植物生态学报 ›› 2010, Vol. 34 ›› Issue (7): 827-838.DOI: 10.3773/j.issn.1005-264x.2010.07.008
收稿日期:
2009-08-11
接受日期:
2010-04-21
出版日期:
2010-08-11
发布日期:
2010-07-01
通讯作者:
孙杉
作者简介:
* E-mail: sunshan@mail.kib.ac.cn
SUN Shan*(), CAO Guo-Xing, LUO Yan-Jiang, LI Qing-Jun
Received:
2009-08-11
Accepted:
2010-04-21
Online:
2010-08-11
Published:
2010-07-01
Contact:
SUN Shan
摘要:
花柱多态型的型比通过选型交配形成的负频率依赖的选择来维持。种群空间格局对于频率依赖选择所维持的花柱多态型是至关重要的。该研究使用摆放实验构建不同型比的马来良姜(Alpinia mutica)人工种群, 以评价花柱卷曲性促进选型交配的功能。使用邻域模型调查了云南草蔻(Alpinia blepharaocalyx)种群中的目标植株的坐果数、结籽数及其邻域中花序的数量和型偏以探讨在局域尺度上目标植株的雌性适合度是否受负频率依赖的选择。通过比较云南草蔻上举型和下垂型各繁殖性状的差异以及两型的功能性别的计算以评估两型功能性别可能存在的特化趋势。摆放实验结果说明花柱卷曲性形成的二型种群可以促进型间花粉传递。在局域尺度上, 云南草蔻两型倾向于同型聚集分布, 这一分布特点反映出其克隆生长特性。统计结果说明: 邻域中异型花粉供体相对丰富度对两型目标植株都无影响; 目标植株雌性适合度在邻域中不受负频率依赖选择的影响。这一结果并不支持负频率依赖的选择在该研究尺度上是影响种群型比结构的主要机制。云南草蔻两型个体在花期和果期的繁殖指标上不存在显著差异; 功能性别计算结果显示选型交配是种群内唯一的交配模式时, 下垂型功能性别偏雌性而上举型功能性别偏雄性。这一功能性别特化的趋势可能是通过对上举型的雄性功能的选择而造成的, 并可能与种群型结构相关。当功能性别计算只考虑自交而忽略近交衰退时, 种群功能性别偏向雌性。
孙杉, 操国兴, 罗燕江, 李庆军. 花柱卷曲性的维持及功能性别特化. 植物生态学报, 2010, 34(7): 827-838. DOI: 10.3773/j.issn.1005-264x.2010.07.008
SUN Shan, CAO Guo-Xing, LUO Yan-Jiang, LI Qing-Jun. Maintenance and functional gender specialization of flexistyly. Chinese Journal of Plant Ecology, 2010, 34(7): 827-838. DOI: 10.3773/j.issn.1005-264x.2010.07.008
图1 不同实验摆放对每株种子总数的作用。 A, 全上举型; C, 全下垂型; D, 二型。误差棒表示平均值的标准偏差。不同的字母表示不同差异显著(p < 0.05)。
Fig. 1 The effect of experimental array on the total number of seeds per plant. A, entirely anaflexistylous; C, entirely cataflexistylous; D, dimorphic. Bars indicate SD of mean values. Different letters indicate significantly different at p < 0.05 level.
图2 上举型和下垂型目标植株邻域中平均型偏。 误差棒表示平均值的标准偏差。***, p < 0.001。
Fig. 2 Mean morph bias in neighborhoods surrounding anaflexistylous (Ana-M) and cataflexistlyous (Cata-M) focal plants. Bars indicate SD of mean values. ***, p < 0.001.
图3 型偏与目标植株雌性适合度的散点图。 详细的统计结果见表1。Ana-M, Cata-M, 同图2。
Fig. 3 Scatter plot between female fitness of focal plant and morph bias. Statistical details see Table 1. Ana-M, Cata-M, See Fig. 2.
变量 Variable | 2007 (n = 56) | 2008 (n = 89) | |||
---|---|---|---|---|---|
估计值 Estimate | p | 估计值 Estimate | p | ||
坐果数/株 Fruits/plant | |||||
(型偏)2 (Morph bias)2 | -0.482 2 | 0.480 8 | -1.421 2 | 0.150 1 | |
型偏 Morph bias | -0.274 2 | 0.498 2 | -0.296 1 | 0.595 0 | |
目标植株的表型 Focal plant morph | 0.074 1 | 0.730 9 | -0.500 9 | 0.093 0 | |
邻域密度 Neighborhood density | 0.007 8 | 0.326 6 | -0.000 2 | 0.990 6 | |
(型偏)2 ×目标植株表型 (Morph bias)2 × focal plant morph | 0.477 6 | 0.596 5 | 1.514 4 | 0.200 6 | |
型偏×目标植株表型 Morph bias × focal plant morph | 0.054 3 | 0.914 7 | 0.178 0 | 0.788 4 | |
目标植株的花数 No. flowers in focal plant | 0.026 9 | 0.000 4 *** | 0.019 0 | 0.112 0 | |
结籽数/株 Seeds/plant | |||||
(型偏)2 (Morph bias)2 | -0.362 3 | 0.597 6 | -2.038 0 | 0.104 9 | |
型偏 Morph bias | -0.169 7 | 0.671 1 | -0.634 9 | 0.380 7 | |
目标植株的表型 Focal plant morph | 0.121 8 | 0.574 9 | -0.469 0 | 0.163 7 | |
邻域密度 Neighborhood density | 0.000 9 | 0.909 2 | -0.002 1 | 0.896 7 | |
(型偏)2 ×目标植株表型 (Morph bias)2 × focal plant morph | 0.076 3 | 0.933 2 | 1.946 6 | 0.176 6 | |
型偏×目标植株表型 Morph bias × focal plant morph | 0.134 0 | 0.794 0 | 0.463 4 | 0.573 6 | |
目标植株的花数 No. flowers in focal plant | 0.025 2 | 0.001 0 *** | 0.032 4 | 0.018 6 * |
表1 2007年和2008年型偏的平方、型偏、目标植株表型、邻域密度和目标植株花数对目标植株雌性适合度的作用。分析使用广义线性模型(准泊松分布)
Table 1 The effect of (morph bias)2, morph bias, morph, neighborhood density and the number of flowers per focal plant on female fitness of focal plants in 2007 and 2008, respectively, analyzed with generalized linear models with the log link function for quasipoisson variables
变量 Variable | 2007 (n = 56) | 2008 (n = 89) | |||
---|---|---|---|---|---|
估计值 Estimate | p | 估计值 Estimate | p | ||
坐果数/株 Fruits/plant | |||||
(型偏)2 (Morph bias)2 | -0.482 2 | 0.480 8 | -1.421 2 | 0.150 1 | |
型偏 Morph bias | -0.274 2 | 0.498 2 | -0.296 1 | 0.595 0 | |
目标植株的表型 Focal plant morph | 0.074 1 | 0.730 9 | -0.500 9 | 0.093 0 | |
邻域密度 Neighborhood density | 0.007 8 | 0.326 6 | -0.000 2 | 0.990 6 | |
(型偏)2 ×目标植株表型 (Morph bias)2 × focal plant morph | 0.477 6 | 0.596 5 | 1.514 4 | 0.200 6 | |
型偏×目标植株表型 Morph bias × focal plant morph | 0.054 3 | 0.914 7 | 0.178 0 | 0.788 4 | |
目标植株的花数 No. flowers in focal plant | 0.026 9 | 0.000 4 *** | 0.019 0 | 0.112 0 | |
结籽数/株 Seeds/plant | |||||
(型偏)2 (Morph bias)2 | -0.362 3 | 0.597 6 | -2.038 0 | 0.104 9 | |
型偏 Morph bias | -0.169 7 | 0.671 1 | -0.634 9 | 0.380 7 | |
目标植株的表型 Focal plant morph | 0.121 8 | 0.574 9 | -0.469 0 | 0.163 7 | |
邻域密度 Neighborhood density | 0.000 9 | 0.909 2 | -0.002 1 | 0.896 7 | |
(型偏)2 ×目标植株表型 (Morph bias)2 × focal plant morph | 0.076 3 | 0.933 2 | 1.946 6 | 0.176 6 | |
型偏×目标植株表型 Morph bias × focal plant morph | 0.134 0 | 0.794 0 | 0.463 4 | 0.573 6 | |
目标植株的花数 No. flowers in focal plant | 0.025 2 | 0.001 0 *** | 0.032 4 | 0.018 6 * |
图4 种群中抽样植株功能性别的累积频度图。 M, Gj= 0, 个体只有雄性功能; F, Gj = 1, 个体只有雌性功能。实线和虚线分别表示忽略自交和存在自交。
Fig. 4 The cumulative frequency distribution of functional gender for plants sampled from the population. M, Gj = 0, for pure males; F, Gj = 1 for pure females. Values of the functional gender were calculated both assuming negligible selfing (solid) and using selfing rates (dotted).
表型 Morph | 2007 | 2008 | |||||
---|---|---|---|---|---|---|---|
下垂型 Cata-M | 上举型 Ana-M | p | 下垂型 Cata-M | 上举型 Ana-M | p | ||
花序花数/株 Flowers/plant | 46.89 ± 9.54 (69) | 47.84 ± 10.13 (58) | 0.59 | 41.24 ± 10.97 (89) | 39.80 ± 12.90 (81) | 0.43 | |
花展示 Display | 2.11 ± 0.46 (38) | 2.01 ± 0.45 (33) | 0.59 | 2.67 ± 0.34 (41) | 2.47 ± 0.26 (38) | 0.13 | |
花粉数/型/日 Pollen grains/morph/day | 78 300 ± 17 500 (38) | 82 000 ± 21 900 (33) | 0.63 | - | - | ||
胚珠数/型/日 Ovules/morph /day | 79 ± 20 (38) | 76 ± 18 (33) | 0.78 | - | - | ||
种子数/花 Seeds/flower | 8.24 ± 5.02 (31) | 9.56 ± 4.68 (25) | 0.31 | 3.54 ± 3.32 (45) | 2.85 ± 3.80 (44) | 0.37 | |
坐果数/株 Fruits/plant | 14.00 ± 7.56 (31) | 12.29 ± 8.51 (25) | 0.43 | 6.51 ± 5.95 (45) | 5.16 ± 5.78 (44) | 0.28 | |
结籽数/株 Seeds/plant | 378.81 ± 252.00 (31) | 430.84 ± 238.50 (25) | 0.43 | 159.84 ± 147.50 (45) | 131.86 ± 175.85 (44) | 0.45 |
表2 云南草蔻两型繁殖特征(平均值±标准偏差)
Table 2 Reproductive character means for anaflexistylous (Ana-M) and cataflexistylous (Cata-M) morphs of Alpinia blepharaocaly (mean ± SD)
表型 Morph | 2007 | 2008 | |||||
---|---|---|---|---|---|---|---|
下垂型 Cata-M | 上举型 Ana-M | p | 下垂型 Cata-M | 上举型 Ana-M | p | ||
花序花数/株 Flowers/plant | 46.89 ± 9.54 (69) | 47.84 ± 10.13 (58) | 0.59 | 41.24 ± 10.97 (89) | 39.80 ± 12.90 (81) | 0.43 | |
花展示 Display | 2.11 ± 0.46 (38) | 2.01 ± 0.45 (33) | 0.59 | 2.67 ± 0.34 (41) | 2.47 ± 0.26 (38) | 0.13 | |
花粉数/型/日 Pollen grains/morph/day | 78 300 ± 17 500 (38) | 82 000 ± 21 900 (33) | 0.63 | - | - | ||
胚珠数/型/日 Ovules/morph /day | 79 ± 20 (38) | 76 ± 18 (33) | 0.78 | - | - | ||
种子数/花 Seeds/flower | 8.24 ± 5.02 (31) | 9.56 ± 4.68 (25) | 0.31 | 3.54 ± 3.32 (45) | 2.85 ± 3.80 (44) | 0.37 | |
坐果数/株 Fruits/plant | 14.00 ± 7.56 (31) | 12.29 ± 8.51 (25) | 0.43 | 6.51 ± 5.95 (45) | 5.16 ± 5.78 (44) | 0.28 | |
结籽数/株 Seeds/plant | 378.81 ± 252.00 (31) | 430.84 ± 238.50 (25) | 0.43 | 159.84 ± 147.50 (45) | 131.86 ± 175.85 (44) | 0.45 |
表型 Morph | 自交率 Selfing rate | 雄性适合度 Male fitness | 雌性适合度 Female fitness | 总适合度 Total fitness |
---|---|---|---|---|
下垂型 Cata-M | 0.385 | 0.485 (0.465) | 0.415 (0.415) | 0.447 (0.435) |
上举型 Ana-M | 0.249 | 0.659 (0.441) | 0.409 (0.369) | 0.514 (0.397) |
表3 2007年两型雄性、雌性和总适合度
Table 3 Male, female and total prospective fitness for the morphs in 2007
表型 Morph | 自交率 Selfing rate | 雄性适合度 Male fitness | 雌性适合度 Female fitness | 总适合度 Total fitness |
---|---|---|---|---|
下垂型 Cata-M | 0.385 | 0.485 (0.465) | 0.415 (0.415) | 0.447 (0.435) |
上举型 Ana-M | 0.249 | 0.659 (0.441) | 0.409 (0.369) | 0.514 (0.397) |
表型 Morph | 表型频率 Morph frequency $\frac{{{n}_{x}}}{{{n}_{a}}+{{n}_{c}}}$ | 功能性别 Functional gender | |
---|---|---|---|
$\overline{{{d}_{x}}}$ | $\overline{{{G}_{x}}}$ | ||
2007 | |||
下垂型 Cata-M | 0.556 | 378.81 | 0.524 (0.565) |
上举型 Ana-M | 0.444 | 430.84 | 0.476 (0.574) |
2008 | |||
下垂型 Cata-M | 0.561 | 159.84 | 0.610 (0.615) |
上举型 Ana-M | 0.439 | 131.86 | 0.390 (0.468) |
表4 2007年和2008年上举型和下垂型植株功能性别($\overline{{{G}_{x}}}$)
Table 4 Functional gender ($\overline{{{G}_{x}}}$) of anaflexistylous and cataflexitylous plants for 2007 and 2008
表型 Morph | 表型频率 Morph frequency $\frac{{{n}_{x}}}{{{n}_{a}}+{{n}_{c}}}$ | 功能性别 Functional gender | |
---|---|---|---|
$\overline{{{d}_{x}}}$ | $\overline{{{G}_{x}}}$ | ||
2007 | |||
下垂型 Cata-M | 0.556 | 378.81 | 0.524 (0.565) |
上举型 Ana-M | 0.444 | 430.84 | 0.476 (0.574) |
2008 | |||
下垂型 Cata-M | 0.561 | 159.84 | 0.610 (0.615) |
上举型 Ana-M | 0.439 | 131.86 | 0.390 (0.468) |
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