中国农业科学 ›› 2022, Vol. 55 ›› Issue (13): 2613-2628.doi: 10.3864/j.issn.0578-1752.2022.13.011
陈旭1(),郝雅琼1,聂兴华1,杨海莹1,刘松1,王雪峰2,曹庆芹1,秦岭1(
),邢宇1(
)
收稿日期:
2021-10-18
接受日期:
2022-03-01
出版日期:
2022-07-01
发布日期:
2022-07-08
联系方式:
陈旭,Tel:13463609439;E-mail: 13463609439@163.com。
基金资助:
CHEN Xu1(),HAO YaQiong1,NIE XingHua1,YANG HaiYing1,LIU Song1,WANG XueFeng2,CAO QingQin1,QIN Ling1(
),XING Yu1(
)
Received:
2021-10-18
Accepted:
2022-03-01
Published:
2022-07-01
Online:
2022-07-08
摘要:
【目的】通过测定113份板栗品种(系)的数量、质量和假质量性状,分析其遗传变异,比较不同组群之间的性状差异,将SSR标记与性状进行关联,获得更多与SSR标记显著关联的性状,挖掘优异的等位变异位点,为开展板栗分子辅助育种的研究提供参考。【方法】测定并分析板栗总苞和坚果的38个数量、质量和假质量性状,利用SPSS和Graphpad软件对数量性状进行差异显著性和相关性分析,基于SSR标记进行遗传多样性分析,最后用TASSEL 2.1软件通过一般线性模型(general linear model,GLM)和混合线性模型(mixed linear model,MLM)分别对性状和标记进行关联分析。【结果】在遗传多样性分析中,21对SSR引物的有效等位基因数(Ne)、Shannon指数(I)、多态性信息含量(PIC>0.5)、观察杂合度(Ho)和期望杂合度(He)的平均值分别为3.164、1.269、0.589、0.593和0.635。根据群体结构分析分为2个主要组群,为了更好地比较性状差异,在聚类分析时将中间类型的品种单独划分为一组。在质量和假质量性状分析中,多样性指数变化范围为0.139—1.567,遗传多样性最高的性状是坚果光泽,最低的是底座接线;坚果形状、光泽、颜色等性状组群间频率分布差异明显。在数量性状分析中,变异系数的范围在3.96%—36.31%,苞重、总苞重和单粒重的变异系数均在30%以上,遗传变异程度高;果形指数和含水量变异系数均在10%以下,具有稳定的遗传特性;总苞和坚果的外观性状之间有强相关性,相关系数均在0.6以上;组1-组2和组2-组3在果实形状和重量中存在显著差异(P<0.05)。在关联分析中,GLM模型中有13个标记位点与18个表型性状极显著关联,表型变异解释率范围为15.12%—54.99%;MLM模型中有6个标记位点与7个表型性状极显著关联,表型变异解释率范围在8.66%—26.93%。【结论】本研究将SSR标记与表型性状进行关联分析,共发现13个标记位点与坚果单粒重等20个表型性状极显著关联,为开展板栗分子辅助育种的研究奠定了基础。
陈旭,郝雅琼,聂兴华,杨海莹,刘松,王雪峰,曹庆芹,秦岭,邢宇. 板栗总苞和坚果主要性状与SSR标记的关联分析[J]. 中国农业科学, 2022, 55(13): 2613-2628.
CHEN Xu,HAO YaQiong,NIE XingHua,YANG HaiYing,LIU Song,WANG XueFeng,CAO QingQin,QIN Ling,XING Yu. Association Analysis of Main Characteristics of Bur and Nut with SSR Markers in Chinese Chestnut[J]. Scientia Agricultura Sinica, 2022, 55(13): 2613-2628.
表1
SSR分子标记的多态性"
引物 Marker | 观察等位基因数 Na | 有效等位基因数 Ne | Shannon指数 I | 观测杂合度 Ho | 期望杂合度 He | 多态性信息含量 PIC |
---|---|---|---|---|---|---|
CmSI0396 | 4 | 2.553 | 1.069 | 0.708 | 0.608 | 0.542 |
CmSI0509 | 5 | 2.217 | 1.040 | 0.504 | 0.549 | 0.503 |
CmSI0561 | 8 | 1.988 | 1.132 | 0.460 | 0.497 | 0.477 |
CmSI0617 | 8 | 4.414 | 1.637 | 0.814 | 0.773 | 0.738 |
CmSI0658 | 6 | 4.454 | 1.589 | 0.796 | 0.775 | 0.740 |
CmSI0702 | 5 | 2.887 | 1.177 | 0.637 | 0.654 | 0.589 |
CmSI0735 | 6 | 2.753 | 1.199 | 0.646 | 0.637 | 0.568 |
CmSI0742 | 6 | 3.149 | 1.280 | 0.558 | 0.682 | 0.624 |
CmSI0800 | 13 | 5.240 | 1.959 | 0.708 | 0.809 | 0.786 |
CmSI0809 | 4 | 3.015 | 1.193 | 0.625 | 0.668 | 0.604 |
CmSI0853 | 6 | 3.835 | 1.518 | 0.681 | 0.739 | 0.699 |
CmSI0871 | 5 | 1.895 | 0.768 | 0.531 | 0.472 | 0.384 |
CmSI0883 | 6 | 3.776 | 1.531 | 0.681 | 0.735 | 0.702 |
CmSI0933 | 3 | 1.822 | 0.799 | 0.469 | 0.451 | 0.408 |
CmSI0938 | 4 | 1.774 | 0.740 | 0.204 | 0.436 | 0.368 |
CmSI0922 | 8 | 4.884 | 1.766 | 0.664 | 0.795 | 0.769 |
CmSI0930 | 11 | 4.617 | 1.838 | 0.779 | 0.783 | 0.761 |
CmSI0881 | 6 | 3.278 | 1.442 | 0.478 | 0.695 | 0.658 |
CmSI0614 | 6 | 4.464 | 1.590 | 0.805 | 0.776 | 0.740 |
CmSI0404 | 3 | 1.399 | 0.510 | 0.212 | 0.285 | 0.253 |
CmSI0747 | 3 | 2.042 | 0.880 | 0.496 | 0.510 | 0.456 |
均值 Average | 6 | 3.164 | 1.269 | 0.593 | 0.635 | 0.589 |
表2
板栗品种(系)质量性状的频率分布"
性状 Character | 频率分布 Frequency distribution (%) | 多样性指数H' Diversity index | ||||
---|---|---|---|---|---|---|
1 | 2 | 3 | 4 | 5 | ||
开裂方式Bur crack | 32.74 | 63.72 | 3.54 | 0.771 | ||
刺束分支角Prickle angle | 4.42 | 65.49 | 30.09 | 0.777 | ||
刺束密度Prickle density | 5.31 | 29.20 | 65.49 | 0.793 | ||
刺束颜色Prickle color | 11.50 | 88.50 | 0.357 | |||
坚果颜色均匀度Nut color evenness | 71.68 | 22.12 | 6.19 | 0.745 | ||
坚果光泽Nut glossiness | 21.24 | 30.09 | 13.27 | 20.35 | 15.04 | 1.567 |
果顶果肩Apex | 26.55 | 35.40 | 35.40 | 2.65 | 1.184 | |
茸毛分布Fuzz distribution | 56.64 | 22.12 | 21.24 | 0.985 | ||
茸毛颜色Fuzz color | 25.66 | 74.34 | 0.570 | |||
茸毛密度Fuzz density | 30.97 | 25.66 | 28.32 | 15.04 | 1.354 | |
筋线明显程度Obvious of stripes | 76.11 | 20.35 | 3.54 | 0.650 | ||
底座大小Hilum size | 9.73 | 50.44 | 39.82 | 0.939 | ||
底座光滑度Hilum smoothing | 44.25 | 55.75 | 0.687 | |||
涩皮剥离难易程度Difficulty of peeling inner skin | 12.39 | 87.61 | 0.375 | |||
刺苞形状Bur shape | 24.78 | 74.34 | 0.88 | 0.608 | ||
坚果颜色Nut color | 2.65 | 36.28 | 14.16 | 41.59 | 5.31 | 1.262 |
边果形状Nut shape | 77.88 | 22.12 | 0.528 | |||
底座接线Hilum margin | 97.35 | 0.88 | 1.77 | 0.139 | ||
种仁颜色Kernel color | 5.31 | 87.61 | 7.08 | 0.459 |
表3
板栗品种(系)的质量性状频率分布差异"
性状 Character | 组别 Groups | 频率分布 Frequency distribution (%) | ||||
---|---|---|---|---|---|---|
1 | 2 | 3 | 4 | 5 | ||
开裂方式 Bur crack | 1 | 47.06 | 47.06 | 5.88 | ||
2 | 29.33 | 68.00 | 2.67 | |||
3 | 33.33 | 61.90 | 4.76 | |||
刺束分支角 Prickle angle | 1 | 5.88 | 58.82 | 35.29 | ||
2 | 5.33 | 65.33 | 29.33 | |||
3 | 71.43 | 28.57 | ||||
刺束密度 Prickle density | 1 | 5.88 | 23.53 | 70.59 | ||
2 | 5.33 | 32.00 | 62.67 | |||
3 | 4.76 | 23.81 | 71.43 | |||
刺束颜色 Prickle color | 1 | 11.76 | 88.24 | |||
2 | 10.67 | 89.33 | ||||
3 | 14.29 | 85.71 | ||||
坚果颜色均匀度 Nut color evenness | 1 | 70.59 | 23.5 | 5.88 | ||
2 | 76.00 | 17.33 | 6.67 | |||
3 | 57.14 | 38.10 | 4.76 | |||
坚果光泽 Nut glossiness | 1 | 29.41 | 41.18 | 11.76 | 11.76 | 5.88 |
2 | 20.00 | 26.67 | 10.67 | 22.67 | 20.00 | |
3 | 19.05 | 33.33 | 23.81 | 19.05 | 4.76 | |
果顶果肩 Apex | 1 | 17.65 | 29.41 | 52.94 | ||
2 | 28.00 | 36.00 | 33.33 | 2.67 | ||
3 | 28.57 | 38.10 | 28.57 | 4.76 | ||
茸毛分布 Fuzz distribution | 1 | 88.24 | 5.88 | 5.88 | ||
2 | 48.00 | 25.33 | 26.67 | |||
3 | 61.90 | 23.81 | 14.29 | |||
茸毛颜色 Fuzz color | 1 | 23.53 | 76.47 | |||
2 | 26.67 | 73.33 | ||||
3 | 23.81 | 76.19 | ||||
茸毛密度 Fuzz density | 1 | 35.29 | 41.18 | 11.76 | 11.76 | |
2 | 29.33 | 18.67 | 32.00 | 20.00 | ||
3 | 33.33 | 38.10 | 28.57 | |||
筋线明显程度 Obvious of stripes | 1 | 76.47 | 23.53 | |||
2 | 78.67 | 16.00 | 5.33 | |||
3 | 66.67 | 33.33 | ||||
底座大小 Hilum size | 1 | 23.53 | 58.82 | 17.65 | ||
2 | 6.67 | 41.33 | 52.00 | |||
3 | 9.52 | 76.19 | 14.29 | |||
性状 Character | 组别 Groups | 频率分布 Frequency distribution (%) | ||||
1 | 2 | 3 | 4 | 5 | ||
底座光滑度 Hilum smoothing | 1 | 29.41 | 70.59 | |||
2 | 48.00 | 52.00 | ||||
3 | 42.86 | 57.14 | ||||
涩皮剥离难易程度 Difficulty of peeling inner skin | 1 | 23.53 | 76.47 | |||
2 | 9.33 | 90.67 | ||||
3 | 14.29 | 85.71 | ||||
刺苞形状 Bur shape | 1 | 17.65 | 82.35 | |||
2 | 24.00 | 74.67 | 1.33 | |||
3 | 33.33 | 66.67 | ||||
坚果颜色 Nut color | 1 | 17.65 | 70.59 | 11.76 | ||
2 | 2.67 | 42.67 | 18.67 | 33.33 | 2.67 | |
3 | 4.76 | 28.57 | 9.52 | 47.62 | 9.52 | |
边果形状 Nut shape | 1 | 47.06 | 52.94 | |||
2 | 85.33 | 14.67 | ||||
3 | 76.19 | 23.81 | ||||
底座接线 Hilum margin | 1 | 88.24 | 11.76 | |||
2 | 98.67 | 1.33 | ||||
3 | 100.00 | |||||
种仁颜色 Kernel color | 1 | 17.65 | 82.35 | |||
2 | 4.00 | 92.00 | 4.00 | |||
3 | 76.19 | 23.81 |
表4
板栗品种(系)主要数量性状的变异分析"
指标 Character | 极小值 Minimum | 极大值 Maximum | 平均值 Average | 标准差 S | 变异系数 CV (%) |
---|---|---|---|---|---|
总苞重 Total bur weight (g) | 11.93 | 120.83 | 65.91 | 22.02 | 33.41 |
苞重 Bur weight (g) | 7.76 | 79.95 | 38.66 | 14.04 | 36.31 |
刺苞宽Bur width (mm) | 50.80 | 105.31 | 80.54 | 10.77 | 13.38 |
刺苞厚Bur thickness (mm) | 42.58 | 88.06 | 66.45 | 8.14 | 12.25 |
刺苞高Bur height (mm) | 39.55 | 80.09 | 61.99 | 7.87 | 12.69 |
苞肉厚Bur skin thickness (mm) | 1.52 | 5.48 | 3.63 | 0.974 | 26.80 |
刺束长Prickle length (mm) | 6.80 | 18.66 | 13.77 | 2.21 | 16.08 |
刺苞的坚果数Nuts No./Bur | 1.38 | 3.10 | 2.50 | 0.305 | 12.21 |
单粒重Single nut weight (g) | 2.67 | 20.68 | 10.99 | 3.81 | 34.69 |
坚果厚Nut thickness (mm) | 13.86 | 32.05 | 20.60 | 3.25 | 15.76 |
坚果宽Nut width (mm) | 19.42 | 39.53 | 31.08 | 3.67 | 11.79 |
坚果高Nut height (mm) | 17.45 | 32.63 | 26.60 | 2.71 | 10.19 |
果形指数Nut shape index | 0.788 | 0.953 | 0.858 | 0.034 | 3.96 |
出实率Kernel yield (%) | 22.23 | 56.84 | 41.27 | 5.80 | 14.05 |
含水量Water content (%) | 44.46 | 60.69 | 52.03 | 3.57 | 6.85 |
可溶性糖含量Soluble sugar (%, DW) | 8.63 | 19.75 | 13.21 | 2.08 | 15.76 |
直链淀粉含量Amylose (%, DW) | 4.44 | 18.53 | 11.86 | 3.20 | 26.94 |
支链淀粉含量Amylopectin (%, DW) | 28.52 | 75.64 | 53.15 | 10.15 | 19.09 |
总淀粉含量Total starch (%, DW) | 40.22 | 85.18 | 65.01 | 9.39 | 14.45 |
表5
板栗品种(系)的数量性状差异比较"
指标 Character | 组1 Group1 | 组2 Group2 | 组3 Group3 |
---|---|---|---|
总苞重Total bur weight (g) | 56.29±17.47a | 72.26±21.37b | 51.03±16.40a |
苞重Bur weight (g) | 32.80±10.55a | 42.68±13.88b | 29.05±9.71a |
刺苞宽Bur width (mm) | 75.29±10.62a | 83.62±9.81b | 73.78±9.07a |
刺苞厚Bur thickness (mm) | 62.15±7.98a | 68.94±7.12b | 61.05±7.40a |
刺苞高Bur height (mm) | 57.34±7.09a | 64.58±6.58b | 56.54±7.96a |
苞肉厚Bur skin thickness (mm) | 3.30±0.82a | 3.84±0.96b | 3.18±0.89a |
刺束长Prickle length (mm) | 12.97±2.02a | 14.18±2.10a | 12.95±2.31a |
刺苞的坚果数Nuts No./Bur | 2.53±0.28a | 2.49±0.30a | 2.52±0.32a |
单粒重Single nut weight (g) | 9.24±3.05a | 12.03±3.74b | 8.67±2.87a |
坚果厚Nut thickness (mm) | 19.22±2.52a | 21.42±3.12b | 18.75±2.99a |
坚果宽Nut width (mm) | 29.78±3.47a | 32.06±3.34b | 28.63±3.35a |
坚果高Nut height (mm) | 25.63±2.71a | 27.32±2.38b | 24.82±2.71a |
果形指数Nut shape index | 0.86±0.04a | 0.85±0.04a | 0.87±0.02a |
出实率Kernel yield (%) | 41.69±4.50a | 40.71±6.31a | 42.95±4.10a |
含水量Water content (%) | 51.58±3.17a | 52.13±3.56a | 52.04±3.77a |
可溶性糖含量Soluble sugar (%, DW) | 12.62±2.11a | 13.19±1.98a | 13.83±2.19a |
直链淀粉含量Amylose (%, DW) | 12.94±2.93a | 11.77±3.26a | 11.21±2.81a |
支链淀粉含量Amylopectin (%, DW) | 52.51±11.56a | 52.82±9.68a | 55.09±9.96a |
总淀粉含量Total starch (%, DW) | 65.45±10.82a | 64.58±9.02a | 66.29±8.96a |
表6
GLM和MLM模型的关联分析结果"
性状 Character | 位点 Loci | GLM | MLM | ||
---|---|---|---|---|---|
P<0.01 | R2 (%) | P<0.01 | R2 (%) | ||
总苞重Total bur weight | CmSI0883 | 0.0017 | 36.84 | ||
CmSI0930 | 0.0021 | 46.90 | |||
苞重Bur weight | CmSI0883 | 0.0023 | 36.43 | ||
CmSI0930 | 0.001 | 49.07 | 0.0095 | 16.31 | |
刺苞宽Bur width | CmSI0883 | 0.0072 | 33.37 | ||
刺苞厚Bur thickness | CmSI0883 | 0.0045 | 33.90 | ||
刺苞高Bur height | CmSI0883 | 7.96E-04 | 37.47 | ||
刺束长Prickle length | CmSI0702 | 0.0018 | 28.87 | ||
CmSI0883 | 0.0084 | 34.38 | |||
刺苞的坚果数Nuts No./Bur | CmSI0800 | 0.0074 | 53.33 | 0.0027 | 19.99 |
单粒重Single nut weight | CmSI0853 | 0.0021 | 38.84 | ||
CmSI0930 | 0.0092 | 42.86 | |||
坚果宽Nut width | CmSI0930 | 0.0065 | 45.21 | ||
坚果高Nut height | CmSI0883 | 0.0035 | 36.59 | ||
含水量Water content | CmSI0735 | 6.71E-04 | 36.42 | ||
可溶性糖含量Soluble sugar | CmSI0800 | 0.0085 | 26.93 | ||
刺束密度Prickle density | CmSI0735 | 0.0064 | 26.79 | 0.0065 | 8.66 |
CmSI0742 | 0.0053 | 29.04 | |||
坚果颜色均匀度Nut color evenness | CmSI0871 | 8.14E-05 | 30.05 | ||
筋线明显程度Obvious of stripes | CmSI0800 | 0.0037 | 20.21 | ||
涩皮剥离难易程度Difficulty of peeling inner skin | CmSI0561 | 7.94E-05 | 42.52 | ||
刺苞形状Bur shape | CmSI0404 | 0.0061 | 15.12 | ||
坚果颜色Nut color | CmSI0881 | 0.0076 | 36.49 | ||
底座接线Hilum margin | CmSI0509 | 1.73E-06 | 45.52 | 1.20E-03 | 14.41 |
CmSI0881 | 5.96E-07 | 54.99 | 3.60E-03 | 16.29 | |
种仁颜色Kernel color | CmSI0617 | 0.0037 | 47.72 | 0.0041 | 17.93 |
CmSI0881 | 0.0033 | 37.95 | 0.0013 | 15.01 |
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