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核桃(Juglans regia L.)与泡核桃(J. sigillata Dode)均属胡桃科(Juglandaceae),核桃属(Juglans L.),核桃组(Section Juglans),在我国四川省凉山彝族自治州均有栽培和分布[1]。凉山彝族自治州位于四川省西南部,该地区东接四川盆地,西跨横断山系,北抵川西高原,南临金沙江畔,处于青藏高原和云贵高原与四川盆地的过渡地带;由于独特的地质演化,形成以高山深谷为主,并有平坝、河谷和丘陵相间的特殊地貌,所以形成了最为显著的山地垂直地带性气候,加之是光热资源丰富的低纬度地区,是生物多样性极其丰富的地区[2-4]。凉山州的核桃和泡核桃资源存量丰富,区内德昌、会理、雷波、木里、冕宁、盐源等县是传统产地,并存在一定规模的天然居群和数量可观的实生类型群体[5-6],是核桃和泡核桃宝贵的遗传资源和潜在的种质库。之前对凉山州核桃表型性状的研究较少,已有报道涉及的采样地或资源数量有限,未开展广泛调查及进一步对坚果性状的详细研究。近年来,凉山州开始大力发展核桃产业,首先面临的问题是对现有资源的调查,在摸清本地区核桃和泡核桃资源现状的基础上,方能提出有效的发展规划,最直接和有效的方法是从坚果的主要表型性状及其特色入手。因此,本文以凉山州15个实生居群330棵单株的坚果为试材,利用18个坚果表型性状揭示该地区核桃和泡核桃种质资源的多样性,明确农家实生资源的丰富程度,以期为制定该地区核桃和泡核桃资源保存与利用策略及构建核心种质提供参考。
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根据当地林业部门的相关资料,在核桃和泡核桃资源相对集中的区域,走访当地群众获取树龄、果实特异性等相关信息,选择树龄30 a以上的实生核桃和泡核桃树采集试验样品。每个样本采集30个大小基本一致、饱满的坚果带回实验室,晾干至恒质量后进行表型性状的测定。本研究的试材采集范围包括凉山州15个县(市),15个核桃和泡核桃实生居群名称和采集地生态、地理信息及样本数见表 1。
表 1 采样地生态、地理信息及样本数
Table 1. The ecological and geographical information and sample size of populations in this study
居群代码
Code采样地
sampling site经度(E)
Langitude纬度(N)
Latitude年降水量
Annual precipitation/mm年均气温
Annual temperature/℃年日照时数
Annual sunshine hours/h海拔Elevation 样本数
Sample size/个范围
Range/m平均值
Mean/mXC 西昌 102°05′09″ 28°04′07″ 1 013 15.0 1 648 1 500~2 500 1 542 9 YY 盐源 101°29′35″ 27°25′53″ 855 12.1 2 600 1 800~2 700 2 540 8 DC 德昌 102°10′37″ 27°24′21″ 1 049 17.7 2 147 1 300~2 500 1 380 38 HL 会理 102°14′33″ 26°39′16″ 1 130 15.1 2 055 1 000~2 600 1 788 46 HD 会东 102°34′24″ 26°38′08″ 1 001 16.1 2 445 1 100~2 300 2 075 24 NN 宁南 102°44′00″ 27°03′48″ 970 19.4 2 257 800~1 900 1 643 17 PG 普格 102°32′22″ 27°22′35″ 1 300 16.8 2 095 1 040~2 680 1 860 12 JY 金阳 103°15′08″ 27°41′33″ 820 15.8 1 574 900~2 200 1 452 13 ZJ 昭觉 102°50′58″ 28°00′28″ 1 013 19.0 1 622 1 260~1 600 1 430 15 MN 冕宁 102°13′48″ 28°32′52″ 1 110 13.3 1 878 1 700~2 500 1 774 14 YX 越西 102°30′31″ 28°38′06″ 1 133 13.3 1 687 1 620~2 700 1 662 12 GL 甘洛 102°47′48″ 29°02′25″ 850 17.0 1 661 870~1 980 1 170 12 MG 美姑 103°06′17″ 28°26′23″ 815 11.2 1 791 1 938~2 211 2 074 33 LB 雷波 103°34′19″ 28°15′45″ 850 12.2 1 225 900~1 800 1 353 43 ML 木里 101°16′18″ 27°56′09″ 800 14.3 2 303 2 000~2 800 2 667 34 合计Tota l330 -
坚果表型性状测定参照《植物新品种特异性、一致性和稳定性测试指南-核桃属》[7]进行特征描述和赋值,调查测定了核桃和泡核桃的18个坚果表型性状,测试指南中未列出的特征用“其他”表示[8](表 2)。坚果数量性状中的纵径、横径、侧径用游标卡尺(精度0.01 mm)测量,单果质量、核仁质量用天平(精度0.01 g)测量,果壳厚度用螺旋测微仪(精度0.001 mm)测量。3个数量性状的计算公式为:
$ 坚果圆度指数=(侧径+横径)/(2×纵径)\\ 出仁率=核仁质量/单果质量×100\%\\ 三径均值=(纵径+侧径+横径)/3 $
表 2 凉山州核桃和泡核桃坚果表型特征赋值及描述
Table 2. The description and assignment of nut quality character for walnut accessions in Liangshan prefecture
代码
Code坚果表型质量性状
Nut quality traits赋值及描述
Description and assignment1 沿缝合线纵切面形状
Shape in longitudinalsection through suture1:椭圆形Elliptic;2:阔椭圆形Broad elliptic;3:长圆形Long circular;4:圆形Circular;
5:卵圆形Ovate;6:阔卵形Broad ovate;7:三角形Triangular;8:梯形Trapezium2 垂直缝合线纵切面形状
Shape in longitudinal section perpendicular to suture1:圆形Elliptic;2:扁圆形Oblate;3:倒卵圆形Inverted Ovate;4:心形Heart-shaped;
5:阔卵圆形Broad ovate;6:三角形Triangular;7:方(梯)形Squareness(Trapezium);
8:长椭圆形Long circular;9:卵圆形Ovate;10:其他Other (图 1)3 坚果横切面形状Shape in cross section 2:扁圆形Oblate;3:圆形Circular;4:椭圆形Elliptic;5:其他Other 4 顶尖突出程度Prominence of apical tip 3:平Weak flat;5:凸Medium bulge;7:尖Strong pointed 5 果肩形状Shape of apex perpendicular to suture 1:圆形Rounded;2:平Truncate 6 果底形状Shape of base perpendicular to suture 1:圆形Rounded;2:扁圆形Oblate;3:楔形Cuneate;4:方形Truncate;5:其他Other 7 刻窝Pit on the shell 1:少Slightly;3:中Moderately;5:多Strongly;7:很多Embossed; 8 刻纹Groove on the shell 1:平Flat;3:浅Shallow;5:中Medium;7:深Deep 9 缝合线条数Number of pads on suture 1:2条Two;2:2~3条Two-three;3:3条Three;4:2~4条Two-four;5:3~4条Three-four;6:4条Four (图 2) 10 缝合线突出程度Prominence of pad on suture 3:平Flat;5:凸Bulge 11 横隔膜质地Diaphragm texture 1:膜质Membranous;2:纸质Papery;3:革质Leathery;4:骨质Bony 12 内褶壁质地Inner pleat wall of shell 1:膜质Membranous;2:革质Leathery;3:骨质Bony;4:退化Degeneration 13 核仁皮色Skin color of kernel 1:白(黄白)White (Yellow white);2:浅黄Light yellow;3:黄Yellow;4:浅褐Light brown;5:褐Brown;6:红褐(紫红)Red brown (Purple red);7:紫褐(紫)Purple brown (Purple);8:乌Black (图 3) 图 1 凉山州核桃和泡核桃坚果垂直于缝合线纵切面形状
Figure 1. Shape in longitudinal section perpendicular to suture of walnut in Liangshan prefecture
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运用Microsoft Excel 2007,SPSS19.0[9]进行数据统计分析,包括分布频率、Simpson指数和Shannon-Wiener指数,并绘制数量性状频率数据分布直方图[10],用SAS 8.1[11]软件统计坚果表型性状的平均值和标准差,用类平均法计算居群间的Manhattan距离并进行聚类分析。用性状的变异系数(CV)表示居群表型多样性水平[12],并对其做Duncan检验;对各性状观测值采用巢式设计方差分析比较居群间和居群内的差异显著性[13];用表型分化系数(VST)反应居群间表型分化程度[14]。
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由表 3可知:13个坚果表型的质量性状Simpson指数为0.201~0.855,Shannon-Wiener指数为0.649~2.873,其中,沿缝合线纵切面形状(代码1)的分别为0.855和2.873,垂直缝合线纵切面(代码2)的分别为0.771和2.546,刻窝数量(代码7)的分别为0.731和1.940,核仁皮色(代码13)的分别为0.761和2.490,坚果缝合线条数(代码9)和缝合线突出程度(代码10)的多样性指数较低。在数据采集和分析中发现倒卵圆形、三角形和方(梯)形3种垂直缝合线纵切面形状和乌仁(一种仁色特征),这在核桃属DUS测试指南中未见描述,另有部分资源描述暂列为其他,因此,将垂直缝合线纵切面形状分为10级,核仁皮色分为8级(表 2)。
表 3 凉山州330份样本13个坚果表型的质量性状变异状况
Table 3. Variation of thirteen quality traits for 330 genotypes nut phenotypic in Liangshan prefecture
质量性状代码
Quality traits code各级所占比例The proportion of each level/% 辛普森指数
Simpson index香农-维纳指数
Shannon-Wiener index1 2 3 4 5 6 7 8 9 10 1 15.15 10.91 13.94 21.52 3.33 8.18 13.94 13.03 0.855 2.873 2 38.79 16.97 2.42 3.03 3.33 1.21 2.12 12.12 17.58 2.42 0.771 2.546 3 39.39 58.48 1.52 0.61 0.502 1.118 4 34.85 47.88 17.27 0.619 1.476 5 58.48 41.52 0.486 0.979 6 46.06 9.09 14.24 29.09 1.52 0.674 1.840 7 14.24 30.91 31.21 23.64 0.731 1.940 8 45.15 40.61 9.39 4.00 0.620 1.578 9 89.09 6.67 0.30 3.33 0.30 0.30 0.201 0.649 10 26.97 73.03 0.394 0.841 11 13.33 46.97 28.48 11.21 0.668 1.770 12 13.94 24.85 6.97 54.24 0.620 1.642 13 10.00 17.88 41.82 3.64 3.64 5.45 6.97 10.61 0.761 2.490 表 2、3表明:沿缝合线纵切面形状分为8种,其中,圆形较多(21.52%),卵圆形最少(3.33%),其它6种形状占比为8.18%~15.15%;垂直缝合线纵切面形状中,圆形所占比例较大(38.79%),其次为卵圆形(17.58%)、扁圆形(16.97%)、长椭圆形(12.12%),其余阔卵圆形、心形、倒卵圆形、三角形和方(梯)形所占比例较小,为3.33%~1.21%;坚果横切面形状以圆形和扁圆形为主;果肩形状和果底形状各级占比均以圆形居多,分别占58.48%和46.06%;顶尖突出程度中,凸的资源最多,占47.88%,进一步印证了顶尖突出程度明显的坚果在该地区是主要类型。本研究将缝合线条数分为6级,缝合线条数以调查结果所占比例超过样本总数的一半为准,其中缝合线条数2条的占比较多,为89.09%,2条以上的占比较低,为6.67%~0.30%。刻窝各级占比为多和很多的各占31.21%和23.64%,说明所调查地区泡核桃资源的坚果表型以深刻窝为主。横隔膜质地以纸质的资源最多,占46.97%。内褶壁质地退化的资源最多,占54.24%,说明该地区大部分薄壳核桃和泡核桃较易取仁。该地区核仁皮色仍以黄色为主,占41.82%,除其它6种核仁皮色,还有10.61%的乌仁资源。
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表 4表明:单果质量为3.29~28.18 g,平均11.99 g;壳厚为0.11~4.50 mm,平均0.96 mm;出仁率为10.00%~78.48%,平均48.06%;圆度指数为0.66~1.13,平均0.88;三径均值为21.24~47.41 mm,平均34.17 mm。坚果核壳厚度的变异系数最大,为44.59%;其次为单果质量,为31.02%;再次是出仁率,为19.03%;最后是三径均值和圆度指数,分别为10.41%和9.22%。
表 4 凉山州330份样本坚果表型5个数量性状的变异状况
Table 4. Variation of five quantitative traits for 330 genotypes nut phenotypic in Liangshan prefecture
数量性状代码
Quantitative traits Code数量性状
Quantitative traits最大值
Max.最小值
Min.平均值
Mean极差
Range标准差
SD变异系数
CV/%14 单果质量Weight of nut/g 28.18 3.29 11.99 24.89 3.72 31.02 15 壳厚Shell thickness/mm 4.50 0.11 0.96 4.39 0.43 44.59 16 出仁率Kernel rate/% 78.48 10.00 48.06 68.48 9.14 19.03 17 圆度指数Index of roundness 1.13 0.66 0.88 0.47 0.08 9.22 18 三径均值Geometric mean diameter/mm 47.41 21.24 34.17 24.17 3.56 10.41 各坚果表型数量性状频率分布直方图(图 4)表明:各性状的分布存在差异,单果质量、出仁率和圆度指数均符合正态分布,但核壳厚度和三径均值近似服从χ2分布。单果质量大多为10.04~13.94 g,占46.67%。出仁率集中在43.26%~52.86%,占51.52%。三径均值在32.31~38.73 mm集中分布,但32.31~36.04 mm的性状频率分布最大,占46.97%。圆度指数较分散,其中,0.78~0.84的占23.33%,0.84~0.92的占35.76%,0.92~0.98的占21.52%。坚果核壳厚度0.74~1.19 mm的性状频率分布最大,占55.76 %。
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由表 5可知:18个坚果表型性状变异系数为8.46%~59.47%,差异显著(P<0.05),平均为35.26%,其中,变异程度最大的是刻纹,为59.47%;变异程度最小的是出仁率,为8.46%。15个居群内18个坚果表型性状变异系数平均值为29.49%~39.28%,居群间差异不显著。
表 5 凉山州15个实生居群坚果表型性状变异系数及多重比较
Table 5. Variation coefficients (CV) and multiple comparisons of phenotypic traits in 15 populations in Liangshan prefecture
表型性状代码
Code of phenotypic traits居群Population 均值
Mean/%DC GL HD HL JY LB MG ML MN NN PG XC YX YY ZJ 1 63.21 36.47 63.56 63.55 66.89 47.24 54.99 37.17 46.04 43.86 58.86 69.02 52.26 37.64 49.13 52.66g 2 31.13 25.69 36.77 38.53 54.20 54.20 29.13 44.78 22.85 40.05 28.75 39.09 42.74 32.63 35.77 37.09def 3 26.08 20.89 18.84 20.59 15.84 18.83 17.65 17.09 18.81 20.82 21.10 21.43 22.60 21.38 21.82 20.25b 4 29.41 30.06 36.12 22.87 31.35 34.07 30.97 33.60 30.59 29.33 25.88 29.27 19.93 39.40 31.40 30.28cd 5 45.20 34.82 30.43 35.49 45.17 38.10 35.48 36.83 35.95 34.99 33.36 36.08 34.82 26.45 36.22 35.96cde 6 57.76 60.54 58.23 55.16 60.46 54.40 60.60 68.91 63.60 60.91 60.80 49.93 44.20 46.29 61.93 57.58g 7 44.91 38.56 60.53 47.22 75.31 48.54 39.66 45.05 39.05 27.62 51.31 30.57 30.76 50.65 43.04 44.85f 8 69.00 49.49 86.92 60.20 65.81 64.11 59.84 52.05 60.31 59.46 67.42 68.82 40.20 46.61 41.82 59.47g 9 82.22 0.00 37.68 44.70 25.75 54.55 31.91 68.29 24.94 0.00 26.65 70.19 33.36 77.14 78.49 43.73ef 10 20.79 22.72 16.32 20.53 11.45 15.81 10.09 20.97 25.94 19.31 26.11 27.11 26.86 27.60 14.87 20.43b 11 36.51 34.88 27.24 29.93 35.36 36.89 32.68 30.28 30.94 29.22 37.26 39.67 55.16 61.72 26.35 36.27cde 12 40.11 44.72 31.65 40.37 48.86 21.46 32.30 34.09 37.15 40.70 51.31 43.27 69.28 70.71 29.41 42.36ef 13 47.83 70.20 51.94 64.46 65.56 45.61 57.29 68.67 38.12 52.70 40.18 55.22 63.21 49.68 57.47 55.21g 14 37.51 36.71 24.67 21.48 31.82 27.99 25.59 34.58 33.64 17.31 22.11 21.66 18.50 37.60 34.33 28.37c 15 32.76 36.26 34.91 19.31 38.83 43.80 72.44 42.27 25.33 30.32 36.12 23.26 25.43 35.54 35.15 35.45cde 16 10.57 8.06 6.92 9.33 5.20 10.65 9.38 8.42 6.17 7.33 11.54 6.19 7.94 10.14 9.02 8.46a 17 15.29 16.07 15.26 16.56 18.91 20.35 29.95 24.58 12.70 10.87 17.60 8.10 11.60 14.11 14.27 16.41ab 18 10.16 12.71 10.27 7.97 10.31 11.08 8.74 11.35 14.91 5.98 4.77 8.62 5.39 15.16 9.46 9.79a 均值Mean 38.92 32.16 36.01 34.35 39.28 35.98 35.48 37.72 31.50 29.49 34.51 35.97 33.57 38.91 35.00 35.26 注:居群代码见表 1,表型性状代码1~18见表 2,下同。“均值”列不同字母表示性状间在0.05水平上差异显著。
Note:Code of population see table 1, codes of phenotypic traits 1-18 see table 2, the same below. Paired populations with the different letters are significantly different (P<0.05) shown in ‘Mean’ line,and Paired traits with the different letters are significantly different (P<0.05) shown in ‘Mean’ column.对15个核桃和泡核桃实生居群的18个表型性状进行巢式方差分析,计算出各分量占变异的比例。由表 6可知:居群内方差分量占总变异的93.97%,居群间方差分量占6.03%;居群间18个表型性状的表型分化系数为0.16%~11.03%,平均为6.03%,18个表型性状分化最大的是刻纹,最小的是果底形状;18个性状方差分量百分比的居群内变异均大于居群间,说明居群内变异是该区域坚果表型变异的主要来源。
表 6 凉山州核桃坚果表型方差分量和表型分化系数
Table 6. Variance components and phenotypic differentiation coefficients of nut phenotypic traits of walnut populations in Liangshan prefecture
表型性状代码
Code of phenotypic traits方差分量Variance component 方差分量百分比Percentage of variance component /% 居群间表型分化系数
VST /%居群间
Among populations居群内
Within populations居群间
Among populations居群内
Within populations1 3.63×10-1 5.22 6.50 93.50 6.50 2 6.03×10-2 11.64 0.52 99.48 0.52 3 1.56×10-2 2.86×10-1 5.19 94.81 5.19 4 1.68×10-2 1.98 0.84 99.16 0.84 5 1.84×10-3 2.81×10-1 0.65 99.35 0.65 6 2.91×10-3 1.82 0.16 99.84 0.16 7 2.97×10-1 3.69 7.46 92.54 7.46 8 3.00×10-1 2.42 11.03 88.97 11.03 9 6.05×10-3 4.70×10-1 1.27 98.73 1.27 10 7.35×10-2 7.23×10-1 9.23 90.77 9.23 11 6.61×10-2 6.67×10-1 9.01 90.99 9.01 12 1.37×10-1 1.23 10.07 89.93 10.07 13 4.06×10-1 4.17 8.87 91.13 8.87 14 1.47 12.49 10.53 89.47 10.53 15 1.46×10-2 1.72×10-1 7.81 92.19 7.81 16 3.09×10-4 6.30×10-3 4.68 95.32 4.68 17 5.73 78.38 6.81 93.19 6.81 18 1.01 11.72 7.92 92.08 7.92 均值Mean 5.54×10-1 7.63 6.03 93.97 6.03 -
以所测的18个坚果表型性状的平均值为参数,利用核桃和泡核桃实生居群表型性状Manhattan距离进行UPGMA聚类分析,结果(图 5)表明:当遗传距离为0.8时,15个核桃和泡核桃实生居群聚为7类。DC、HL、XC、PG聚为安宁河谷类;JY、LB、MG、ZJ聚为大凉山类;GL、NN、MN聚为一类;HD单独一类,在遗传距离为0.9时与大凉山类聚为一类;YX、YY在遗传距离为1.1时聚为一类;ML与其它居群的遗传距离较远。聚类分析结果呈现了与地理距离或立地条件显著正相关的趋势。
凉山州核桃坚果表型多样性研究
Phenotypic Diversities of Nuts of Walnut Populations Originated from Seedlings in Liangshan Prefecture
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摘要:
目的 以四川省凉山州核桃和泡核桃农家类型的坚果为样本,研究其表型丰富度和变异特点,为其资源挖掘和合理利用提供理论依据。 方法 以15个核桃和泡核桃居群的330棵实生单株为研究材料,利用18个坚果表型相关性状进行多样性和聚类分析。 结果 表明:(1)凉山州核桃和泡核桃实生居群坚果表型性状变异系数为8.46%~59.47%,平均为35.26%,单果质量极大(>20.0 g)和极小(<5.0 g)的资源均占一定比例;该地区坚果资源在《植物新品种特异性、一致性和稳定性测试指南-核桃属》中所描述坚果表型性状的基础上,新增了3个垂直于缝合线纵切面形状和1个核仁皮色特征;居群遗传性状Simpson指数为0.201~0.855,Shannon-Wiener指数为0.649~2.873,说明凉山州核桃和泡核桃坚果表型多样性比较丰富。(2)18个坚果表型性状的居群内变异均大于居群间,居群间的表型分化系数为6.03%,说明居群内变异是凉山州坚果表型多样性的主要来源。(3)UPGMA聚类分析结果表明,凉山州坚果表型性状Manhattan距离表现出与地理距离或气候条件显著正相关的趋势。 结论 凉山州核桃和泡核桃资源坚果表型丰富度高,是重要的核桃资源多样性分布地区。 Abstract:Objective Using the nut of Juglans regia and Juglans sigillata in Liangshan of Sichuan as samples to study the phenotype abundance and variation characteristics for detecting resource and providing theory evidence in using resource reasonable. Method Taking 330 seedlings from 15 J. regia and J. sigillata populations as research materials, the diversity and cluster analysis were studied using 18 nut phenotypic traits. Result (1) The coefficient of variation of phenotypic traits in Liangshan walnut population originated from seedlings was between 8.46% and 59.47%, with an average of 35.26%. The nuts with very large (> 20.0 g) and very small (< 5.0 g) single nut weight occupied a certain proportion. The phenotypic traits meet almost all the trait descriptions in the "Guidelines for the conduct of tests for distinctness, uniformity and stability-Juglans (Juglans L.)". Three features in shape and one in skin color of kernels were added. The Simpson index of population genetic traits was 0.201-0.855, Shannon-Wiener index was between 0.649 and 2.873, indicating abundant variations in nut phenotypic diversity in this area. (2) The phenotypic differentiation of 18 phenotypic traits in populations was larger than that among populations in this area and the differentiation coefficients among populations was 6.03%, suggesting the phenotypic differentiation in populations was the main variation source. (3) UPGMA cluster analysis showed that the Manhattan distance of nut phenotypic traits were positively related to the geographic distance or climatic conditions. Conclusion The nut phenotype of J. regia and J. sigillata in Liangshan was abundant. It is the important area for distribution of diversity walnut resources in China. -
Key words:
- walnut(Juglans regia)
- / phenotype diversity
- / liangshan
- / population
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表 1 采样地生态、地理信息及样本数
Table 1. The ecological and geographical information and sample size of populations in this study
居群代码
Code采样地
sampling site经度(E)
Langitude纬度(N)
Latitude年降水量
Annual precipitation/mm年均气温
Annual temperature/℃年日照时数
Annual sunshine hours/h海拔Elevation 样本数
Sample size/个范围
Range/m平均值
Mean/mXC 西昌 102°05′09″ 28°04′07″ 1 013 15.0 1 648 1 500~2 500 1 542 9 YY 盐源 101°29′35″ 27°25′53″ 855 12.1 2 600 1 800~2 700 2 540 8 DC 德昌 102°10′37″ 27°24′21″ 1 049 17.7 2 147 1 300~2 500 1 380 38 HL 会理 102°14′33″ 26°39′16″ 1 130 15.1 2 055 1 000~2 600 1 788 46 HD 会东 102°34′24″ 26°38′08″ 1 001 16.1 2 445 1 100~2 300 2 075 24 NN 宁南 102°44′00″ 27°03′48″ 970 19.4 2 257 800~1 900 1 643 17 PG 普格 102°32′22″ 27°22′35″ 1 300 16.8 2 095 1 040~2 680 1 860 12 JY 金阳 103°15′08″ 27°41′33″ 820 15.8 1 574 900~2 200 1 452 13 ZJ 昭觉 102°50′58″ 28°00′28″ 1 013 19.0 1 622 1 260~1 600 1 430 15 MN 冕宁 102°13′48″ 28°32′52″ 1 110 13.3 1 878 1 700~2 500 1 774 14 YX 越西 102°30′31″ 28°38′06″ 1 133 13.3 1 687 1 620~2 700 1 662 12 GL 甘洛 102°47′48″ 29°02′25″ 850 17.0 1 661 870~1 980 1 170 12 MG 美姑 103°06′17″ 28°26′23″ 815 11.2 1 791 1 938~2 211 2 074 33 LB 雷波 103°34′19″ 28°15′45″ 850 12.2 1 225 900~1 800 1 353 43 ML 木里 101°16′18″ 27°56′09″ 800 14.3 2 303 2 000~2 800 2 667 34 合计Tota l330 表 2 凉山州核桃和泡核桃坚果表型特征赋值及描述
Table 2. The description and assignment of nut quality character for walnut accessions in Liangshan prefecture
代码
Code坚果表型质量性状
Nut quality traits赋值及描述
Description and assignment1 沿缝合线纵切面形状
Shape in longitudinalsection through suture1:椭圆形Elliptic;2:阔椭圆形Broad elliptic;3:长圆形Long circular;4:圆形Circular;
5:卵圆形Ovate;6:阔卵形Broad ovate;7:三角形Triangular;8:梯形Trapezium2 垂直缝合线纵切面形状
Shape in longitudinal section perpendicular to suture1:圆形Elliptic;2:扁圆形Oblate;3:倒卵圆形Inverted Ovate;4:心形Heart-shaped;
5:阔卵圆形Broad ovate;6:三角形Triangular;7:方(梯)形Squareness(Trapezium);
8:长椭圆形Long circular;9:卵圆形Ovate;10:其他Other (图 1)3 坚果横切面形状Shape in cross section 2:扁圆形Oblate;3:圆形Circular;4:椭圆形Elliptic;5:其他Other 4 顶尖突出程度Prominence of apical tip 3:平Weak flat;5:凸Medium bulge;7:尖Strong pointed 5 果肩形状Shape of apex perpendicular to suture 1:圆形Rounded;2:平Truncate 6 果底形状Shape of base perpendicular to suture 1:圆形Rounded;2:扁圆形Oblate;3:楔形Cuneate;4:方形Truncate;5:其他Other 7 刻窝Pit on the shell 1:少Slightly;3:中Moderately;5:多Strongly;7:很多Embossed; 8 刻纹Groove on the shell 1:平Flat;3:浅Shallow;5:中Medium;7:深Deep 9 缝合线条数Number of pads on suture 1:2条Two;2:2~3条Two-three;3:3条Three;4:2~4条Two-four;5:3~4条Three-four;6:4条Four (图 2) 10 缝合线突出程度Prominence of pad on suture 3:平Flat;5:凸Bulge 11 横隔膜质地Diaphragm texture 1:膜质Membranous;2:纸质Papery;3:革质Leathery;4:骨质Bony 12 内褶壁质地Inner pleat wall of shell 1:膜质Membranous;2:革质Leathery;3:骨质Bony;4:退化Degeneration 13 核仁皮色Skin color of kernel 1:白(黄白)White (Yellow white);2:浅黄Light yellow;3:黄Yellow;4:浅褐Light brown;5:褐Brown;6:红褐(紫红)Red brown (Purple red);7:紫褐(紫)Purple brown (Purple);8:乌Black (图 3) 表 3 凉山州330份样本13个坚果表型的质量性状变异状况
Table 3. Variation of thirteen quality traits for 330 genotypes nut phenotypic in Liangshan prefecture
质量性状代码
Quality traits code各级所占比例The proportion of each level/% 辛普森指数
Simpson index香农-维纳指数
Shannon-Wiener index1 2 3 4 5 6 7 8 9 10 1 15.15 10.91 13.94 21.52 3.33 8.18 13.94 13.03 0.855 2.873 2 38.79 16.97 2.42 3.03 3.33 1.21 2.12 12.12 17.58 2.42 0.771 2.546 3 39.39 58.48 1.52 0.61 0.502 1.118 4 34.85 47.88 17.27 0.619 1.476 5 58.48 41.52 0.486 0.979 6 46.06 9.09 14.24 29.09 1.52 0.674 1.840 7 14.24 30.91 31.21 23.64 0.731 1.940 8 45.15 40.61 9.39 4.00 0.620 1.578 9 89.09 6.67 0.30 3.33 0.30 0.30 0.201 0.649 10 26.97 73.03 0.394 0.841 11 13.33 46.97 28.48 11.21 0.668 1.770 12 13.94 24.85 6.97 54.24 0.620 1.642 13 10.00 17.88 41.82 3.64 3.64 5.45 6.97 10.61 0.761 2.490 表 4 凉山州330份样本坚果表型5个数量性状的变异状况
Table 4. Variation of five quantitative traits for 330 genotypes nut phenotypic in Liangshan prefecture
数量性状代码
Quantitative traits Code数量性状
Quantitative traits最大值
Max.最小值
Min.平均值
Mean极差
Range标准差
SD变异系数
CV/%14 单果质量Weight of nut/g 28.18 3.29 11.99 24.89 3.72 31.02 15 壳厚Shell thickness/mm 4.50 0.11 0.96 4.39 0.43 44.59 16 出仁率Kernel rate/% 78.48 10.00 48.06 68.48 9.14 19.03 17 圆度指数Index of roundness 1.13 0.66 0.88 0.47 0.08 9.22 18 三径均值Geometric mean diameter/mm 47.41 21.24 34.17 24.17 3.56 10.41 表 5 凉山州15个实生居群坚果表型性状变异系数及多重比较
Table 5. Variation coefficients (CV) and multiple comparisons of phenotypic traits in 15 populations in Liangshan prefecture
表型性状代码
Code of phenotypic traits居群Population 均值
Mean/%DC GL HD HL JY LB MG ML MN NN PG XC YX YY ZJ 1 63.21 36.47 63.56 63.55 66.89 47.24 54.99 37.17 46.04 43.86 58.86 69.02 52.26 37.64 49.13 52.66g 2 31.13 25.69 36.77 38.53 54.20 54.20 29.13 44.78 22.85 40.05 28.75 39.09 42.74 32.63 35.77 37.09def 3 26.08 20.89 18.84 20.59 15.84 18.83 17.65 17.09 18.81 20.82 21.10 21.43 22.60 21.38 21.82 20.25b 4 29.41 30.06 36.12 22.87 31.35 34.07 30.97 33.60 30.59 29.33 25.88 29.27 19.93 39.40 31.40 30.28cd 5 45.20 34.82 30.43 35.49 45.17 38.10 35.48 36.83 35.95 34.99 33.36 36.08 34.82 26.45 36.22 35.96cde 6 57.76 60.54 58.23 55.16 60.46 54.40 60.60 68.91 63.60 60.91 60.80 49.93 44.20 46.29 61.93 57.58g 7 44.91 38.56 60.53 47.22 75.31 48.54 39.66 45.05 39.05 27.62 51.31 30.57 30.76 50.65 43.04 44.85f 8 69.00 49.49 86.92 60.20 65.81 64.11 59.84 52.05 60.31 59.46 67.42 68.82 40.20 46.61 41.82 59.47g 9 82.22 0.00 37.68 44.70 25.75 54.55 31.91 68.29 24.94 0.00 26.65 70.19 33.36 77.14 78.49 43.73ef 10 20.79 22.72 16.32 20.53 11.45 15.81 10.09 20.97 25.94 19.31 26.11 27.11 26.86 27.60 14.87 20.43b 11 36.51 34.88 27.24 29.93 35.36 36.89 32.68 30.28 30.94 29.22 37.26 39.67 55.16 61.72 26.35 36.27cde 12 40.11 44.72 31.65 40.37 48.86 21.46 32.30 34.09 37.15 40.70 51.31 43.27 69.28 70.71 29.41 42.36ef 13 47.83 70.20 51.94 64.46 65.56 45.61 57.29 68.67 38.12 52.70 40.18 55.22 63.21 49.68 57.47 55.21g 14 37.51 36.71 24.67 21.48 31.82 27.99 25.59 34.58 33.64 17.31 22.11 21.66 18.50 37.60 34.33 28.37c 15 32.76 36.26 34.91 19.31 38.83 43.80 72.44 42.27 25.33 30.32 36.12 23.26 25.43 35.54 35.15 35.45cde 16 10.57 8.06 6.92 9.33 5.20 10.65 9.38 8.42 6.17 7.33 11.54 6.19 7.94 10.14 9.02 8.46a 17 15.29 16.07 15.26 16.56 18.91 20.35 29.95 24.58 12.70 10.87 17.60 8.10 11.60 14.11 14.27 16.41ab 18 10.16 12.71 10.27 7.97 10.31 11.08 8.74 11.35 14.91 5.98 4.77 8.62 5.39 15.16 9.46 9.79a 均值Mean 38.92 32.16 36.01 34.35 39.28 35.98 35.48 37.72 31.50 29.49 34.51 35.97 33.57 38.91 35.00 35.26 注:居群代码见表 1,表型性状代码1~18见表 2,下同。“均值”列不同字母表示性状间在0.05水平上差异显著。
Note:Code of population see table 1, codes of phenotypic traits 1-18 see table 2, the same below. Paired populations with the different letters are significantly different (P<0.05) shown in ‘Mean’ line,and Paired traits with the different letters are significantly different (P<0.05) shown in ‘Mean’ column.表 6 凉山州核桃坚果表型方差分量和表型分化系数
Table 6. Variance components and phenotypic differentiation coefficients of nut phenotypic traits of walnut populations in Liangshan prefecture
表型性状代码
Code of phenotypic traits方差分量Variance component 方差分量百分比Percentage of variance component /% 居群间表型分化系数
VST /%居群间
Among populations居群内
Within populations居群间
Among populations居群内
Within populations1 3.63×10-1 5.22 6.50 93.50 6.50 2 6.03×10-2 11.64 0.52 99.48 0.52 3 1.56×10-2 2.86×10-1 5.19 94.81 5.19 4 1.68×10-2 1.98 0.84 99.16 0.84 5 1.84×10-3 2.81×10-1 0.65 99.35 0.65 6 2.91×10-3 1.82 0.16 99.84 0.16 7 2.97×10-1 3.69 7.46 92.54 7.46 8 3.00×10-1 2.42 11.03 88.97 11.03 9 6.05×10-3 4.70×10-1 1.27 98.73 1.27 10 7.35×10-2 7.23×10-1 9.23 90.77 9.23 11 6.61×10-2 6.67×10-1 9.01 90.99 9.01 12 1.37×10-1 1.23 10.07 89.93 10.07 13 4.06×10-1 4.17 8.87 91.13 8.87 14 1.47 12.49 10.53 89.47 10.53 15 1.46×10-2 1.72×10-1 7.81 92.19 7.81 16 3.09×10-4 6.30×10-3 4.68 95.32 4.68 17 5.73 78.38 6.81 93.19 6.81 18 1.01 11.72 7.92 92.08 7.92 均值Mean 5.54×10-1 7.63 6.03 93.97 6.03 -
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