姜科植物繁殖生态学研究
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摘要
姜科Zingiberaceae是一个有着50余属1,500多种的由动物传粉的热带单子叶大科,和姜科植物丰富的物种多样性和显著的花展示相比,长期以来对姜科植物繁殖生态学的研究相对较为缺乏,但这些相对有限的研究结果显示了姜科植物具有极其多样性的传粉和繁育系统,是研究有花植物传粉和繁育系统进化的理想植物类群。本研究在对姜科植物繁殖生态学研究进行全面总结和把握的基础上,选择具有不同花部特征的3属4种姜科植物,以花寿命的适应意义为关注重点,对这些植物的繁殖生态学进行研究,期望从不同角度来探讨姜科植物传粉和繁育系统的多样性及其形成机制。
大叶拟姜黄Curcumorpha longiflora为自交亲和植物,花寿命为2天,花在开放的第1天处于雄性阶段,第2天则为雌性阶段,花粉只能在不同花之间进行传递,从而完全避免了自花授粉的发生,同时,自然居群中的个体具有每隔1天开1朵花的特性,这使得在自然条件下同株异花授粉很少发生。大叶拟姜黄通过2天的花寿命来实现雄性先熟,同时配合特殊的开花格局来促进异交的机制是在姜科植物中的首次发现和报道。一种熊蜂Bombus sp.和小蜜蜂Apis florae是大叶拟姜黄的有效传粉者,它们对大叶拟姜黄不同性阶段的花有选择的访问是为了获取不同的回报物。
喙花姜Rhynchanthus beesianus具有典型的鸟传粉综合特征,花展示极为显著,但自然居群的结实率极低。在长达138个小时的访花动物观察中并没有发现喙花姜的有效传粉者,雌性黑胸太阳鸟Aethopyga saturata和一种熊蜂是喙花姜有规律的访花者,但它们都只是盗蜜者。人工授粉试验中,自交授粉的结实率(57.55±4.08%)和异交授粉的结实率(64.32±4.42%)没有显著差异,说明喙花姜是完全自交亲和的,同时自然居群存在着严重的花粉限制。生境片断化导致的鸟类绝灭可能使喙花姜的传粉者减少,是进而导致喙花姜居群结实率极低的主要原因。喙花姜的花寿命为2.5天,但其艳丽的苞片能保持鲜艳的红色4-5天,替代花瓣起着吸引传粉者的作用。
红姜花Hedychium coccineum为自交亲和植物,花序水平上的花展示极为显著,自然居群的结实同时存在着严重的花粉限制和资源限制。蝴蝶是红姜花主要的传粉者,其对花序的访问频率随花序总开花数的增加而显著增加。红姜花的花寿命为2天,花序上的每个蝎尾状聚伞小花序具3-4朵花,小花序内的花依次开放,小花序内不同轮次花的花粉数随轮次显著增加,而胚珠数则随轮次显著减少。红姜花在花序水平上的大量开花增加了对传粉者的吸引,同时有利于花粉的分散输出,增加花粉的供应,提高了红姜花个体的雄性适合度。
毛姜花Hedychium. villosum的2个变种四倍体毛姜花原变种var. villosum和二倍体小毛姜花var. tenuiflorum在形态特征上的差异主要体现在植株大小、花部大小及小花序数、花粉和胚珠数上,在花设计上没有任何差别。四倍体毛姜花原变种比二倍体小毛姜花具有较广的地理分布范围和多样化的生境适应性,两者在花期上已经产生了明显的生殖隔离。小毛姜花为合子后的自交不亲和,毛姜花原变种为完全自交亲和。蝴蝶和天蛾都是2个变种的有效传粉者,它们对2个变种具有同等的传粉效率,但2个变种在传粉者的种类上产生了强烈的分化。毛姜花原变种的花寿命为5天,而小毛姜花则仅有4天的花寿命。二倍体小毛姜花具有较强的无性繁殖能力,自然居群中个体的克隆大小和结实率之间具有显著的负相关性,而四倍体毛姜花原变种则不存在这样的相关性。多倍化过程使毛姜花的自交不亲和系统发生了崩溃,导致了四倍体毛姜花原变种和二倍体小毛姜花在繁殖特性上产生了一系列的差异,使得四倍体毛姜花原变种具有更强的生境适应能力,能分布到二倍体小毛姜花所不能到达的生境。多倍化可能在姜花属植物从热带到高海拔地区的扩散中起着重要的作用。
姜花属Hedychium是姜科唯一一个从热带到高海拔地区分布的大属,属内不同种类植物的花寿命差异很大。毛姜花原变种Hedychium villosum var. villosum的花寿命为5天,通过人工剪除毛姜花的柱头来使花朵具雌性功能的时间缩短,结果显示在不同的处理中,结实率随花朵具雌性功能时间的增长而增加,开花1天后剪除柱头的结实率为1.85±1.59%,极显著低于对照的结实率20.96±4.13%,说明其较长的花寿命能显著的增加传粉的成功率和结实率,长达5天的花寿命有利于雌性适合度的提高。同时,毛姜花原变种开花1天后剩余的平均花粉数和总平均花粉数没有显著差异,而开花后2、3、4天后剩余的平均花粉数则极显著少于总平均花粉数,说明随着花寿命的增加,花粉输出量也显著增加,其长达5天的花寿命也有利于其雄性适合度的提高。较长的花寿命在四倍体毛姜花原变种拓展新的生境,向更高海拔地区的扩散中也许起着重要的作用。
姜科植物有着极其多样性的传粉和繁育系统,花寿命在姜科植物传粉和繁育系统的多样性形成及其进化中起着重要作用,同时,较长的花寿命有效的提高了雌性适合度和雄性适合度,从而使姜科植物能脱离热带生境,向更高海拔的地区扩散,花寿命在姜科植物从热带地区到高海拔地区的分布过程中也起着重要作用。
The Zingiberaceae is a large family of animal-pollinated tropical monocotyledons with approximate 1,500 species in 50 genera. Members of this family display a broad range of pollination and breeding systems. Ginger plants are considered to be an ideal plant group for studying the evolution of pollination and breeding system of flowering plant. In this study, 4 ginger species in 3 genera with different floral characteristics are selected for studying the reproductive ecology with special focus on floral longevity.
Curcumorpha longiflora (Wallich) A. S. Rao & D. M. Verma is a perennial chasmophyte ginger that usually grows in crevices of calcareous rocks and forms patches on the understory of limestone monsoon rainforests. The pollination ecology of C. longiflora was studied by monitoring phenology and flowering behavior, observing pollinator activity, and the quantity and quality of pollination services. We also investigated the germination of pollen grains and growth of pollen tubes after different pollination treatments to detect its breeding system. Based on the results: (1) for the first time in Zingiberaceae a new protandrous mechanism was found with a two-day flowering to avoid autogamy in this species; (2) under field conditions, all individuals of C. longiflora usually produced only one flower every other day to keep geitonogamy to a minimum; (3) germination of pollen grains and growth rates of pollen tubes under different pollination treatments were the same 4 h later after pollination, suggesting that C. longiflora is completely self-compatible; (4) among the limited visitors, Bombus sp. and Apis florae were effective pollinators, but they were active at different times and at different stages of the flower, probably receiving different rewards.
Rhynchanthus beesianus W. W. Smith is an epiphytic tropical ginger with a very conspicuous floral display, but almost no fruit set under field conditions. The reproductive ecology encompassing phenology, floral biology, and pollination and breeding systems was investigated in an evergreen broadleaved forest in Yunnan Province, Southwest China. The flowers possess a typical bird pollination syndrome, but no effective pollinators were observed during 138 h of observation. Female Black-breasted Sunbird (Aethopyga saturata) and bumblebees visited R. beesianus regularly, but they all played roles as nectar robbers. No fruit was found in the bagging treatment, and fruit set following manual self-pollination (57.55±4.08%) was comparable with cross-pollination (64.32±4.42%), suggesting that R. beesianus is self-compatible but spontaneous self-pollination in this species does not occur. R. beesianus was dependent on animals for fertilization and suffered a serious pollinator-limitation. Bird extinctions as a result of habitat fragmentation may lead to a decline of pollination and fruit set in R. beesianus. The floral longevity of R. beesianus is 2.5 days, but its showy red bracts last for 4-5 days as attractant to pollinators.
Hedychium coccineum Smith is a self-compatible ginger with very conspicuous floral display at the inflorescence level, but fruit set in natural population suffered both serious pollen-limitation and resource-limitation. Butterflies are main pollinators of H. coccineum. In our observation, the visit frequency of 3 butterflies to H. coccineum increased greatly following the increase of total flowers on an inflorescence. The floral longevity of H. coccineum is 2 days. A cincinnus contains 3-4 flowers, and flowers bloom in turn. In cincinnus, the pollen grains of different flowers increase significantly in ture, but ovules numbers decline significantly in ture. Large floral display at the inflorescence level is advantageous both to attract pollinators and increase male fittness in H. coccineum.
The two varieties of Hedychium villosum Wallich in Roxburgh, diploidy var. tenuiflorum and tetraploidy var. villosum, are morphologically different in palnt size, floral size and numbers of pollen grains, ovules and cincinnus. There is no any difference in floral design between two varieties. Tetraploidy var. villosum is much widespread than diploidy var. tenuiflorum geographically and ecologically, and the reproductive isolation occur between two varieties through flowering season. Diploidy var. tenuiflorum is post-zygotic self-incompatible and tetraploidy var. villosum is completely self-compatible. Both butterflies and hawkmothes are effective pollinators of two varieties, but strong differentiation in butterfly and hawkmoth species occurred to two varieties. The floral longevity of var. tenuiflorum and var. villosum are 4 days and 5 days respectively. Diploidy var. tenuiflorum shows stronger clonal ability than tetraploidy var. villosum does, and the fruit-set of var. tenuiflorum is significantly negative relative to clone size in natural population. The breakdown of self-incompatibility associated with polyploidy in H. villosum lead to changes in flowering time, pollinators, sexual versus asexual reproduction between diploidy var. tenuiflorum and tetraploidy var. villosum, which may cause tetraploidy var. villosum can explore new habitation and widespread. Polyploidy may play an very important role in distribution of Hedychium specieas from tropical area to high elevation area. Hedychium J. Konig is the only large genera distributed from tropical area to high elevation area in Zingiberaceae. Members of this genera vary greatly in floral longevity. The floral longevity of Hedychium villosum var. villosum is 5 days which is much longer than other sympatric gingers. We reduced the functional floral longevity for female function by clipping stigmas to test whether the long flowering duration of var. villosum increase female fitness by measure the fruit-set of different treatments.
The results show that fruit-set incresed following the increse of female functional duration. Flowers with reduced functional floral longevity to 1 day had only 1.85±1.59% fruit-set which is significantly lower than 20.96±4.13% fruit-set of control flowers with natural longevity. Meanwhile, the mean pollen grains remiand on anthers of flowers 1 day after anthesis has no significant difference with the mean pollen grains of unopened flowers, but the mean pollen grains remiand on anthers of flowers 2, 3, 4 days after anthesis are all significantlly lower than the mean pollen grains of unopened flowers. These results indicate that the long floral longevity is advantageous both to male and female fitness in var. villosum. Long floral longevity may contribute to exploring new habitation and spreading toward high elevation area in tetraploidy var. villosum.
Floral longivity not only plays a key role in form of diversity and evolution of pollination and breeding system of ginger famlily, but also plays an very important role in exploring new habitation and spreading toward high elevation area from tropical area in ginger famlily.
大叶拟姜黄Curcumorpha longiflora为自交亲和植物,花寿命为2天,花在开放的第1天处于雄性阶段,第2天则为雌性阶段,花粉只能在不同花之间进行传递,从而完全避免了自花授粉的发生,同时,自然居群中的个体具有每隔1天开1朵花的特性,这使得在自然条件下同株异花授粉很少发生。大叶拟姜黄通过2天的花寿命来实现雄性先熟,同时配合特殊的开花格局来促进异交的机制是在姜科植物中的首次发现和报道。一种熊蜂Bombus sp.和小蜜蜂Apis florae是大叶拟姜黄的有效传粉者,它们对大叶拟姜黄不同性阶段的花有选择的访问是为了获取不同的回报物。
喙花姜Rhynchanthus beesianus具有典型的鸟传粉综合特征,花展示极为显著,但自然居群的结实率极低。在长达138个小时的访花动物观察中并没有发现喙花姜的有效传粉者,雌性黑胸太阳鸟Aethopyga saturata和一种熊蜂是喙花姜有规律的访花者,但它们都只是盗蜜者。人工授粉试验中,自交授粉的结实率(57.55±4.08%)和异交授粉的结实率(64.32±4.42%)没有显著差异,说明喙花姜是完全自交亲和的,同时自然居群存在着严重的花粉限制。生境片断化导致的鸟类绝灭可能使喙花姜的传粉者减少,是进而导致喙花姜居群结实率极低的主要原因。喙花姜的花寿命为2.5天,但其艳丽的苞片能保持鲜艳的红色4-5天,替代花瓣起着吸引传粉者的作用。
红姜花Hedychium coccineum为自交亲和植物,花序水平上的花展示极为显著,自然居群的结实同时存在着严重的花粉限制和资源限制。蝴蝶是红姜花主要的传粉者,其对花序的访问频率随花序总开花数的增加而显著增加。红姜花的花寿命为2天,花序上的每个蝎尾状聚伞小花序具3-4朵花,小花序内的花依次开放,小花序内不同轮次花的花粉数随轮次显著增加,而胚珠数则随轮次显著减少。红姜花在花序水平上的大量开花增加了对传粉者的吸引,同时有利于花粉的分散输出,增加花粉的供应,提高了红姜花个体的雄性适合度。
毛姜花Hedychium. villosum的2个变种四倍体毛姜花原变种var. villosum和二倍体小毛姜花var. tenuiflorum在形态特征上的差异主要体现在植株大小、花部大小及小花序数、花粉和胚珠数上,在花设计上没有任何差别。四倍体毛姜花原变种比二倍体小毛姜花具有较广的地理分布范围和多样化的生境适应性,两者在花期上已经产生了明显的生殖隔离。小毛姜花为合子后的自交不亲和,毛姜花原变种为完全自交亲和。蝴蝶和天蛾都是2个变种的有效传粉者,它们对2个变种具有同等的传粉效率,但2个变种在传粉者的种类上产生了强烈的分化。毛姜花原变种的花寿命为5天,而小毛姜花则仅有4天的花寿命。二倍体小毛姜花具有较强的无性繁殖能力,自然居群中个体的克隆大小和结实率之间具有显著的负相关性,而四倍体毛姜花原变种则不存在这样的相关性。多倍化过程使毛姜花的自交不亲和系统发生了崩溃,导致了四倍体毛姜花原变种和二倍体小毛姜花在繁殖特性上产生了一系列的差异,使得四倍体毛姜花原变种具有更强的生境适应能力,能分布到二倍体小毛姜花所不能到达的生境。多倍化可能在姜花属植物从热带到高海拔地区的扩散中起着重要的作用。
姜花属Hedychium是姜科唯一一个从热带到高海拔地区分布的大属,属内不同种类植物的花寿命差异很大。毛姜花原变种Hedychium villosum var. villosum的花寿命为5天,通过人工剪除毛姜花的柱头来使花朵具雌性功能的时间缩短,结果显示在不同的处理中,结实率随花朵具雌性功能时间的增长而增加,开花1天后剪除柱头的结实率为1.85±1.59%,极显著低于对照的结实率20.96±4.13%,说明其较长的花寿命能显著的增加传粉的成功率和结实率,长达5天的花寿命有利于雌性适合度的提高。同时,毛姜花原变种开花1天后剩余的平均花粉数和总平均花粉数没有显著差异,而开花后2、3、4天后剩余的平均花粉数则极显著少于总平均花粉数,说明随着花寿命的增加,花粉输出量也显著增加,其长达5天的花寿命也有利于其雄性适合度的提高。较长的花寿命在四倍体毛姜花原变种拓展新的生境,向更高海拔地区的扩散中也许起着重要的作用。
姜科植物有着极其多样性的传粉和繁育系统,花寿命在姜科植物传粉和繁育系统的多样性形成及其进化中起着重要作用,同时,较长的花寿命有效的提高了雌性适合度和雄性适合度,从而使姜科植物能脱离热带生境,向更高海拔的地区扩散,花寿命在姜科植物从热带地区到高海拔地区的分布过程中也起着重要作用。
The Zingiberaceae is a large family of animal-pollinated tropical monocotyledons with approximate 1,500 species in 50 genera. Members of this family display a broad range of pollination and breeding systems. Ginger plants are considered to be an ideal plant group for studying the evolution of pollination and breeding system of flowering plant. In this study, 4 ginger species in 3 genera with different floral characteristics are selected for studying the reproductive ecology with special focus on floral longevity.
Curcumorpha longiflora (Wallich) A. S. Rao & D. M. Verma is a perennial chasmophyte ginger that usually grows in crevices of calcareous rocks and forms patches on the understory of limestone monsoon rainforests. The pollination ecology of C. longiflora was studied by monitoring phenology and flowering behavior, observing pollinator activity, and the quantity and quality of pollination services. We also investigated the germination of pollen grains and growth of pollen tubes after different pollination treatments to detect its breeding system. Based on the results: (1) for the first time in Zingiberaceae a new protandrous mechanism was found with a two-day flowering to avoid autogamy in this species; (2) under field conditions, all individuals of C. longiflora usually produced only one flower every other day to keep geitonogamy to a minimum; (3) germination of pollen grains and growth rates of pollen tubes under different pollination treatments were the same 4 h later after pollination, suggesting that C. longiflora is completely self-compatible; (4) among the limited visitors, Bombus sp. and Apis florae were effective pollinators, but they were active at different times and at different stages of the flower, probably receiving different rewards.
Rhynchanthus beesianus W. W. Smith is an epiphytic tropical ginger with a very conspicuous floral display, but almost no fruit set under field conditions. The reproductive ecology encompassing phenology, floral biology, and pollination and breeding systems was investigated in an evergreen broadleaved forest in Yunnan Province, Southwest China. The flowers possess a typical bird pollination syndrome, but no effective pollinators were observed during 138 h of observation. Female Black-breasted Sunbird (Aethopyga saturata) and bumblebees visited R. beesianus regularly, but they all played roles as nectar robbers. No fruit was found in the bagging treatment, and fruit set following manual self-pollination (57.55±4.08%) was comparable with cross-pollination (64.32±4.42%), suggesting that R. beesianus is self-compatible but spontaneous self-pollination in this species does not occur. R. beesianus was dependent on animals for fertilization and suffered a serious pollinator-limitation. Bird extinctions as a result of habitat fragmentation may lead to a decline of pollination and fruit set in R. beesianus. The floral longevity of R. beesianus is 2.5 days, but its showy red bracts last for 4-5 days as attractant to pollinators.
Hedychium coccineum Smith is a self-compatible ginger with very conspicuous floral display at the inflorescence level, but fruit set in natural population suffered both serious pollen-limitation and resource-limitation. Butterflies are main pollinators of H. coccineum. In our observation, the visit frequency of 3 butterflies to H. coccineum increased greatly following the increase of total flowers on an inflorescence. The floral longevity of H. coccineum is 2 days. A cincinnus contains 3-4 flowers, and flowers bloom in turn. In cincinnus, the pollen grains of different flowers increase significantly in ture, but ovules numbers decline significantly in ture. Large floral display at the inflorescence level is advantageous both to attract pollinators and increase male fittness in H. coccineum.
The two varieties of Hedychium villosum Wallich in Roxburgh, diploidy var. tenuiflorum and tetraploidy var. villosum, are morphologically different in palnt size, floral size and numbers of pollen grains, ovules and cincinnus. There is no any difference in floral design between two varieties. Tetraploidy var. villosum is much widespread than diploidy var. tenuiflorum geographically and ecologically, and the reproductive isolation occur between two varieties through flowering season. Diploidy var. tenuiflorum is post-zygotic self-incompatible and tetraploidy var. villosum is completely self-compatible. Both butterflies and hawkmothes are effective pollinators of two varieties, but strong differentiation in butterfly and hawkmoth species occurred to two varieties. The floral longevity of var. tenuiflorum and var. villosum are 4 days and 5 days respectively. Diploidy var. tenuiflorum shows stronger clonal ability than tetraploidy var. villosum does, and the fruit-set of var. tenuiflorum is significantly negative relative to clone size in natural population. The breakdown of self-incompatibility associated with polyploidy in H. villosum lead to changes in flowering time, pollinators, sexual versus asexual reproduction between diploidy var. tenuiflorum and tetraploidy var. villosum, which may cause tetraploidy var. villosum can explore new habitation and widespread. Polyploidy may play an very important role in distribution of Hedychium specieas from tropical area to high elevation area. Hedychium J. Konig is the only large genera distributed from tropical area to high elevation area in Zingiberaceae. Members of this genera vary greatly in floral longevity. The floral longevity of Hedychium villosum var. villosum is 5 days which is much longer than other sympatric gingers. We reduced the functional floral longevity for female function by clipping stigmas to test whether the long flowering duration of var. villosum increase female fitness by measure the fruit-set of different treatments.
The results show that fruit-set incresed following the increse of female functional duration. Flowers with reduced functional floral longevity to 1 day had only 1.85±1.59% fruit-set which is significantly lower than 20.96±4.13% fruit-set of control flowers with natural longevity. Meanwhile, the mean pollen grains remiand on anthers of flowers 1 day after anthesis has no significant difference with the mean pollen grains of unopened flowers, but the mean pollen grains remiand on anthers of flowers 2, 3, 4 days after anthesis are all significantlly lower than the mean pollen grains of unopened flowers. These results indicate that the long floral longevity is advantageous both to male and female fitness in var. villosum. Long floral longevity may contribute to exploring new habitation and spreading toward high elevation area in tetraploidy var. villosum.
Floral longivity not only plays a key role in form of diversity and evolution of pollination and breeding system of ginger famlily, but also plays an very important role in exploring new habitation and spreading toward high elevation area from tropical area in ginger famlily.
引文
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