Identification and Evaluation of Waterlogging Tolerance of 43 Clematis varieties
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摘要:目的
耐涝性是铁线莲园林应用的关键因素，耐涝性鉴定对铁线莲引种驯化和抗逆性育种具有重要意义。
方法通过分析43个铁线莲园艺品种涝渍胁迫后的表型，将叶片形态、叶片颜色、茎秆形态、茎秆颜色4个表型指标定性分级，结合相对电导率、相对含水量、叶绿素含量等生理指标，采用主成分分析和聚类分析法，综合评价其耐涝性。
结果涝渍胁迫后铁线莲叶片相对含水量、叶绿素a、叶绿素b、总叶绿素及类胡萝卜素含量降低，相对电导率增加；叶片含水量、叶色、叶形态、茎色、茎形态均与相对电导率显著相关，耐涝性综合评分与相对电导率、叶色、叶形态、茎色、茎形态显著相关；聚类分析将43个铁线莲品种分为耐涝型、中间型、不耐涝型、极不耐涝型，分别有2个、16个、6个和19个。
结论初步明确43个铁线莲品种耐涝性分级的特征，为铁线莲耐涝性鉴定提供参考。
Abstract:Waterlogging tolerance was the key to Clematis application, and identification of waterlogging tolerance was important for introduction and domestication of Clematis and stress resistance breeding. In this study, the phenotype of 43 Clematis horticultural varieties after waterlogging stress was analyzed, and four appearance morphological indexes （leaf color, leaf shape, stem color and stem shape） were qualitatively graded. Combined with the physiological indexes of relative conductivity, relative water content and chlorophyll content, principal component analysis and cluster analysis were used to comprehensively evaluate their waterlogging tolerance. The results showed that the relative water content, chlorophyll a, chlorophyll b, total chlorophyll and carotenoid contents of Clematis leaves decreased after waterlogging stress, while the relative conductivity increased. Leaf water content, leaf color, leaf shape, stem color and stem shape were significantly correlated with electrical conductivity, and the total evaluation score of waterlogging tolerance was significantly correlated with electrical conductivity, leaf color, leaf shape, stem color and stem shape. Forty-three Clematis horticultural varieties were classified into waterlogging tolerant types, intermediate types, intolerant types and extremely intolerant types by cluster analysis, including 2 waterlogging tolerant types, 16 intermediate types, 6 waterlogging intolerant varieties and 19 extremely intolerant Clematis. The characteristics of waterlogging tolerance classification of 43 Clematis avarieties were preliminatively identified. An evaluation system of Clematis waterlogging tolerance was initially established to provide reference for identification of Clematis waterlogging tolerance.
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Keywords:
- Clematis /
- waterlogging tolerance /
- morphological criteria /
- evaluation system
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表 1 供试试验材料
Table 1 Plant materials used in this study
序号 品种 来源 序号 品种 来源 1 ‘小鸭’‘Piilu’ 爱沙尼亚 23 ‘志愿者’ ‘Volunteer’ 英国 2 ‘红衣主教’ ‘Rouge Cardinal’ 法国 24 ‘别致’ ‘Bieszczady’ 英国 3 ‘琉璃’ ‘Ruriokoshi’ 日本 25 ‘红星’ ‘Red Star’ 日本 4 ‘包查德女伯爵’ ‘Comtesse de Bouchaud’ 法国 26 ‘变心’ ‘Change of Heart’ 波兰 5 ‘倒影’ ‘Reflections’ 英国 27 ‘爱莎’ ‘Asao’ 日本 6 ‘多蓝’‘Multi Blue’ 荷兰 28 ‘瑞贝卡’ ‘Rebecca’ 英国 7 ‘乌托邦’ ‘Utopia’ 日本 29 ‘神秘面纱’ ‘Night Veil’ 日本 8 ‘宝石蓝光’ ‘Blue Light’ 荷兰 30 ‘努比亚’ ‘Nubia’ 英国 9 ‘黎明的天空’ ‘Dawn’ 波兰 31 ‘薇安’ ‘Vyvyan Pennell’ 英国 10 ‘水晶喷泉’ ‘Crystal Fountain’ 日本 32 ‘里昂村庄’ ‘Ville de Lyon’ 法国 11 ‘繁星’ ‘Nelly Moser’ 法国 33 ‘莎拉芬娜’ ‘Serafina’ 波兰 12 ‘天幕坠落’ ‘Skyfall’ 波兰 34 ‘撒玛利亚’ ‘Samaritan Jo’ 英国 13 ‘波罗尼亚’ ‘Viva Polonia’ 波兰 35 ‘丹空’ ‘Yuan’ 日本 14 ‘向阳’ ‘Sunny Side’ 日本 36 ‘钻石球’ ‘Diamond Ball’ 波兰 15 ‘斯丽’‘Thyrislund’ 英国 37 ‘哥白尼’ ‘Kopernik’ 波兰 16 ‘粉香槟’ ‘Pink Champagne’ 日本 38 ‘薇妮沙’ ‘Venosa Violacea’ 法国 17 ‘白王冠’ ‘Hakaookan’ 日本 39 ‘欧啦啦’ ‘Ooh La La’ 英国 18 ‘钻石’ ‘Diamantina’ 英国 40 ‘倪碧欧’ ‘Niobe’ 波兰 19 ‘蜥蜴’ ‘Basilisk’ 日本 41 ‘朱莉安’ ‘Juliane’ 英国 20 ‘金银丝’ ‘Filigree’ 英国 42 ‘玛丽’ ‘Mary Rose’ 英国 21 ‘大河’ ‘Taiga’ 日本 43 ‘中国红’ ‘Westerplatte’ 波兰 22 ‘巴黎风情’ ‘Parisienne’ 英国 
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表 2 叶片形态指标定级范围及得分标准
Table 2 Leaf morphology index rating range and scoring criteria
实测值定级范围 形态 颜色 叶片形态 得分/分 叶颜色 得分/分 Ⅰ 叶片自然伸展 7 全株叶片绿色 5 Ⅱ 植株下层叶片萎蔫下垂 6 近水处边缘失绿 4 Ⅲ 植株中下层叶片萎蔫下垂 5 黄色或褐色腐烂叶<1/3 3 Ⅳ 植株上层叶片稍萎蔫，中下层叶片萎蔫下垂 4 黄色或褐色腐烂叶1/3～～1/2 2 Ⅴ 全株叶片萎蔫下垂 3 黄色或褐色腐烂叶>1/2 1 Ⅵ 植株下层叶片腐烂，中上层叶片干枯、褶皱 2 Ⅶ 大部分叶片干枯腐烂 1
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表 3 茎形态指标定级范围及得分标准
Table 3 Stem morphology index rating range and scoring criteria
实测值定级范围 形态 颜色 茎形态 得分/分 茎颜色 得分/分 Ⅰ 正常 4 自然色 4 Ⅱ 表面水渍状膨大 3 浅褐色 3 Ⅲ 水渍状肥肿 2 褐化 2 Ⅳ 水渍状腐烂 1 发黑 1
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表 4 43个铁线莲园艺品种形态涝害指数得分
Table 4 Morphological waterlogging damage index scores of 43 Clematis.
序号 品种 各评价指标/分 综合评分/分 评价等级 叶形态 叶色 茎形态 茎色 1 ‘小鸭’ 1 1 1 1 4 Ⅳ 2 ‘红衣主教’ 4 3 3 4 14 Ⅱ 3 ‘琉璃’ 4 3 3 2 12 Ⅱ 4 ‘包查德女伯爵’ 5 2 4 4 15 Ⅱ 5 ‘倒影’ 2 1 4 3 10 Ⅲ 6 ‘多蓝’ 5 3 4 1 13 Ⅱ 7 ‘乌托邦’ 1 1 1 1 4 Ⅳ 8 ‘蓝光’ 4 2 3 4 13 Ⅱ 9 ‘黎明的天空’ 5 2 4 4 15 Ⅱ 10 ‘水晶喷泉’ 2 1 2 1 6 Ⅳ 11 ‘繁星’ 3 1 4 1 9 Ⅲ 12 ‘天幕坠落’ 5 3 4 4 16 Ⅱ 13 ‘波罗尼亚’ 3 5 2 3 13 Ⅱ 14 ‘向阳’ 5 3 4 4 16 Ⅱ 15 ‘斯丽’ 1 1 4 2 8 Ⅲ 16 ‘粉香槟’ 2 1 4 1 8 Ⅲ 17 ‘白王冠’ 2 2 2 1 7 Ⅳ 18 ‘钻石’ 2 2 2 1 7 Ⅳ 19 ‘蜥蜴’ 2 1 4 2 9 Ⅲ 20 ‘金银丝’ 2 1 1 1 5 Ⅳ 21 ‘大河’ 1 1 1 1 4 Ⅳ 22 ‘巴黎风情’ 6 3 4 4 17 Ⅰ 23 ‘志愿者’ 1 1 1 1 4 Ⅳ 24 ‘别致’ 7 5 4 4 20 Ⅰ 25 ‘红星’ 1 1 3 1 6 Ⅳ 26 ‘变心’ 1 1 1 1 4 Ⅳ 27 ‘爱莎’ 3 2 2 1 8 Ⅲ 28 ‘瑞贝卡’ 2 1 2 1 6 Ⅳ 29 ‘神秘面纱’ 5 3 4 4 16 Ⅱ 30 ‘努比亚’ 1 1 1 1 4 Ⅳ 31 ‘薇安’ 1 1 2 1 5 Ⅳ 32 ‘里昂村庄’ 5 3 3 2 13 Ⅱ 33 ‘塞拉菲娜’ 4 2 4 3 13 Ⅱ 34 ‘撒玛利亚’ 3 2 4 3 12 Ⅱ 35 ‘円空’ 2 2 2 1 7 Ⅳ 36 ‘钻石球’ 1 1 1 1 4 Ⅳ 37 ‘哥白尼’ 3 3 4 3 13 Ⅱ 38 ‘薇尼莎’ 6 3 3 1 13 Ⅱ 39 ‘哦啦啦’ 2 1 2 1 6 Ⅳ 40 ‘倪欧碧’ 1 1 1 1 4 Ⅳ 41 ‘朱莉安’ 3 2 4 3 12 Ⅱ 42 ‘玛丽玫瑰’ 1 1 2 1 5 Ⅳ 43 ‘中国红’ 1 2 2 2 7 Ⅳ
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表 5 43个铁线莲园艺品种主要生理指标
Table 5 Main physiological indexes of 43 Clematis.
序号 品种 相对含水量/% 相对电导率/（mS·cm−1） 叶绿素a/（mg·g−1） 叶绿素b/（mg·g−1） 总叶绿素/（mg·g−1） 类胡萝卜素/（mg·g−1） CK T CK T CK T CK T CK T CK T 1 ‘小鸭’ 87.29±0.45 76.68±0.48 34.29±1.60 72.31±2.60 2.80±0.05 1.83±0.02 1.64±0.04 1.02±0.01 4.47±0.09 2.87±0.03 10.86±0.21 7.22±0.07 2 ‘红衣主教’ 78.69±0.20 73.95±0.56 50.64±1.77 52.72±0.43 2.59±0.04 2.36±0.02 1.39±0.03 1.29±0.01 4.01±0.07 3.67±0.04 10.10±0.18 9.11±0.09 3 ‘琉璃’ 83.14±0.41 78.41±0.09 26.32±0.45 38.96±0.51 2.45±0.04 1.75±0.08 1.27±0.02 0.93±0.05 3.75±0.06 2.70±0.13 9.50±0.17 7.06±0.33 4 ‘包查德女伯爵’ 83.33±0.19 81.19±0.29 34.45±0.21 50.33±2.79 2.48±0.03 1.40±0.05 1.37±0.02 0.78±0.03 3.88±0.05 2.19±0.08 9.59±0.12 5.41±0.18 5 ‘倒影’ 76.52±0.76 74.91±0.37 41.67±1.29 53.87±1.81 2.87±0.04 1.92±0.05 1.75±0.04 1.02±0.03 4.65±0.08 2.95±0.08 11.35±0.19 7.39±0.24 6 ‘多蓝’ 82.52±0.29 79.51±0.67 26.34±0.34 35.38±0.79 1.91±0.05 1.17±0.03 1.08±0.03 0.59±0.02 3.01±0.08 1.78±0.05 7.75±0.22 4.44±0.12 7 ‘乌托邦’ 77.24±0.73 54.52±0.55 32.50±0.41 93.20±0.44 2.27±0.06 1.97±0.07 1.24±0.04 1.12±0.04 3.53±0.10 3.11±0.11 8.99±0.27 8.87±0.31 8 ‘蓝光’ 79.88±0.38 79.62±0.34 35.45±0.38 38.62±0.57 1.82±0.07 1.14±0.04 0.96±0.03 0.62±0.02 2.80±0.10 1.77±0.06 7.00±0.23 4.61±0.16 9 ‘黎明的天空’ 85.22±0.25 80.58±0.16 35.92±0.93 55.09±2.38 2.54±0.03 1.63±0.11 1.35±0.02 0.91±0.06 3.92±0.05 2.56±0.17 9.69±0.11 7.02±0.48 10 ‘水晶喷泉’ 80.80±0.17 77.88±0.14 44.91±0.59 67.54±0.91 2.43±0.02 1.48±0.04 1.30±0.01 0.83±0.02 3.76±0.03 2.33±0.06 9.40±0.09 5.95±0.18 11 ‘繁星’ 79.49±0.42 74.85±0.26 32.72±0.27 85.13±0.76 2.04±0.05 1.37±0.02 1.10±0.03 0.74±0.01 3.16±0.08 2.12±0.03 8.06±0.20 5.90±0.08 12 ‘天幕坠落’ 82.79±0.06 76.84±0.23 49.66±0.81 51.12±0.51 2.24±0.03 1.51±0.03 1.19±0.02 0.83±0.02 3.45±0.04 2.35±0.05 8.57±0.11 5.87±0.13 13 ‘波罗尼亚’ 81.69±0.08 77.14±0.51 28.86±0.41 48.76±0.89 2.33±0.06 0.97±0.02 1.23±0.04 0.52±0.01 3.59±0.10 1.51±0.03 8.85±0.26 4.08±0.07 14 ‘向阳’ 83.19±0.25 78.45±0.28 27.26±0.60 40.53±1.35 2.11±0.06 1.50±0.02 1.11±0.03 0.82±0.01 3.25±0.09 2.34±0.04 8.16±0.23 6.04±0.10 15 ‘斯丽’ 83.70±0.18 67.30±2.13 23.36±0.35 91.67±0.73 1.85±0.08 1.07±0.01 1.01±0.05 0.57±0.01 2.88±0.13 1.64±0.02 7.27±0.31 4.42±0.05 16 ‘粉香槟’ 80.75±0.07 75.13±0.43 30.83±0.27 41.70±1.28 2.23±0.06 1.72±0.03 1.22±0.03 0.98±0.02 3.47±0.09 2.72±0.04 8.54±0.22 6.76±0.11 17 ‘白王冠’ 80.78±0.53 70.21±0.18 26.71±0.44 91.47±1.43 1.92±0.05 0.98±0.05 1.05±0.03 0.55±0.03 3.00±0.08 1.54±0.08 7.35±0.19 4.24±0.20 18 ‘钻石’ 79.80±0.19 79.30±0.37 33.68±0.64 79.28±1.58 1.88±0.08 0.72±0.01 1.04±0.04 0.40±0.01 2.94±0.12 1.13±0.02 7.39±0.30 3.34±0.06 19 ‘蜥蜴’ 79.35±0.36 71.37±0.47 34.26±1.02 69.02±0.82 2.65±0.07 2.54±0.02 1.50±0.04 1.43±0.02 4.18±0.11 4.00±0.04 10.41±0.28 10.18±0.11 20 ‘金银丝’ 78.14±1.93 68.27±1.48 25.20±0.08 99.43±0.36 1.91±0.03 1.72±0.05 1.06±0.02 0.94±0.03 2.99±0.05 2.68±0.07 7.22±0.13 6.64±0.17 21 ‘大河’ 78.25±0.16 76.57±0.89 27.99±0.46 52.48±1.25 2.28±0.05 2.12±0.05 1.29±0.03 1.16±0.03 3.59±0.09 3.30±0.08 9.63±0.22 8.87±0.22 22 ‘巴黎风情’ 56.80±3.94 37.15±7.63 91.42±0.84 98.69±0.20 2.24±0.03 1.99±0.02 1.28±0.01 1.19±0.01 3.54±0.04 3.20±0.03 8.62±0.07 7.73±0.04 23 ‘志愿者’ 81.08±2.00 80.49±2.02 45.25±0.29 70.25±0.46 2.77±1.16 0.04±0.01 1.45±0.63 0.01±0.01 4.25±1.80 0.05±0.01 12.20±4.82 0.25±0.04 24 ‘别致’ 79.16±0.49 78.37±0.73 28.92±0.68 31.59±0.62 2.47±0.05 2.34±0.07 1.42±0.03 1.28±0.05 3.92±0.08 3.65±0.12 9.74±0.18 8.77±0.26 25 ‘红星’ 77.48±3.12 60.98±3.64 28.57±0.44 91.04±0.50 1.91±0.04 1.60±0.05 1.07±0.02 0.88±0.03 3.00±0.06 2.50±0.09 7.70±0.16 6.37±0.21 26 ‘变心’ 77.97±2.05 69.19±1.31 51.23±0.81 97.64±0.44 1.70±0.05 1.53±0.03 0.95±0.03 0.85±0.02 2.68±0.08 2.40±0.04 6.69±0.19 6.37±0.12 27 ‘爱莎’ 79.74±0.18 79.21±0.62 32.75±1.67 44.63±0.63 1.86±0.08 1.30±0.03 1.05±0.05 0.72±0.02 2.93±0.13 2.03±0.04 7.43±0.31 4.96±0.11 28 ‘瑞贝卡’ 72.00±2.88 60.55±2.34 29.25±0.29 91.83±0.75 2.44±0.04 2.49±0.07 1.38±0.03 1.44±0.05 3.84±0.07 3.95±0.13 9.63±0.18 9.83±0.35 29 ‘神秘面纱’ 83.19±1.44 74.67±0.94 32.67±0.49 38.33±1.36 2.71±0.08 1.52±0.01 1.53±0.05 0.80±0.01 4.27±0.13 2.34±0.02 10.91±0.37 6.21±0.04 30 ‘努比亚’ 85.31±1.53 78.16±0.88 44.47±0.47 76.90±2.50 3.59±2.60 0.20±0.12 1.66±1.38 0.05±0.04 5.29±4.01 0.24±0.16 14.78±12.48 0.90±0.81 31 ‘薇安’ 77.03±0.57 74.46±0.91 34.39±0.36 38.29±0.36 2.41±0.04 1.30±0.04 1.33±0.02 0.72±0.02 3.77±0.06 2.03±0.07 9.20±0.18 5.23±0.16 32 ‘里昂村庄’ 80.13±0.31 79.51±0.58 30.18±0.83 33.09±0.35 2.41±0.09 2.23±0.04 1.34±0.06 1.24±0.02 3.78±0.15 3.50±0.07 9.58±0.37 9.02±0.17 33 ‘塞拉菲娜’ 80.16±0.34 77.99±0.69 38.24±0.49 48.50±1.00 2.58±0.05 1.91±0.04 1.46±0.03 1.08±0.02 4.07±0.08 3.01±0.06 10.26±0.18 7.46±0.14 34 ‘撒玛利亚’ 78.75±0.13 74.25±0.19 48.54±0.18 60.67±0.60 3.41±0.15 1.99±0.02 1.89±0.07 1.17±0.02 5.34±0.12 3.18±0.04 13.86±0.65 7.76±0.09 35 ‘円空’ 81.01±0.43 77.13±0.23 35.82±0.97 66.36±0.60 2.57±0.07 2.10±0.05 1.44±0.05 1.23±0.03 4.03±0.13 3.35±0.08 9.74±0.29 8.37±0.21 36 ‘钻石球’ 80.37±0.40 73.83±0.27 48.57±1.48 94.17±0.65 1.99±0.03 1.94±0.05 1.06±0.02 1.06±0.03 3.07±0.05 3.03±0.08 8.13±0.13 7.40±0.18 37 ‘哥白尼’ 78.13±0.22 75.84±0.35 59.78±0.76 77.79±0.87 2.41±0.10 1.84±0.08 1.44±0.09 1.04±0.05 3.88±0.19 2.90±0.13 9.47±0.41 7.40±0.31 38 ‘薇尼莎’ 80.03±0.19 77.27±0.19 38.77±0.47 62.81±2.70 2.51±0.04 2.31±0.02 1.39±0.03 1.29±0.01 3.92±0.07 3.63±0.03 9.96±0.13 9.52±0.09 39 ‘哦啦啦’ 79.91±0.23 69.10±0.24 35.60±0.36 83.28±1.35 2.66±0.04 2.27±0.04 1.49±0.03 1.23±0.02 4.18±0.07 3.52±0.06 10.17±0.19 9.06±0.14 40 ‘倪欧碧’ 79.94±0.30 68.97±0.47 34.99±0.69 70.45±0.60 2.33±0.02 2.01±0.03 1.25±0.01 1.05±0.02 3.60±0.03 3.08±0.04 8.79±0.08 7.87±0.10 41 ‘朱莉安’ 74.36±0.21 73.00±0.29 61.06±0.45 78.40±2.56 2.63±0.05 2.00±0.07 1.45±0.03 1.05±0.04 4.11±0.08 3.07±0.11 10.00±0.18 7.67±0.27 42 ‘玛丽玫瑰’ 80.23±0.22 65.84±1.10 50.54±2.24 50.93±0.78 2.19±0.02 2.47±0.03 1.22±0.02 1.36±0.02 3.44±0.04 3.85±0.06 8.58±0.11 10.02±0.14 43 ‘中国红’ 76.24±0.60 70.66±0.49 60.08±0.65 81.67±0.19 2.34±0.10 1.75±0.03 1.36±0.08 0.92±0.02 3.72±0.18 2.68±0.05 9.01±0.45 7.12±0.13
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表 6 11个指标之间相关性分析
Table 6 Correlation analysis among 11 indicators
指标 相对电导率 相对含水量 叶绿素a 叶绿素b 总叶绿素 类胡萝卜素 叶形态 叶色 茎形态 茎色 总得分 相对电导率 1 相对含水量 −0.604** 1 叶绿素a −0.102 −0.247 1 叶绿素b −0.109 −0.255 0.994** 1 总叶绿素 −0.106 −0.249 0.999** 0.998** 1 类胡萝卜素 −0.084 −0.254 0.994** 0.989** 0.993** 1 叶形态 −0.518** 0.161 0.189 0.200 0.195 0.169 1 叶色 −0.481** 0.199 0.054 0.058 0.057 0.040 0.776** 1 茎形态 −0.373* 0.065 0.189 0.212 0.198 0.157 0.585** 0.377* 1 茎色 −0.394** 0.079 0.108 0.120 0.113 0.076 0.697** 0.613** 0.690** 1 总得分 −0.524** 0.148 0.167 0.182 0.174 0.138 0.921** 0.807** 0.778** 0.878** 1 注：**表示在 P<0.01水平极显著性相关，*表示在 P<0.05水平显著相关。
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表 7 铁线莲耐涝性分级评价
Table 7 Grading evaluation of Clematis waterlogging tolerance
评价等级 分值/分 耐涝性 特征描述 Ⅰ ≥17 耐涝型 植株叶片自然伸展或下层叶片少许萎蔫下垂，叶片绿色或近水处边缘失绿，茎秆正常呈自然色，叶片相对电导率增加10%以下，叶绿素含量降低10%以下 Ⅱ 11～16 中间型 植株中下层叶片萎蔫下垂，黄色或褐色腐烂叶片小于1/3，茎秆正常，颜色呈自然色或浅褐色，叶片相对电导率增加10%～70% Ⅲ 8～10 不耐涝型 植株大部分叶片萎蔫下垂，黄色或褐色腐烂叶1/3至1/2，茎秆形态正常，颜色褐化或发黑，叶片相对电导率增加70%以上 Ⅳ ≤7 极不耐涝型 植株大部分叶片干枯腐烂，黄色或褐色腐烂叶>1/2，颜色褐化或发黑。呈水渍状腐烂，叶片相对电导率增加110%以上
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