3种毛茛科地被植物对遮阴的生理响应及耐阴性评价
为探究3种毛茛科野生地被植物在不同遮阴条件下的耐阴状况，以毛茛(Ranunculus japonicus)、华东唐松草(Thalictrum fortunei)和卵瓣还亮草(Delphinium anthriscifolium var. savatieri)为试验材料，以全光照为对照(CK)，人工搭棚设置30%、50%、70%和90%遮阴处理，测定上述3种毛茛科植物的形态和生理指标并进行耐阴性综合分析，确定3种植物的耐阴性强弱。结果表明，随着遮阴程度增加，3种毛茛科植物的叶面积、叶长、叶宽、株高、株幅、茎粗总体呈先升后降的趋势；过氧化物酶(POD)活性、丙二醛(MDA)含量总体呈先降后升的趋势；比叶重呈下降趋势。随着遮阴程度的增加，毛茛和卵瓣还亮草的超氧化物歧化酶(SOD)活性总体呈下降趋势，华东唐松草的SOD活性呈先降后升的趋势；毛茛的过氧化氢酶(CAT)活性呈下降趋势，华东唐松草和卵瓣还亮草的CAT活性呈先降后升的趋势；毛茛和华东唐松草脯氨酸(Pro)含量呈先降后升的趋势，卵瓣还亮草的Pro含量呈下降趋势。不同植物各指标的变化存在差异，通过隶属函数法进行耐阴性综合评价，结果显示，3种植物的耐阴性存在差异，3种毛茛科植物耐阴性强弱表现为华东唐松草 > 卵瓣还亮草 > 毛茛。本研究结果可为其在不同遮阴程度的环境下栽培提供理论参考。
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图 1 不同遮阴处理下3种毛茛科植物SOD和POD活性的变化
不同小写字母表示同种植物内不同处理间差异显著(P < 0.05)，下图同。
Figure 1. Changes in superoxide dismutase (SOD) and peroxidase (POD) activities of three Ranunculaceae species under different shade treatments
Different lowercase letters indicate significant differences between the different treatments within the same species at the 0.05 level. This is applicable for the following figures as well.
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图 2 不同遮阴处理下3种毛茛科植物CAT活性和MDA含量的变化
Figure 2. Changes in catalase (CAT) activity and malondialdehyde (MDA) content of three Ranunculaceae species under different shade treatments
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图 3 不同遮阴处理下3种毛茛科植物脯氨酸含量和比叶重的变化
Figure 3. Changes in proline content and specific leaf weight of three Ranunculaceae species under different shade treatments
表 1 3种毛茛科植物在不同遮阴处理下的形态指标
Table 1 Morphological indexes of three Ranunculaceae species under different shade treatments
植物
Plant处理
Treatment叶面积
Leaf area/cm2叶长
Leaf length/cm叶宽
Leaf width/cm株高
Plant height/cm株幅
Plant width/cm茎粗
Stem diameter/cmM CK 36.18 ± 1.46c 7.37 ± 0.15b 9.11 ± 0.74b 37.30 ± 2.21bc 34.85 ± 1.24c 0.52 ± 0.01ab T1 51.23 ± 1.94a 8.63 ± 0.23a 9.73 ± 0.12ab 42.03 ± 4.61ab 46.65 ± 1.92b 0.58 ± 0.02a T2 52.85 ± 2.75a 9.17 ± 0.90a 10.53 ± 1.31a 41.43 ± 2.20ab 45.30 ± 1.34b 0.47 ± 0.05bc T3 45.40 ± 0.88b 8.77 ± 0.06a 10.33 ± 0.31ab 44.93 ± 1.03a 51.85 ± 2.57a 0.46 ± 0.05bc T4 28.24 ± 1.40d 7.57 ± 0.65b 9.17 ± 0.45ab 35.90 ± 1.65c 33.87 ± 0.84c 0.43 ± 0.06c H CK 1.87 ± 0.17d 1.85 ± 0.15c 1.47 ± 0.06b 28.00 ± 0.46bc 28.07 ± 1.56b 0.19 ± 0.01bc T1 2.02 ± 0.26cd 2.25 ± 0.13bc 1.60 ± 0.10b 29.23 ± 0.49bc 29.73 ± 0.48ab 0.24 ± 0.01a T2 2.47 ± 0.14b 2.50 ± 0.35ab 2.43 ± 0.29a 29.53 ± 1.38b 30.08 ± 1.26a 0.23 ± 0.01a T3 2.91 ± 0.11a 2.70 ± 0.20a 2.47 ± 0.21a 33.10 ± 0.96a 29.30 ± 0.18ab 0.22 ± 0.03ab T4 2.19 ± 0.07bc 2.22 ± 0.24bc 2.33 ± 0.23a 26.43 ± 2.90c 29.30 ± 0.58ab 0.17 ± 0.02c L CK 3.63 ± 0.23c 3.37 ± 0.25c 4.23 ± 0.12ab 25.00 ± 0.40c 19.78 ± 1.13c 0.25 ± 0.00b T1 5.97 ± 0.47a 5.20 ± 0.30a 4.43 ± 0.21a 35.63 ± 2.35a 26.55 ± 0.69a 0.31 ± 0.03a T2 6.20 ± 0.53a 4.53 ± 0.35b 4.67 ± 0.50a 30.17 ± 0.96b 27.93 ± 1.13a 0.25 ± 0.03b T3 4.50 ± 0.22b 4.33 ± 0.31b 4.40 ± 0.26a 29.03 ± 0.25b 22.93 ± 0.63b 0.24 ± 0.00b T4 1.70 ± 0.17d 3.37 ± 0.07c 3.73 ± 0.15b 18.07 ± 1.10d 9.67 ± 0.15d 0.18 ± 0.01c M：毛茛；H：华东唐松草；L：卵瓣还亮草。CK：对照(全光照)；T1：30%遮阴度；T2：50%遮阴度；T3：70%遮阴度；T4：90%遮阴度。同列不同小写字母表示同种植物不同处理间差异显著(P < 0.05)。下同。
M: R. japonicus; H: T. fortunei; L: D. anthriscifolium var. savatieri. CK: control (full light); T1: 30% shade; T2: 50% shade; T3: 70% shade; T4: 90% shade. Different lowercase letters within the same column indicate significant differences between the different treatments within the same species at the 0.05 level. This is applicable for the following tables and figures as well.
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表 2 3种毛茛科植物各指标的耐阴系数α值
Table 2 Shade tolerance coefficients (α) of three Ranunculaceae species
植物
PlantLA LL LW PH PW SD SLW SOD POD CAT MDA Pro M 1.463 1.246 1.167 1.113 1.301 0.823 0.813 0.388 0.572 0.501 0.895 0.526 H 1.329 1.349 1.656 1.055 1.074 1.242 0.880 0.858 0.886 0.858 0.848 0.897 L 1.708 1.353 1.105 1.206 1.413 1.007 0.740 0.801 0.288 0.280 0.782 0.809 LA：叶面积；LL：叶长；LW：叶宽；PH：株高；PW：株幅；SD：茎粗；SLW：比叶重；SOD：超氧化物歧化酶活性；POD：过氧化物酶活性；CAT：过氧化氢酶活性；MDA：丙二醛含量；Pro：脯氨酸含量。
LA: leaf area; LL: leaf length; LW: leaf width; PH: plant height; PW: plant width; SD: stem diameter; SLW: specific leaf weight; SOD: superoxide dismutase activity; POD: peroxidase activity; CAT: catalase activity; MDA: malondialdehyde content; Pro: proline content.
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表 3 综合指标系数及贡献率
Table 3 Comprehensive index coefficients and proportions
指标
Index综合指标系数
Comprehensive index
coefficient1 2 叶面积 Leaf area −0.335 0.182 叶长 Leaf length 0.069 0.468 叶宽 Leaf width 0.356 0.093 株高 Plant height −0.340 0.166 株幅 Plant width −0.363 0.014 茎粗 Stem diameter 0.264 0.327 比叶重 Specific leaf weight 0.351 −0.117 SOD活性 Superoxide dismutase activity 0.119 0.451 POD活性 Peroxidase activity 0.356 −0.092 CAT活性 Catalase activity 0.361 −0.042 MDA含量 Malondialdehyde content 0.140 −0.440 脯氨酸含量 Proline content 0.158 0.429 特征值 Eigenvalue 7.604 4.396 贡献率 Proportion/% 63.369 36.631 累积贡献率 Cumulative proportion/% 63.369 100.000
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表 4 3种毛茛科植物综合指标值(CI)、隶属函数值(u)及权重、综合评价值(D)及耐阴性排序
Table 4 The comprehensive index value, subordinate function value and weight, comprehensive evaluation value, and order of shade tolerance of three Ranunculaceae species
植物 Plant CI (1) CI (2) μ1 μ2 D 排序 Order 毛茛 R. japonicus −0.705 −2.361 0.304 0.000 0.192 3 华东唐松草 T. fortunei 3.042 0.717 1.000 0.768 0.915 1 卵瓣还亮草 D. anthriscifolium var. savatieri −2.337 1.645 0.000 1.000 0.366 2 权重 Weight 0.634 0.366 CI (1)：综合指标值1；CI (2)：综合指标值2；μ1：隶属函数值1；μ2：隶属函数值2；D：耐阴性综合评价值。
CI (1): comprehensive index value 1; CI (2): comprehensive index value 2; μ1: subordinate function value 1; μ2: subordinate function value 2; D: comprehensive evaluation value of shade tolerance.
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