Morphological Characteristics of Pollens of Varieties of Luffa
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摘要:
目的 观察丝瓜品种间花粉粒形态特征,研究其品种间的差异性及各品种间的亲缘关系,为丝瓜品种的分类鉴定提供孢粉学依据。 方法 利用扫描电镜对20个丝瓜栽培品种的花粉进行观察,对所考察性状进行主成分分析和类平均法(UPGMA)聚类分析。 结果 参试品种的花粉粒形态相似,均为长球形,赤道面观为长椭圆形,极面观为三裂圆形,具3条萌发沟,裂至接近两极,未相接,均属于N3P4C5型花粉;品种间的花粉极轴长107.03~112.74 µm、赤道轴长57.17~61.04 µm、花粉大小6164.15~6814.02 µm2、花粉外壁均为网状纹饰,网孔密度为0.129~0.220 n·µm−2,均存在显著差异;经主成分分析可知,影响花粉分类的主要性状指标有极轴长、赤道长、花粉大小、花粉形状和外壁纹饰;对考察的8个花粉性状的聚类分析,在欧氏距离23的水平上,将20个品种聚类为三大类群。 结论 花粉粒极轴长、赤道轴长、形状、大小和网孔密度等性状差异可用作丝瓜品种间亲缘关系及分类鉴定的孢粉学依据。 Abstract:Objective Pollens morphology of a variety of Luffa cylindrica was characterized for taxonomic classification of the plants. Methods The palynological characteristics of 20 varieties of luffa were examined under a scanning electron microscope. The observation was analyzed for classification on the plant species using principal component analysis and UPGMA clustering method. Result Belonging to the N3P4C5 type, the pollens were similarly prolate in shape—oblong equatorially with a diameter ranging from 57.17 μm to 61.04 μm and trilobate-circular latitudinally with an axis ranging between 107.03 μm and 112.74 μm. The pollen size varied from 6 164.15 μm2 to 6 814.02 μm2. There were 3 separate apertures that almost reached both ends of a pollen. The exine was reticulated with a spinule density of 0.129-0.220 n·μm−2, which differed significantly among the varieties. The major palynological traits for taxonomic differentiation included polar axis length, equatorial diameter, size, shape, and exine appearance of the pollens. On a Euclidean distance of 23 on 8 traits, the 20 luffa varieties were clustered into 3 categories. Conclusion Differences on polar axis length, equatorial diameter, shape, size, and spinule density of pollens provided the basis to classify the genetic relationship of 20 different species of luffa. -
Key words:
- Luffa cylindrica /
- pollen morphology /
- scanning electron microscope /
- classification
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图 1 扫描电镜下不同丝瓜栽培品种花粉形态
1:三比2号;2:三比6号;3:优比1号;4:春香;5:白雪玉;6:秀丽;7:早秀1号;8:大皱皮;9:玉兔;10:早帅201;11:早靓1号;12:荷兰秀美;13:早佳玉;14:银田短棒;15:农杂八号;16:雅丽皱皮;17:泰国玉龙;18:精选雅绿;19:延陵丝瓜;20:农福805。a:花粉粒极面观;b:花粉粒赤道观面;c:花粉粒表面纹饰。
Figure 1. SEM images of luffa pollen morphology
1: Sanbi No. 2; 2:Sanbi No. 6; 3: Youbi No. 1; 4: Spring fragrance; 5: White snow jade; 6: Xiuli; 7: Zaoxiu No. 1; 8: Large wrinkled skin; 9: Jade rabbit; 10: Zaoshuai No. 201; 11: Zaoliang No. 1; 12: Beautiful Netherlands; 13: ZaoJia jade; 14: Yintian short stick; 15: Nongza No. 8; 16: Elegant wrinkled skin; 17: Thailand Yulong; 18: Selected elegant green; 19: Yanling luffa; 20: Nongfu 805. a: Latitudinal view of pollen; b: Equatorial view of pollen; c: Surface appearance of pollen.
表 1 20个丝瓜栽培品种来源
Table 1. Origins of 20 luffa varieties
序号
Number品种
Variety熟性类型
Maturity type品种来源 Variety source 公司 Company 地区 Area 1 三比2号 SanbiNo.2 早熟 鼎牌种苗 湖南常德 2 三比6号 Sanbi No.6 早熟 鼎牌种苗 湖南常德 3 优比1号 Youbi No.1 特早熟 鼎牌种苗 湖南常德 4 春香 Spring fragrance 极早熟 银田蔬菜种子实业 湖南长沙 5 白雪玉 White snow jade 早熟 三诚种业 浙江金华 6 秀丽 Xiuli 特早熟 富新源种业 四川成都 7 早秀1号 Zaoxiu No.1 早熟 四川科喜种业 四川成都 8 大皱皮 Large wrinkled skin 早熟 四川科喜种业 四川成都 9 玉兔 Jade rabbit 中早熟 广东和利农 广东澄海 10 早帅201 Zaoshuai No.201 极早熟 银田蔬菜种子实业 湖南长沙 11 早靓1号 Zaoliang No.1 特早熟 四川泸旺种业 四川宜宾 12 荷兰秀美 Beautiful Netherlands 早熟 春秋种业 山东寿光 13 早佳玉 ZaoJia jade 早熟 种都高科种业 四川成都 14 银田短棒 Yintian short stick 早熟 银田蔬菜 湖南长沙 15 农杂八号 Nongza No. 8 早熟 农之子种业 湖南株洲 16 雅丽皱皮 Elegant wrinkled skin 早熟 种都高科种业 四川成都 17 泰国玉龙 Thailand Yulong 早熟 兴强种业 山东寿光 18 精选雅绿 Selected elegant green 早熟 青岛沃田种业 山东青岛 19 延陵丝瓜 Yanling luffa 中早熟 地方品种 福建泉州 20 农福805 Nongfu 805 早熟 超大种业 福建福州 表 2 不同丝瓜栽培品种花粉形态特征比较
Table 2. Morphological characteristics of pollens of L. cylindrica varieties
序号
Number品种
Variety极轴长
Polar axis
length/µm赤道轴长
Equator axis
length/µm萌发沟长
Germinal furrow
length/µm沟脊宽度
Ridge width/
µm花粉大小 P×E
Pollen grain
size/µm2花粉形状 P/E
Pollen grain
shape1 三比2号 SanbiNo.2 107.03±2.69 g 58.49±2.87 defgh 96.94±3.32 hi 34.88±2.08 fghi 6265.39±435.36 ef 1.83±0.01 efg 2 三比6号 Sanbi No.6 109.95±2.82 cde 60.72±2.28 a 100.47±2.76 bcdef 36.79±2.84 abcde 6681.13±397.95 abc 1.81±0.01 fg 3 优比1号 Youbi No.1 108.84±2.04 ef 60.66±1.96 a 95.73±3.61 i 36.35±1.71 bcdefg 6603.64±285.78 abcd 1.80±0.01 fg 4 春香 Spring fragrance 109.53±2.73 de 60.07±2.21 abc 98.92±3.41 fgh 35.74±2.93 cdefg 6582.17±352.32 bcd 1.82±0.01 fg 5 白雪玉 White snow jade 109.74±2.09 de 59.88±1.95 abcd 98.08±3.12 gh 36.26±2.2 bcdefg 6573.33±300.51 bcd 1.83±0.01 efg 6 秀丽Xiuli 111.56±1.87 abc 59.1±1.76 bcdef 102.24±2.71 abc 35.90±2.09 cdefg 6594.29±275.14 abcd 1.89±0.01 abc 7 早秀1号 Zaoxiu No.1 112.74±3.10 a 60.43±2.07 ab 102.80±4.15 a 33.65±2.66 ij 6814.02±341.61 a 1.87±0.02 cde 8 大皱皮 Large wrinkled skin 112.71±2.06 a 60.28±2.08 ab 102.36±2.99 ab 32.71±2.85 j 6795.15±297.77 ab 1.87±0.01 cde 9 玉兔 Jade rabbit 109.76±2.04 de 57.17±1.44 h 99.87±2.72 defg 33.77±1.81 hij 6275.6±235.16 ef 1.92±0.01 ab 10 早帅201Zaoshuai No.201 112.22±2.1 ab 58.15±1.62 efgh 101.87±2.07 abcd 35.39±1.69 defgh 6526.41±259.72 cd 1.93±0.01 a 11 早靓1号 Zaoliang No.1 107.55±1.82 fg 57.30±1.76 gh 98.17±2.76 gh 36.40±2.41 bcdef 6164.15±247.41 f 1.88±0.01 bcd 12 荷兰秀美 Beautiful Netherlands 110.74±2.20 bcd 58.39±2.11 defgh 101.41±3.08 abcde 38.40±2.12 a 6467.21±294.14 cde 1.90±0.02 abc 13 早佳玉 ZaoJia jade 110.83±2.63 bcd 57.76±2.13 fgh 100.12±2.55 cdefg 35.33±2.85 efghi 6403.88±338.55 de 1.92±0.01 ab 14 银田短棒 Yintian short stick 109.80±1.81 de 58.36±2.2 efgh 101.44±2.35 abcde 38.30±3.17 a 6409.97±321.64 de 1.88±0.01 bc 15 农杂八号 Nongza No. 8 112.69±2.60 a 58.71±1.68 cdefg 102.88±3.12 a 36.88±2.41 abcde 6617.71±283.97 abcd 1.92±0.01 ab 16 雅丽皱皮 Elegant wrinkled skin 109.30±1.87 de 61.04±1.58 a 100.29±2.53 bcdefg 37.78±2.42 ab 6672.28±223.07 abc 1.79±0.01 g 17 泰国玉龙 Thailand Yulong 110.12±2.72 cde 60.50±1.76 ab 99.84±3.33 defg 34.61±2.54 ghi 6664.38±298.36 abc 1.82±0.01 fg 18 精选雅绿 Selected elegant green 108.55±2.11 efg 59.67±2.29 abcde 99.05±2.04 fg 37.88±2.94 ab 6479.07±331.39 cde 1.82±0.01 fg 19 延陵丝瓜 Yanling luffa 109.47±2.26 de 58.71±3.05 cdefg 100.17±3.52 cdefg 37.22±1.94 abc 6427.82±381.32 de 1.87±0.02 cde 20 农福805Nongfu 805 108.54±2.32 efg 59.14±2.08 bcdef 99.38±3.23 efg 37.12±2.17 abcd 6418.71±264.45 de 1.84±0.02 def 同列数据后不同小写字母表示差异显著(P<0.05),表3同。
Data with different lowercase letters on same column indicate significant difference (P<0.05). Same for Table 3.表 3 不同丝瓜栽培品种花粉表面纹饰特征比较
Table 3. Pollen appearance and morphological characteristics of luffa varieties
序号
Number品种
Variety网脊宽
Strip ridge width/nm网孔密度
Pore density/(n·µm−2)外壁纹饰
External ornamentation1 三比2号 SanbiNo.2 639.41±115.03 g 0.207±0.031 ab 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 2 三比6号 Sanbi No.6 829.13±109.67 ab 0.170±0.019 def 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 3 优比1号 Youbi No.1 830.34±167.15 a 0.144±0.018 gh 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 4 春香 Spring fragrance 713.77±103.95 defg 0.155±0.020 fg 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 5 白雪玉 White snow jade 523.74±137.87 h 0.147±0.030 g 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 6 秀丽 Xiuli 717.47±74.19 defg 0.176±0.020 de 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 7 早秀1号 Zaoxiu No.1 761.23±152.07 abcde 0.129±0.014 h 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 8 大皱皮 Large wrinkled skin 676.56±105.48 fg 0.160±0.014 efg 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 9 玉兔 Jade rabbit 741.83±84.15 cdef 0.150±0.021 g 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 10 早帅201Zaoshuai No.201 773.01±109.5 abcde 0.148±0.026 g 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 11 早靓1号 Zaoliang No.1 756.11±84.95 abcdef 0.170±0.020 def 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 12 荷兰秀美 Beautiful Netherlands 719.43±130.2 defg 0.197±0.037 bc 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 13 早佳玉 ZaoJia jade 745.68±111.28 bcdef 0.145±0.022 gh 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 14 银田短棒 Yintian short stick 766.25±93.5 abcde 0.175±0.022 de 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 15 农杂八号 Nongza No. 8 814.35±131.2 abc 0.180±0.033 d 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 16 雅丽皱皮 Elegant wrinkled skin 791.57±96.59 abcd 0.199±0.03 bc 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 17 泰国玉龙 Thailand Yulong 751.57±102.78 abcdef 0.182±0.032 cd 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 18 精选雅绿 Selected elegant green 769.91±114.53 abcde 0.202±0.037 b 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 19 延陵丝瓜 Yanling luffa 724.58±117.19 def 0.198±0.018 bc 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 20 农福805 Nongfu 805 688.01±81.86 efg 0.220±0.035 a 网状纹饰,穿孔,孔穴不规则,网脊凸起明显 表 4 主成分的特征根、贡献率和累积贡献率
Table 4. Eigenvalue, contribution rate and total contribution of principal components
主成分
Principal
component特征根
Eigenvalue贡献率
Contribution
rate/%累积贡献率
Total contribution
rate/%第一主成分
Principal component 12.934 36.678 36.678 第二主成分
Principal component 22.305 28.816 65.494 第三主成分
Principal component 31.401 17.519 83.013 第四主成分
Principal component 40.88 11.005 94.018 第五主成分
Principal component 50.405 5.062 99.080 第六主成分
Principal component 60.073 0.918 99.998 第七主成分
Principal component 70.000 0.001 99.999 第八主成分
Principal component 80.000 0.001 100.000 表 5 主成分分析特征
Table 5. Eigenvectors of principal components analysis
性状
Characters第一主成分
Principal
component 1第二主成分
Principal
component 2第三主成分
Principal
component 3第四主成分
Principal
component 4极轴长
Polar axis length0.970 −0.059 0.127 0.128 赤道轴长
Equator axis length0.166 0.984 0.008 0.062 花粉形状
P/E Pollen shape0.469 −0.874 0.077 0.025 花粉大小
P×E Size pollen0.796 −0.208 0.443 0.286 萌发沟长度
Germinal sulcus length0.196 0.103 0.611 −0.731 网脊宽
Reticular ridge width−0.506 0.037 0.723 0.056 沟脊宽
Strip ridge width−0.597 0.032 0.531 0.475 网孔密度
Pore density0.679 0.716 0.073 0.120 -
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