植物生态学报 ›› 2017, Vol. 41 ›› Issue (11): 1190-1198.DOI: 10.17521/cjpe.2017.0153
收稿日期:
2017-06-14
接受日期:
2017-11-20
出版日期:
2017-11-10
发布日期:
2017-11-10
通讯作者:
李巧峡
基金资助:
Qiao-Xia LI*(), Xiao-Xia HUANG, Wen CHEN, Yong WANG, Kun SUN
Received:
2017-06-14
Accepted:
2017-11-20
Online:
2017-11-10
Published:
2017-11-10
Contact:
Qiao-Xia LI
摘要:
紫花地丁(Viola philippica)是具典型开放花与闭锁花混合繁育系统的植物, 其两型花的形态差异为花器官发育研究提供了极好的模型。该文以不同光周期下生长的或在不同光周期间调换的紫花地丁植株为研究对象, 基于形态解剖学方法, 研究了过渡闭锁花的形态结构、开放花到完全闭锁花或完全闭锁花到开放花的变化式样, 以及完全闭锁花与过渡闭锁花中不发育雄蕊与花瓣的位置效应。结果显示: 该物种在短日照与中日照下均有开放花与过渡闭锁花形成, 短日照下大多数花芽为开放花, 少数为过渡闭锁花, 中日照下大多数花芽为过渡闭锁花, 少数为开放花; 长日照下全为完全闭锁花。在过渡闭锁花芽中, 存在着一系列雄蕊与花瓣数目不同的过渡类型, 既有偏向开放花的5雄3瓣的过渡类型, 也有偏向完全闭锁花的2雄1瓣的过渡类型。其中, 前一种类型在短日照下的过渡闭锁花芽中所占比例较大, 而后一种类型在中日照下所占比例较大。另外, 过渡闭锁花与完全闭锁花的雄蕊与花瓣发育程度均存在一定的位置效应, 其花芽腹侧的下花瓣(对应于开放花的下花瓣)与相邻的2枚雄蕊普遍发育最好, 而后花瓣(相对于前花瓣)的发育与完全闭锁花的基本一样, 为器官原基状, 与后花瓣相邻的2枚雄蕊也普遍最小, 而且也最容易发育为无小孢子发生的膜质状结构或原基状结构。同时, 将具完全闭锁花的植株置于短日照下或将具开放花的植株置于长日照下一段时间后, 重新诱导的花芽经历一系列过渡闭锁花类型后发生花型的相互转变。因此, 不同光周期对紫花地丁两型花发育的影响是渐进的, 长日照会抑制部分花瓣与雄蕊的发育, 短日照能拮抗并破除长日照对雄蕊与花瓣的抑制。
李巧峡, 黄小霞, 陈纹, 王镛, 孙坤. 紫花地丁两型花相互转变过程中花部形态结构的变化式样研究. 植物生态学报, 2017, 41(11): 1190-1198. DOI: 10.17521/cjpe.2017.0153
Qiao-Xia LI, Xiao-Xia HUANG, Wen CHEN, Yong WANG, Kun SUN. Patterns of flower morphology and structural changes during interconversion between chasmogamous and cleistogamous flowers in Viola philippica. Chinese Journal of Plant Ecology, 2017, 41(11): 1190-1198. DOI: 10.17521/cjpe.2017.0153
图1 紫花地丁两型花的表型变异。A, 开放花。B, 完全闭锁花。C-J, 过渡闭锁花。K, 开放花花芽的横切面。L, 完全闭锁花花芽的横切面。M-O, 过渡闭锁花花芽的横切面。ca, 心皮; fi, 花丝; pe, 花瓣; se, 花萼; st, 雄蕊。A-J, 比例尺为500 μm; K, 比例尺为200 μm; L-O, 比例尺为100 μm。
Fig. 1 The phenotype variation of dimorphic flower in Viola philippica. A, Chasmogamous flower. B, Cleistogamous flower. C-J, Intermediate cleistogamous flower. K, The cross section of chasmogamous flower. L, The cross section of cleistogamous flower. M-O, The cross section of intermediate cleistogamous flower. ca, carpel; fi, filament; pe, petal; se, sepal; st, stamen. A-J, bar = 500 μm; K, bar = 200 μm; L-O, bar = 100 μm.
图2 紫花地丁两型花中雄蕊形态与数量的变异。A, 开放花中的5枚雄蕊。B-L, 过渡闭锁花中不同数目的雄蕊。M, 完全闭锁花中的2枚雄蕊。an, 花药; fi, 花丝; sc, 附属结构-雄蕊帽。比例尺为500 μm。
Fig. 2 The morphological and number variation of stamens in dimorphic flower of Viola philippica. A, The five stamens in chasmogamous flower. B-L, The different number of stamens in intermediate cleistogamous flower. M, The two stamens in cleistogamous flower. an, anther; fi, filament; sc, stamen cap, the membranous appendage structure. Bar = 500 μm.
光周期 Photoperiod (light/dark) | 开放花植株比率 The ratio of plant with CH flowers (%) | 5枚或4枚雄蕊、3个花瓣过渡闭锁花植株比率 The ratio of plant with inCL flowers of 5 or 4 stamens and 3 petals (%) | 3枚雄蕊、3个花瓣过渡闭锁花植株比率 The ratio of plant with inCL flowers of 3 stamens and 3 petals (%) | 2枚雄蕊、3个花瓣过渡闭锁花植株比率 The ratio of plant with inCL flowers of 2 stamens and 3 petals (%) | 2枚雄蕊、1个花瓣过渡闭锁花植株比率 The ratio of plant with inCL flowers of 2 stamens and 1 petal (%) | 完全闭锁花植株比率 The ratio of plant with CL flowers (%) |
---|---|---|---|---|---|---|
10 h/14 h | 81.93 ± 0.016d | 6.40± 0.011c | 4.73 ± 0.012c | 4.87 ± 0.006c | 2.10 ± 0.008b | 0.00 ± 0.000a |
12 h/12 h | 3.50 ± 0.004b | 4.70 ± 0.014b | 9.27 ± 0.009c | 23.80 ± 0.032d | 59.57 ± 0.008e | 0.00 ± 0.000a |
16 h/8 h | 0.00 ± 0.000a | 0.00 ± 0.000a | 0.00 ± 0.000a | 0.00 ± 0.000a | 0.00 ± 0.000a | 100.00 ± 0.000b |
表1 紫花地丁不同光周期对过渡闭锁花形态结构的影响(平均值±标准误差)
Table 1 The effect of different photoperiod on the morphological structure of intermediate cleistogamous flowers of Viola philippica (mean ± SE)
光周期 Photoperiod (light/dark) | 开放花植株比率 The ratio of plant with CH flowers (%) | 5枚或4枚雄蕊、3个花瓣过渡闭锁花植株比率 The ratio of plant with inCL flowers of 5 or 4 stamens and 3 petals (%) | 3枚雄蕊、3个花瓣过渡闭锁花植株比率 The ratio of plant with inCL flowers of 3 stamens and 3 petals (%) | 2枚雄蕊、3个花瓣过渡闭锁花植株比率 The ratio of plant with inCL flowers of 2 stamens and 3 petals (%) | 2枚雄蕊、1个花瓣过渡闭锁花植株比率 The ratio of plant with inCL flowers of 2 stamens and 1 petal (%) | 完全闭锁花植株比率 The ratio of plant with CL flowers (%) |
---|---|---|---|---|---|---|
10 h/14 h | 81.93 ± 0.016d | 6.40± 0.011c | 4.73 ± 0.012c | 4.87 ± 0.006c | 2.10 ± 0.008b | 0.00 ± 0.000a |
12 h/12 h | 3.50 ± 0.004b | 4.70 ± 0.014b | 9.27 ± 0.009c | 23.80 ± 0.032d | 59.57 ± 0.008e | 0.00 ± 0.000a |
16 h/8 h | 0.00 ± 0.000a | 0.00 ± 0.000a | 0.00 ± 0.000a | 0.00 ± 0.000a | 0.00 ± 0.000a | 100.00 ± 0.000b |
图3 紫花地丁开放花与完全闭锁花植株在一定光周期下的花型变化趋势。A-E, 具有开放花的植株置于16 h光照时间后花芽形态的变化趋势。A, 0天时的花芽形态。B, 10天时的花芽形态。C, 20天时的花芽形态。D, 30天时的花芽形态。E, 40天时的花芽形态。F-J, 具有完全闭锁花的植株置于10 h光照时间后花芽形态的变化趋势。F, 0天时的花芽形态。G, 20天时的花芽形态。H, 40天时的花芽形态。I, 60天时的花芽形态。J, 80天时的花芽形态。fi, 花丝; ne, 蜜腺体; pe, 花瓣; st, 雄蕊。比例尺500 μm。
Fig. 3 The variation trends of flowers type of the plants with chasmogamous or cleistogamous flowers under different photoperiod in Viola philippica. A-E, The variation trends of flowers type of the plants with chasmogamous flowers under 16 h daylight. A, The morphological structure of flowers at 0 days. B, The morphological structure of flowers at 10 days. C, The morphological structure of flowers at 20 days. D, The morphological structure of flowers at 30 days. E, The morphological structure of flowers at 40 days. F-J, The variation trends of flowers type of the plants with cleistogamous flowers under 10 h daylight. F, The morphological structure of flowers at 0 days. G, The morphological structure of flowers at 20 days. H, The morphological structure of flowers at 40 days. I, The morphological structure of flowers at 60 days. J, The morphological structure of flowers at 80 days. fi, filament; ne, nectar; pe, petal; st, stamen. Bar = 500 μm.
图4 不同光周期下紫花地丁两型花花器官大小的变化趋势。A, B, 开放花植株在16 h日照下新诱导花芽花器官大小的变化趋势。C, D, 完全闭锁花植株在10 h日照下新诱导花芽花器官大小的变化趋势。AnL, 花药的长度; FiL, 花丝的长度; LpeL, 下花瓣的长度; LpeW, 下花瓣的宽度。不同小写字母表示不同时间段下花器官大小之间存在显著差异(p < 0.05)。
Fig. 4 The variation trends of floral organ size in dimorphic flowers of Viola philippica under different photoperiod. A, B, The variation trends of flowers organ size in newly developed floral buds of the plants with chasmogamous flowers under 16 h daylight. C, D, the variation trends of flowers organ size in newly developed floral buds of the plants with cleistogamous flowers under 10 h daylight. AnL, the length of anther; FiL, the length of filament; LpeL, the length of lower petal; LpeW, the width of lower petal. The different lowercase letters indicated that there were significant differences in the size of floral organs as the time went on (p < 0.05).
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