两性花序 FFN/BC	12.64	1.77	12.91	1.63	8.47	2.01	17.07	1.41	8.53	1.54	6.18	1.00	10.50	0.56	10.64	1.10	16.83	1.21	7.36	1.04
雄花序比例 MCN/CN	6.52	1.97	10.96	1.35	8.84	1.84	5.76	1.85	4.21	1.85	11.67	1.28	5.37	1.04	0.75	1.10	0.37	1.39	1.97	1.04
两性花序比例 BCN/CN	52.75	1.94	68.36	1.35	60.00	1.89	47.99	1.77	39.07	1.97	122.47	1.28	59.91	1.04	17.42	1.10	55.41	1.39	21.55	1.04
均值Mean	25.91	1.86	29.08	1.51	32.97	1.66	26.46	1.40	20.97	1.50	35.53	1.30	23.72	0.95	16.96	0.95	28.14	1.32	19.15	1.13

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对不同群体板栗资源表型性状H′值进行分析可知,河北群体的平均H′最高为1.86,并且有11个性状H′都是在群体间最高。浙江和安徽群体的平均H′最低为0.95,其中浙江群体有花序粗度、花轴粗度、雄蕊长度、花序长粗比、每两性花序上雌花个数、每果枝总花序个数、每果枝雄花序个数、雄花序比例和两性花序比例9个性状H′最低,安徽群体有花序长度、花序小花簇密度、花轴长粗比、序轴粗度比、每果枝雌花个数5个性状H′最低。由此可知,河北板栗群体的性状频率分布较均匀,浙江和安徽群体的性状频率分布较集中。

2.3 板栗不同群体间的花序表型变异特征

由表4、5可知,15个板栗花序表型性状中,除雄蕊长度和序轴粗度比两个性状无明显差异外,其余13个性状在群体间的差异均达到显著水平,这种相同性状在不同群体差异明显的特点,说明长期的环境异质对群体表型差异有显著影响。北京、广西、河北、山东4个群体资源果枝雌花比较高,广西、山东、湖南、浙江、陕西群体雄花序花簇较稀疏。进一步从群体所处地理位置上看,板栗种质自北向南呈现出雄花序由细长渐变为粗短,雄花序比例逐渐升高,两性花序比例和每果枝雌花数逐渐减少的表型变化特征。

Table 4
表4
表4不同群体间板栗花序形态比较
Table 4Comparision of Catkin morphology for Chinese chestnut in different populations

群体 Population	花序长度 CL (cm)	花序粗度 CW (mm)	花序小花簇密度 DC (No./cm)	花轴粗度 FAW (mm)	雄蕊长度 SL (mm)	花序长粗比 CL/CW	花轴长粗比 FAL/FAW	序轴粗度比FAW/CW
河北Hebei	18.38±2.58bB	11.39±1.75abA	16.51±2.95aAB	3.27±0.75abcAB	4.07±0.78aA	16.71±3.25abA	60.58±15.67abA	0.29±0.06aA
北京Beijing	18.38±1.68bB	11.57±0.97abA	17.21±1.83aAB	3.62±0.74bcAB	3.96±0.61aA	16.17±2.35abA	54.43±13.22abA	0.38±0.07aA
山东Shandong	17.64±3.41bB	10.77±2.87abA	16.12±1.91aA	3.06±0.64abcAB	3.86±1.34aA	18.57±8.98abA	62.06±18.54abA	0.30±0.08aA
湖北Hubei	18.48±2.55bB	11.37±1.49abA	17.81±5.20abAB	3.48±0.83abcAB	3.95±0.70aA	16.59±2.90abA	56.73±10.66abA	0.31±0.07aA
湖南Hunan	17.22±1.80bAB	11.35±1.86abA	15.43±1.50aA	3.06±0.88abcAB	4.15±0.64aA	15.82±3.06abA	63.28±17.04abA	0.27±0.05aA
江苏Jiangsu	16.16±2.91bAB	10.52±0.95abA	17.79±5.14abAB	2.94±0.33abcAB	3.80±0.42aA	15.74±3.71abA	56.93±13.89abA	0.28±0.03aA
浙江Zhejiang	17.06±1.85bAB	11.56±2.68abA	16.06±0.86aA	3.15±0.15abcAB	4.21±1.29aA	15.42±3.12abA	57.00±8.02abA	0.28±0.05aA
安徽Anhui	17.17±1.56bAB	12.21±0.63bA	20.83±3.79bB	3.58±0.63cB	4.18±0.33aA	14.24±0.72abA	46.62±8.13aA	0.32±0.05aA
广西Guangxi	13.34±2.74aA	10.26±1.72abA	14.90±2.29aA	2.88±1.11abAB	3.68±0.52aA	13.65±3.53aA	56.54±17.73abA	0.28±0.07aA
陕西Shaanxi	18.04±4.49bB	9.27±1.90aA	16.33±1.23aA	2.64±0.66aA	3.32±0.68aA	19.61±1.90bA	70.38±14.83bA	0.29±0.04aA

大小写字母分别表示不同群体间1%和5%水平差异显著。下同

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The capital and lowercase letters indicate significant differences at the 0.01 and 0.05 level probability. The same as below

Table 5
表5
表5不同群体间板栗花序数量比较
Table 5Comparision of Catkin numbers for Chinese chestnut in different populations

群体 Population	雄花序个数/果枝MCN/BS	两性花序个数/果枝BCN/BS	雌花个数/果枝FFN/BS	总花序个数/果枝CN/BS	雌花个数/两性花序FFN/BC	雄花序比例 MCN/CN	两性花序比例BCN/CN
河北Hebei	7.14±1.95abA	0.97±0.50abAB	1.12±0.57abcAB	8.11±2.10abA	1.16±0.15abA	0.89±0.06abAB	0.11±0.06abAB
北京Beijing	7.75±2.74abA	1.29±0.89bB	1.62±1.31cB	9.04±3.28abA	1.21±0.16bA	0.86±0.09aAB	0.14±0.09bAB
山东Shandong	7.02±2.01abA	0.99±0.64abAB	1.10±0.73abcAB	8.01±2.29abA	1.11±0.10abA	0.88±0.07abAB	0.12±0.07abAB
湖北Hubei	6.37±1.86aA	0.74±0.45abAB	0.81±0.48abAB	7.08±2.05aA	1.13±0.19abA	0.90±0.06abAB	0.11±0.05abAB
湖南Hunan	6.50±1.78aA	0.71±0.30abAB	0.81±0.35abAB	7.22±1.94abA	1.15±0.10abA	0.90±0.04abAB	0.10±0.04abAB
江苏Jiangsu	8.89±2.59abA	0.63±0.63abAB	0.66±0.64abAB	9.53±2.59abA	1.04±0.06aA	0.91±0.11abAB	0.09±0.11abAB
浙江Zhejiang	8.33±4.13abA	0.63±0.23abAB	0.67±0.31abAB	8.95±4.00abA	1.06±0.11aA	0.92±0.05abAB	0.08±0.05abAB
安徽Anhui	9.53±1.62bA	0.35±0.13aA	0.41±0.18aA	9.89±1.70bA	1.13±0.12abA	0.96±0.01bB	0.04±0.01aA
广西Guangxi	7.31±2.38abA	1.23±0.60bB	1.37±0.63bcAB	8.55±2.52abA	1.11±0.19abA	0.84±0.09aA	0.16±0.09bB
陕西Shaanxi	7.98±1.33abA	0.71±0.23abAB	0.83±0.25abAB	8.68±1.38abA	1.18±0.09abA	0.92±0.02abAB	0.08±0.02abAB

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2.4 板栗花序表型性状相关性分析

由表6可知,8个花序形态表型性状多呈显著或极显著相关性,7个花序数量表型性状亦呈现相同特征,但大多数花序形态和数量表型性状间相关性不显著。序轴粗度比与每果枝总花序个数和雄花序个数呈极显著正相关,与每果枝两性花序个数显著正相关;每两性花序上雌花个数与每果枝雌花个数极显著正相关,与每果枝总花序个数、每果枝雄花序个数显著负相关。每果枝雌花个数与每果枝总花序个数、每果枝雄花序个数、每果枝两性花序个数和两性花序比例成极显著正相关,与雄花序比例极显著负相关。综上可知,板栗花序花轴越粗,果枝上的总花序、雄花序和两性花序越多;两性花序上的雌花越多,则果枝上的总雌花数越多;果枝上的总雌花数越大,花序、雄花序、混合花序的数量都随之增大,但混合花序比例升高,雄花序比例降低。

2.5 板栗花序表型性状主成分分析

以特征值大于1.0为标准提取主成分。由表7可知,花序表型性状中前5个主成分的特征值均大于1,说明这5个主成分在板栗花序表型变异性构成中作用较大,累计反映了总信息量的84.18%,其中第1主成分贡献率占23.52%,对它作用最大的性状包括每果枝两性花序个数（0.931）、每果枝雌花个数(0.828)、雄花序比例（-0.821）和两性花序比例(0.820),由此可知第1主成分主要是和花序数量有关的性状并与雌花数量呈正相关;第2主成分占21.10%,对它作用最大的性状为花序粗度(0.912)、花序长粗比(-0.889)、雄蕊长度（0.806）,说明第2主成分主要是和花序形态有关的性状;第3、4和5主成分贡献率分别占17.537%、11.963%和10.065%。每果枝两性花序个数(0.931)、花序粗度(0.912)、花序长粗比(-0.889)、花序长度（0.864）、每果枝雌花个数（0.828）、雄花序比例（-0.821）、两性花序比例（0.820）、雄蕊长度（0.806）这8个性状的相关系数都在0.8以上,认为其是造成板栗花序表型变异的主要因素,与上述变异系数分析结果基本吻合。以第1主成分为横坐标,第2主成分为纵坐标做成散点图（图1）,将211份板栗资源划分为5大类群。类群Ⅰ只有1份资源,来源于山东,花序短粗,雌花量较少;类群Ⅱ包括了大部分板栗种质,性状表现比较集中;类群Ⅲ有5份资源,来源于河北、北京和广西,果枝上雌花丰富;类群Ⅵ有2份资源,全部来自河北,花序长粗比例大;类群Ⅴ有6份资源,来源于山东、北京、河北和江苏,果枝上雌花多、花序细小。对板栗高产育种来说,选育两性花序比例高、每果枝雌花个数多的资源具有重要意义,以横坐标高于2.0为标准选出高雌花量种质9份。

2.6 板栗花序表型性状聚类分析

由图2可知,通过聚类分析,在遗传距离为0.38处将211份板栗资源聚为8大类群。第1类群有43份资源,占20.38%,主要来源于华北地区的群体,特征以花序粗且长,花轴粗,小花簇稀疏,雄蕊长度中等,总花序量中等,两性花序占比较低,雌花数量少为主;第2类群包含1份资源,来源为广西地区,特征为花序细短,花轴细弱,花簇密度比较稀疏,雄蕊较短,总花序量中等,两性花序占比中庸,雌花数量中等;第3类群包含155份资源,占73.46%,来源涵盖了10个群体,特征以花序粗且长,轴粗中等,花簇密度比较稀疏,总花序量较少,两性花序占比较少,雌花数量少的资源为主;第4类群包含6份资源,来源为华北、安徽、广西地区,特征以花序长度、粗度中等,花簇较密集,总花序量较少,两性花序占比较少,雌花数量少的资源为主;第5类群包含2份资源,来源于山东,特征为花序细长,花簇稀疏,雄蕊较短,总花序量大,两性花序占比高,雌花数高;第6大类群包含1份资源,来自山东,特征为花序短粗,花轴纤细,花簇密度比较稀疏,雄蕊较长,总花序量少,两性花序占比低,雌花数少;第7大类群包含1份资源,来源于广西地区,特征为花序较粗,长度中等,花簇密度中等,总花序量大,两性花序占比高,雌花数多;第8类群包含2份资源,来源于河北和江苏,特征为花序长度、粗度中等,花簇密集,总花序量多,两性花序占比中等,雌花量多。

Table 6
表6
表6板栗花相关表型性状相关性分析
Table 6Correlation analysis of phenotypic traits related catkin for Chinese chestnut

性状 Trait	花序长度 CL	花序粗度CW	花序小花簇密度DC	花轴粗度FAW	雄蕊长度 SL	花序长粗比 CL/CW	花轴长粗比 FAL/ FAW	序轴粗度比FAW/ CW	雄花序个数/ 果枝 MCN/ BS	两性花序个数/果枝BCN/ BS	雌花个数/果枝FFN/BS	总花序个数/ 果枝CN/BS	雌花个数/两性花序FFN/BC	雄花序比例MCN/ CN	两性花序比例BCN/ CN
花序长度CL	1
花序粗度CW	0.133	1
花序小花簇密度DC	-0.174^*	0.088	1
花轴粗度 FAW	0.241^**	0.472^**	0.063	1
雄蕊长度SL	0.051	0.920^**	0.068	0.092	1
花序长粗比 CL/CW	0.496^**	-0.698^**	-0.202^**	-0.248^**	-0.674^**	1
花轴长粗比 FAL/FAW	0.352^**	-0.442^**	-0.182^**	-0.776^**	-0.154^*	0.592^**	1
序轴粗度比FAW/CW	0.135^*	-0.316^**	-0.009	0.657^**	-0.644^**	0.376^**	-0.469^**	1
雄花序个数/果枝MCN/BS	0.0.14	-0.046	0.136^*	0.137^*	-0.116	0.048	-0.113	0.187^**	1
两性花序个数/果枝BCN/BS	0.126	0.025	-0.046	0.164^*	0.043	0.082	-0.090	0.173^*	0.208^**	1
雌花个数/果枝FFN/BS	0.072	-0.016	-0.051	0.094	-0.059	0.087	-0.068	0.127	0.196^**	0.789^**	1
总花序个数/果枝CN/BS	0.043	-0.037	0.115	0.166^*	-0.117	0.064	-0.125	0.214^**	0.972^**	0.432^**	0.370^**	1
雌花个数/两性花序FFN/BC	0.138^*	0.044	-0.157^*	-0.037	0.067	0.057	0.116	-0.018	-0.143^*	-0.007	0.188^**	-0.135^*	1
雄花序比例MCN/CN	-0.074	-0.036	0.108	-0.064	-0.017	-0.034	0.039	-0.050	0.241^**	-0.788^**	-0.620^**	0.032	-0.005	1
两性花序比例BCN/CN	0.076	0.043	-0.118	0.069	0.023	0.030	-0.045	0.051	-0.250^**	0.787^**	0.624^**	-0.041	0.020	-0.996^**	1

* and ** indicate significant correlation at P<0.05 and P<0.01, respectively
*和**分别表示在P<0.05和P<0.01水平上显著相关

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3 讨论

花序是板栗的重要器官,其形态大小,两性花序的比例等对产量影响巨大^[1,2,3]。对板栗种质资源的花序性状进行多样性分析,是开展育种的基础工作,同时也是一种简便有效的筛选特异种质的方法。板栗花序相关性状是一类重要的表型性状,受遗传和环境效应的影响,这类性状表现出复杂的数量性状特点。由于在个体或微观水平上对表型性状进行研究,较难取得突出进展^[24],因此,本研究从全国10个省份（群体）选取211份资源开展花序性状研究,这10个省份涵盖了中国板栗全部5个地方品种群即华北品种群、长江中下游品种群、西北品种群、东南品种群、西南品种群^[1],累计栽培面积约占中国板栗栽培面积的85%^[25],区域内资源丰富,就种质分布而言有较强的代表性,选取这10个省份（群体）资源开展研究,有利于从宏观水平上开展整体趋势分析和探讨,以便更准确探明性状的遗传变异规律。但10个群体中陕西、安徽、浙江等群体在此次研究中选取资源较少,因此板栗花相关的遗传多样性信息有待进一步完善。变异系数和多样性指数是植物多样性分析最常用的两个指标,变异系数越高,表明性状的离散程度越大、变异幅度越大;而多样性指数越高,表明性状多样性程度越丰富、种类越多^[26]。在本研究中,我国板栗资源花序相关性状表现出丰富的变异和表型多样性,其中花序形态表型性状离散程度低但分布频率均匀,整体变异相对较小但多样性程度高,花序数量表型离散程度较高但分布频率相对集中,所以变异范围更大,但多样性程度相对较低,这种现象说明,在板栗的进化过程中,相较于花序的形态,花序数量的变化更大,但分布更集中,这与人们长久以来的育种目的趋于一致,证明板栗资源整体的遗传变异一直在受人为选择的影响。

图1

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图1211份板栗资源花表型性状的主成分分析图

Fig. 1The principal component analysis of 211 Chinese chestnut germplasms

图2

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图2板栗资源聚类图

Fig. 2Cluster dendrogram of Chinese chestnut

Table 7
表7
表7板栗花相关表型性状的主成分矩阵、特征向量和贡献率
Table 7The eigenvalue and contributive percentage of principal components and component scores coefficient matrix of t phenotypic traits related catkin for Chinese chestnut

性状编号 Character code	主成分 Principal component
性状编号 Character code	1主成分 First principal component	2主成分 Second principal component	3主成分 Third principal component	4主成分 Fourth principal component	5主成分 Fifth principal component
花序长度CL	0.194	-0.203	-0.073	-0.073	0.864
花序粗度CW	-0.068	0.912	-0.089	0.063	0.361
花序小花簇密度DC	-0.098	0.217	0.263	0.154	-0.336
花轴粗度FAW	0.305	0.502	0.528	-0.512	0.309
雄蕊长度SL	-0.208	0.806	-0.338	0.295	0.276
花序长粗比CL/CW f	0.203	-0.889	0.015	-0.094	0.274
花轴长粗比FAL/FAW	-0.173	-0.676	-0.504	0.404	0.214
序轴粗度比FAW/CW	0.401	-0.232	0.639	-0.602	0.034
雄花序个数/果枝MCN/BS	0.203	-0.045	0.729	0.619	0.111
两性花序个数/果枝BCN/BS	0.931	0.086	-0.097	0.196	-0.043
雌花个数/果枝FFN/BS	0.828	0.043	-0.101	0.225	-0.011
总花序个数/果枝CN/BS	0.411	-0.021	0.649	0.618	0.091
雌花个数/两性花FFN/BC	0.023	-0.036	-0.263	-0.046	0.390
雄花序比例MCN/CN	-0.821	-0.117	0.472	0.097	0.165
两性花序比例BCN/CN	0.828	0.043	-0.101	0.225	-0.011
特征值Eigenvalue	3.527	3.165	2.631	1.794	1.510
贡献率 Contributive percentage (%)	23.516	21.098	17.537	11.963	10.065
累计贡献率 Cumulative contributive percentage (%)	23.516	44.613	62.150	74.113	84.178

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板栗雄花多、雌花少的特性导致营养分配、物质疏导等方面的不均衡,雄花耗养量大,是限制板栗产量提高的重要因素,生产上常需要采取多种方法解决,如配方施肥、施用微量元素和生长调节剂、人工疏雄等措施,虽可增加雌雄花的比率和坐果率,但也增加了生产成本,所以选育雌花多、雄花序短小、花簇稀疏或雌雄比例大的丰产优质品种,是解决板栗低产的最有效途径之一。由本研究群体看,板栗种质资源自北向南呈现出雄花序由细长渐变为粗短、雄花序比例逐渐升高、两性花序比例和每果枝雌花数逐渐减少的表型变化特征,不同地理来源的板栗资源同一花序表型差异明显,这一结果与江锡兵等^[12]的研究相似,说明长期环境异质可导致群体表型的明显差异,推断复杂多变的地理特征和气象条件,悠久的栽培历史,长期的自然和人工选择,是板栗形成具有特定适应性的地方群体资源的共同作用因子。本研究中,地理来源不同的各群体在性状表型值、变异系数及多样性程度上都有显著的差异,相同群体内的不同性状,差异程度也不尽相同,说明基因型差异和环境异质导致了不同群体间表型变异的不同,体现了群体间种质在育种上的不等值性。此现象与笔者之前对板栗枝条多样性的研究结果一致^[13]。地理隔离是增加多样性的一个重要因素^[27],结合本研究结果,在育种工作中可以根据不同群体间的特点进行亲本选配,打破地理隔离,更易选育出杂合性更高、适应性更强、超亲更明显的品种。比如选取北方的河北群体和南方的广西群体内资源进行杂交,其后代雄花序相关性状变异幅度和多样性水平可能更大,更容易聚合出雄花序短粗、每果枝雌花数等综合品质更优秀的高产品种。

性状相关性分析可以看出,多数花序形态性状和数量性状间相关性不显著,但花轴粗度与两种花序个数显著正相关,花轴越粗,雄花序数和两性花序数都随之变大,由此推测,花轴粗度可以作为判断板栗花量多少的一个佐证。本研究还发现总雌花数量与雄花序数正相关,但与雄花序占比为负相关,因此在育种过程中,不能因为雄花序养分消耗较大^[15],一味选择雄花序数量少的资源,而更应该关注雌雄比例适当的资源。

主成分分析可将多个变量化为少数几个指标,从而简化分析过程,更好地描述总变异构成特征^[28,29],因此用主成分分析来筛选资源,这样筛选出来的资源不是某单一性状突出,而是相关性状综合优质,在育种上更有意义。本研究中,第一主成分的特征向量与花序数量性状有关,第二主成分的特征向量与花序形态有关。由第一、二主成分为坐标做出的散点图中,类群Ⅰ中变异大的‘徐家1号’在聚类分析中被单独聚出,类群Ⅴ中离其他资源较远的‘143野杂’和‘泰安薄壳’在聚类分析中也被单独聚为一类,类群Ⅴ中的‘BJHH1’在聚类分析中虽未被单独聚类,但从图中也可看出‘BJHH1’与其他资源有较远的遗传距离。可见,本研究中主成分分析所分类群与聚类分析结果相似,证明主成分分析中贡献率较高的性状确实能反映资源变异的主要特点。这一结果与李鸿雁等^[30]的研究结果一致。依据主成分分析结果,鉴定筛选出9份两性花数量大、雌雄比例高的高雌花量资源（84-2、长南庄2号、BJHJ1、BJHH1、北京8号、沂蒙短枝、焦杂、广西14-4）,这9份特异资源可以在今后的育种工作中作为花性别调控、提高产量的亲本材料。

描述和分析植物表型性状在种质资源研究的过程中是最基本的方法和不可替代的途径。根据目前的研究结果,在作物种质研究中,已有许多通过利用综合评价种质资源来构建核心种质资源库的成功案例,并成功地提高了育种效率^{[31,32,33,34,35,36,37]},表型性状已成为生物学研究的重要命题,基因组和转录组等生物信息学数据只有和植物表型性状互相结合,共同分析才能解析遗传的特点和机理^[38,39]。由此可知,充分挖掘表型数据并整理分析,将有助于推动育种学跨越式的发展。

4 结论

中国板栗资源花相关表型性状变异丰富,多样性程度高,地域间遗传变异和多样性程度不同,群体间性状差异显著。花序形态表型性状和花序数量表型性状内部的相关性较明显,但花序形态和花序数量间相关性不明显。综合相关性分析和主成分分析,筛选出每果枝两性花序个数、花序粗、花序长粗比、花序长、每果枝雌花数、雄花序数/花序数、两性花序数/花序数、雄蕊长8个性状可作为板栗花形和花量的综合评定指标;84-2、长南庄2号、BJHJ1、BJHH1、北京8号、沂蒙短枝、焦杂、广西14-4共9个高雌花量的板栗资源可作为性别调控、改善产量的亲本材料。聚类分析将211份资源分为8个类群。本研究为进一步丰富中国板栗表型性状遗传多样性信息,挖掘优异基因材料和现有种质资源保护、利用、创新及遗传改良奠定了基础。

参考文献原文顺序
文献年度倒序
文中引用次数倒序
被引期刊影响因子

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, LIU

. Morpholgical properties of wild resources in Medicago ruthenica in Inner Mongolia
Journal of Plant Genetic Resources, 2011,12(5):721-726. (in Chinese)

[本文引用: 1]

[29]

张莹, 曹玉芬, 霍宏亮, 徐家玉, 田路明, 董星光, 齐丹, 张小双, 刘超, 王立东 . 基于枝条和叶片表型性状的梨种质资源多样性
中国农业科学, 2018,51(17):3353-3369.

DOI:10.3864/j.issn.0578-1752.2018.17.010 URL [本文引用: 1]

【Objective】The study on the diversity and variation of current-year twig and leaf phenotypic traits of pear germplasm resources was conducted in order to provide valuable basic data and theory foundation for normalization, standardization, preservation and construction of pear core collections, and to promote the efficient utilization of pear germplasm resources.【Method】Data were collected for 23 phenotypic parameters of current-year twig and leaf from 548 accessions of 13 Pyrus species preserved in National Germplasm Repository of Apple and Pear according to the method described in Descriptors and Data Standard for Pear (Pyrus spp.) methods. The distribution frequency, coefficient of variation, Simpson index, Shannon-weaver index, correlation and principal component analysis of pear current-year twig and leaf were analyzed using the SPSS19.0 software, and the intraspecific and interspecific genetic diversities of pear were also analyzed and compared. The frequency distributions of quantitative characters were analyzed by Origin 8.0. The crisp-fleshed and soft-fleshed pears were clustered using MEGA 5.0, respectively, according to morphological data.【Result】Analyzing of 15 character traits of pear leaf phenotype showed that 8 out of 15 traits were abundant, namely, ovate shape, wide wedge-shaped base, sharp-acuate apex, serrate on leaf margin with seta, enclasped status of leaf surface, downward latitude of leaf and redish-green young leaf, which accounted for 90.51%, 58.03%, 66.97%, 81.93%, 87.23%, 59.27%, 86.68% and 35.04%, respectively. Regarding the phenotype of current-year twig, which was ample among yellow brown, rich in lenticels, leaf bud slightly held out, obtuse leaf bud apex, size of bud support medium, pubescence on flower bud absent, which accounted for 87.23%, 78.28%, 87.96%, 83.76%, 73.91% and 99.27%, respectively. The Shannon indexes of the color of young leaf and leaf base shape were found to be as high as 2.197 and 1.597, respectively. The analyses of 8 numeric traits indicated that the average coefficient of variation of leaf length, leaf width, petiole length, current-year twig length, twig thickness, internode length, length of flower bud and thickness of flower bud was 17.25%, 19.04%, 20.06%, 23.70%, 15.08%, 19.33%, 20.62% and 16.66%, respectively. Reference cultivars and 5 groups of each trait were proposed based on the statistical analysis of frequency distribution of numeric traits of current-year twig and leaf. Eight numeric traits, including current-year twig, length of flower bud, leaf width and petiole length, were put forward as comprehensive assessment indexes according to the results of correlation and principal component analyses. There were significant differences in 8 pear numeric traits of current-year twig and leaf among and within populations, while phenotypic differentiation coefficient (VST) of intraspecies and interspecies were 41.10% and 58.90%, respectively. Cluster analysis showed that 233 crisp-fleshed local pear cultivars could be divided into 12 categories and 87 soft-fleshed P. ussuriensis accessions into 6 categories. It was worth mentioning that pear resources which was from southwest China were found in most of the groups.【Conclusion】There were abundant genetic diversity based on the phenotype of current-year twig and leaf of pear. The diversities of character traits of color of young leaf and shape of leaf base were higher than the others. The variation coefficients of numeric traits of current-year twig and length of flower bud were also more obvious than the others. These 4 traits can therefore reflect the differences among pear varieties. The variation in pear germplasm resources based on twig and leaf phenotype traits among populations was higher than within populations, suggesting that the variation among populations was the main variation source. Finally, 5 numeric traits were selected to be as the important comprehensive evaluation indexes used for pear germplasm resources.

ZHANG

, CAO Y

, HUO H

, XU J

, TIAN L

, DONG X

, QI

, ZHANG X

, LIU

, WANG L

. Diversity of pear germplasm resources based on twig and leaf phenotypic traits
Scientia Agricultura Sinica, 2018,51(17):3353-3369. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2018.17.010 URL [本文引用: 1]

李鸿雁, 李志勇, 黄帆, 师文贵, 李俊, 刘磊, 解永凤 . 内蒙古扁蓿豆种质资源花性状的变异分析
植物遗传资源学报, 2015,16(6):1223-1228.

[本文引用: 1]

LI H

, LI Z

, HUANG

, SHI W

, LI

, LIU

, XIE Y

. Analysis of flower character variation of Medicago ruthenica in Inner Mongolia
Journal of Plant Genetic Resources, 2015,16(6) : 1223-1228. (in Chinese)

[本文引用: 1]

[31]

LIU

, KAKIHARA

, KATO

. Characterization of six varieties of Cucumis melo L. based on morphological and physiological charac-ters, including shelf-life of fruit
Euphytica, 2004,135:305-313.

DOI:10.1023/B:EUPH.0000013330.66819.6f URL [本文引用: 1]

Seventy-two accessions covering six varieties of Cucumis melo were characterized by using 35 morphological characters with emphasis on shelf-life, and the relationships between shelf-life and related characters was investigated. Principal component analysis (PCA) revealed that development period of plant and fruit, size of seed and fruit, shelf-life, stem hair, flesh juiciness, netting, abscission of peduncle, rapid yellowing of epidermis at maturity, Brix value, and color of flesh and epidermis etc. were the principal characters to discriminate melon accessions examined in the present study. According to the scatter diagram, vars. acidulusand makuwa, both of which belong to the Oriental melon, are closely related because of their short growth duration, small seed, thin pericarp and poor shelf-life, while American cantaloupe (var.reticulatus) and European cantaloupe (var. cantalupensis) are rather closely related due to their climacteric fruit with orange flesh, slipped peduncle and rapid yellowing of epidermis at maturity, which is closely related with their shelf-life. PCA also indicated that var. saccharinus was closer to var. inodorus than to the other varieties, due to their requirement of long period for development, large size of seed and fruit, and half- or non-slipped peduncle. Shelf-life of melon fruit was significantly correlated with the following characters: quality of flesh, size of seed and fruit, abscission of peduncle, development periods of plant and fruit, rapid yellowing of epidermis at maturity, Brix value and color of flesh and epidermis. Accessions with good shelf-life were mostly found in vars. saccharinus and inodorus.

[32]

王丽侠, 程须珍, 王素华, 李金榜, 李金秀 . 中国绿豆应用型核心样本农艺性状的分析
植物遗传资源学报, 2009,10(4):589-593.

[本文引用: 1]

WANG L

, CHENG X

, WANG S

, LI J

. Genetic Variability of agronomic traits in mungbean applied core collection of China
Journal of Plant Genetic Resources, 2009,10(4):589-593. (in Chinese)

[本文引用: 1]

[33]

SZAMOSI

, SOLMAZ

, SARI

, BARSONY

. Morphological evaluation andcomparison of Hungarian and Turkish melon ( Cucumis melo L.) germplasm
Scientia Horticulturae, 2010,124:170-182.

DOI:10.1016/j.scienta.2009.12.024 URL [本文引用: 1]

[34]

胡建斌, 马双武, 李建吾, 王吉明, 李琼, 王林忠 . 国外甜瓜种质资源形态性状遗传多样性分析
植物学报, 2013,48(1):42-51.

DOI:10.3724/SP.J.1259.2013.00042 URL [本文引用: 1]

对250份国外甜瓜(Cucumis melo)种质资源的19个形态性状多样性进行研究。结果表明, 国外甜瓜种质资源具有丰富的形态多样性, 平均遗传多样性指数为1.378。9个质量性状(果实形状、果皮底色、覆纹颜色、覆纹形状、网纹密度、网纹粗度、果肉颜色、果肉质地和种子颜色)和4个数量性状(单果鲜重、果肉厚度、可溶性固形物含量和种子千粒重)变异明显, 其Shannon’s指数分别大于1.0和1.9。不同生态区间种质资源遗传多样性差异明显, 多样性指数由高到低依次为: 南亚(1.512)、东北欧(1.404)、西欧(1.372)、北美(1.340)和东亚(1.281)。通过聚类分析将所有甜瓜种质划分为四大组群, 即南亚组群、东北欧组群、西欧北美组群和东亚北美组群。以印度为代表的南亚甜瓜种质形态多样性水平较高, 支持了印度次生大陆为甜瓜起源中心的观点。

HU J

, MA S

, LI J

, WANG J

, LI

, WANG L

. Genetic diversity of foreign melon (Cucumis melo) germplasm resources by morphological characters
Chinese Bulletin of Botany, 2013,48(1):42-51. (in Chinese)

DOI:10.3724/SP.J.1259.2013.00042 URL [本文引用: 1]

邱丽娟, 李英慧, 关荣霞, 刘章雄, 王丽侠, 常汝镇 . 大豆核心种质和微核心种质的构建验证与研究进展
作物学报, 2009,35(4):571-579.

DOI:10.3724/SP.J.1006.2009.00571 URL [本文引用: 1]

China has the most soybean germplasm in the world, but only about 1% of them have been used in soybean breeding program, which results in narrow genetic base for developing varieties. This exiguous use of germplasm is due to lack of reliable information on traits with economic importance, which requires replicated multilocational evaluations to identify useful parents. In order to accelerate evaluation and utilization of germplasm, the projects of ‘Establishment of soybean core collection’ (1998–2003) and ‘Gene diversity of mini core collection in soybean’ (2004–2008) were carried out by the continuous supporting from National Basic Research Program (973 project) in case of large germplasm collections has been evaluated properly on the basis of the guideline from core collections. In this paper, the development of core (2–5% of entire collection) and mini-core (10% of core or 1% of entire collection) is introduced. Meanwhile, the complement of the core or mini-core collections is carried out in order to broaden their representative from national wide into international soybean accessions. For testing the representatives of the core collection, four parameters including SSR loci, sample components, sampling ratio and low frequency allele are analyzed, and then confirmed by the random sampling test. The core and mini-core collections have been used in identifying elite traits and developing new lines, in which the approaches have been introduced that based on core and mini-core collections. The further utilization of core and mini-core collections related to genetic diversity and utilization method of collections is discussed, it is pointed that core collections are basic materials for new trait identification, novel gene mining, germplasm enhancement and new variety development. It has been proved that systematic study and use of soybean germplasm will improve the research of the soybean germplam from conservation into wider use.

QIU L

, LI Y

, GUAN R

, LIU Z

, WANG L

, CHANG R

. Establishment, representative testing and research progress of soybean core collection and mini core collection
Acta Agronomica Sinica, 2009,35(4):571-579. (in Chinese)

DOI:10.3724/SP.J.1006.2009.00571 URL [本文引用: 1]

王海飞, 关建平, 孙雪莲, 马钰, 宗绪晓 . 世界蚕豆种质资源遗传多样性和相似性的ISSR分析
中国农业科学, 2011,44(5):1056-1062.

URL [本文引用: 1]

【Objective】 The purpose of this research is to analyze the genetic diversity of global faba bean germplasm, to explore their genetic similarity and population structure, and to provide essential information for germplasm evaluation and effective utilization of faba bean genetic resources. 【Method】 The genetic similarity of 383 faba bean accessions from 35 countries was analyzed by using ISSR marker. 【Result】 Eleven ISSR primers generated 229 unambiguous bands, of which 212 were polymorphic, and the rate of polymorphic bands was 0.93. Gene diversity index (H) and allelic richness (NA) of different geographic groups of germplasm ranged from 0.16 to 0.28 and from 104 to 193, respectively. Chinese spring-seeding area group showed the highest genetic diversity (H = 0.28, NA = 193), and America group showed the lowest (H = 0.16, NA = 104). The spring faba bean germplasm was clearly separated from winter faba bean germplasm of China in UPGMA clustering analysis based on ISSR molecular marker data. Germplasm from China is quite distinct to that from exotic accessions. The accessions from Europe had a closer genetic similarity with that from North Africa. While the germplasm resources from Asia, Europe and Africa are closely related to their geographical distribution.【Conclusion】Accessions from spring-seeding area of China were most diverse. Germplasm from America showed lowest diversity. The results indicated that the genetic similarity and diversity of faba bean germplasm are closely associated with their growth habit, their geographical origin and ecological distribution.
Key words: faba bean (Vcia faba L.); germplasm resourses; ISSR marker; genetic diversity; genetic similarity

WANG H

, GUAN J

, SUN X

, MA

, ZONG X

. Genetic diversity and similarity of global faba bean (Vcia faba L.) germplasm revealed by ISSR markers
Scientia Agricultura Sinica, 2011,44(5):1056-1062. (in Chinese)

URL [本文引用: 1]

STEPANSKY

, KOVALSKI

, PERL-TRECES

. Intraspecific classification of melons ( Cucumis melo L.) in view of their phenotypic and molecular variation
Plant Systematics and Evolution, 1999,217:313-332.

DOI:10.1007/BF00984373 URL [本文引用: 1]

[38]

潘映红 . 论植物表型组和植物表型组学的概念与范畴
作物学报, 2015,41(2):175-186.

DOI:10.3724/SP.J.1006.2015.00175 URL [本文引用: 1]

Plant phenotyping is a key link in understanding gene function and environmental effects, and with development of plant function genomics and crop molecular breeding, the traditional phenotypic observation has become the main bottleneck. High-throughput plant phenome analysis technology and plant phenomics study is an effective way to solve this problem. Although plant phenome analysis is becoming a hot spot at home and abroad, relevant concepts are still relatively fuzzy, and this situation hinders the development of this emerging discipline. In this paper, the relevant concepts and categories of plant phenome and plant phenomics were analyzed, and the new concepts such as quasi-phenome, identifiable traits, mapped traits, and tolerance of plant phenotype to the changes of inheritance and environment, were introduced. And, plant phenome was defined as “all of physical, physiological and biochemical characteristics and traits which are decided or influenced by genome and environments, and can reflect the plant structures and compositions, or reflect the processes and results of plant growth and development”, and plant phenomics as “the comprehensive controls, complete collections and systematic analyses of plant phenome informations and related environmental parameters”. The scopes, directions, and top design principles of plant phenomics research, were also discussed.

PAN Y

. Analysis of concepts and categories of plant phenome and phenomics
Acta Agronomica Sinica, 2015,41(2):175-186. (in Chinese)

DOI:10.3724/SP.J.1006.2015.00175 URL [本文引用: 1]

穆金虎, 陈玉泽, 冯慧, 李文建, 周利斌 . 作物育种学领域新的革命: 高通量的表型组学时代
植物科学学报, 2016,34(6):962-971.

DOI:10.11913/PSJ.2095-0837.2016.60962 URL [本文引用: 1]

传统的表型组学研究已严重滞后于高速发展的基因组、转录组及蛋白质组学，这也制约了作物育种学、功能基因组学等领域研究的深入开展。为突破这一瓶颈，国内外科研工作者经不懈的努力开发出了各类具有自动化、高精度、高通量特点的表型组学分析平台，并将该平台与各类“组学”研究相结合，这将是作物育种学领域的一次新的技术革命。本文对植物表型组学的概念和研究意义进行了介绍和分析，并对高通量表型组学分析平台进行了详细介绍，同时对未来表型组学的发展和各类组学及生物大数据的综合利用进行了展望。

MU J

, CHEN Y

, FENG

, LI W

, ZHOU L

. A new revolution in crop breeding: The era of high-throughput phenomics

Plant Science Journal,

2016,34(6):962-971. (in Chinese)

DOI:10.11913/PSJ.2095-0837.2016.60962 URL [本文引用: 1]