珍稀克隆植物北极花有性繁殖状态及传粉模式研究
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摘要
有性繁殖是克隆植物适应不稳定环境和物种地理迁移的重要手段.分布于欧洲的克隆植物北极花因其有性繁殖率过低而濒危,新疆位于北极花欧亚分布区的边缘居群,该居群有着怎样的有性繁殖状态与传粉模式仍不清楚.研究表明:1)北极花单花期4～5d,花开放时,花冠内壁具毛状附属物,交叉成网状覆盖在雄蕊上,随着毛状附属物的逐渐倒伏,二强雄蕊两两序次呈现;2)自然状态下,80%以上的柱头上有充足的花粉,且能正常萌发,但能够生长到子房的花粉管数量仅占总花粉数的1/3左右;3)北极花的访花者主要有蝇、食蚜蝇、独居蜂,访花频率为0.102 4±0.056 7次/花/h.但传粉距离很短,主要集中在0～20cm范围之内;4)结实初期北极花的结实率高达82.18±1.48%,而后期大量脱落,仅为34.85±4.18%.北极花的克隆构型、雌雄异位、单花及花粉的序次呈现等成为延长散粉期、促进异交的重要性状;但较短的传粉距离,使其仍然不能避免高比例的同株异花授粉,以及由此导致的雌、雄性适合度代价及低的有性繁殖率.
It's one of the most important methods for clonal plants to fit for unstable environments and species geographic migration through sexual reproduction. The phenomenon has been reported that Linnaea borealis has endangered due to the low sexual reproduction in European populations. The sexual reproduction and pollination mode of populations, a peripheral population of Eurasia distribution area, in Xinjiang locating northwestern China are not clear. So researches on floral syndrome, pollination mode and fruit-set were carried out. Results showed that: 1) The individual flowering lasted for 4-5 days with hair appendages crossing as a web on the corolla inner wall, and coving the stamens during opening. With the hairs lodging, the didynamous stamens appear in pairs sequentially. 2) About 80 % of stigmas owned pollen enough which could germinate easy on stigmas, but only 1/3 of pollen grains can continue to grow with pollen tubes accessing to the ovary. 3) Major visitors were flies, hoverflies and solitary bees with visiting frequency of 0.102 4±0.056 7 times/per flower/per hour. But the pollination distance was very short, which mainly focused on the range of 0-20 cm. 4) The fruit-set rate was high with 82.18±1.48% at the beginning of fruit set period, but falled drop to 34.85±4.18% at mature period due to the fruits falling off gradually. The important traits related to extending floral show and promotting cross-pollination, such as clonal architecture,herkogamy, gradual presentation of flower and pollen et al in Linnaea borealis, fell flat on account of the short pollination distance which leaded to high geitonogamy ratio, male/female fitness cost as well as low sexual reproduction ratio.
It's one of the most important methods for clonal plants to fit for unstable environments and species geographic migration through sexual reproduction. The phenomenon has been reported that Linnaea borealis has endangered due to the low sexual reproduction in European populations. The sexual reproduction and pollination mode of populations, a peripheral population of Eurasia distribution area, in Xinjiang locating northwestern China are not clear. So researches on floral syndrome, pollination mode and fruit-set were carried out. Results showed that: 1) The individual flowering lasted for 4-5 days with hair appendages crossing as a web on the corolla inner wall, and coving the stamens during opening. With the hairs lodging, the didynamous stamens appear in pairs sequentially. 2) About 80 % of stigmas owned pollen enough which could germinate easy on stigmas, but only 1/3 of pollen grains can continue to grow with pollen tubes accessing to the ovary. 3) Major visitors were flies, hoverflies and solitary bees with visiting frequency of 0.102 4±0.056 7 times/per flower/per hour. But the pollination distance was very short, which mainly focused on the range of 0-20 cm. 4) The fruit-set rate was high with 82.18±1.48% at the beginning of fruit set period, but falled drop to 34.85±4.18% at mature period due to the fruits falling off gradually. The important traits related to extending floral show and promotting cross-pollination, such as clonal architecture,herkogamy, gradual presentation of flower and pollen et al in Linnaea borealis, fell flat on account of the short pollination distance which leaded to high geitonogamy ratio, male/female fitness cost as well as low sexual reproduction ratio.
引文
[1] Eckert C G, Barrett S C H. The loss of size in clonal plants[J].Evolutionary Ecology, 2002, 15:501-520.
[2] Wilcock C C, Jenningse S B. Partner limitation and restoration of sexual reproduction in the clonal dwarf shrub Linnaea borealis L.(Caprifoliaceae)[J]. Protoplasma, 1999, 208:76-86.
[3] Araki K, Shimatani K, Ohara M. Floral distribution clonal structure and their effects on pollination success in a selfincompatible Convallaria keiskei population in northern Japan[J]. Plant Ecology, 2007, 189:175-186.
[4]董鸣.克隆植物生态学[M].北京:科学出版社,2011.
[5] Vallejo-Mar′?n M, Dorken ME, Barrett SCH. The ecological and evolutionary consequences of clonality for plant mating[J].Annual Review of Ecology, Evolution and Systematics, 2010, 41:193-213.
[6] Kudoh H, Dhibaike H, Whigham DF, et al. Genet structure and determinants of clonal structure in a temperate de-ciduous woodland herb Uvularia perfoloata[J]. Journal of Ecology, 1999, 87:244-257.
[7] Charpentier A. Consequences of clonal growth for plant mating[J]. Ecol, 2002, 15:521–30.
[8] Albert T, Raspe O, Jacquemart AL. Influence of clonal growth on selfing rate in Vacciniu myrtillus L.[J]. Plant Biology,2008, 10:643–49.
[9] Neiland M R M, Wilcock C C. Reproductive ecology and conservation of rare plant species in Scotland[J]. Botany Social Scotland News, 1997, 68:18-19.
[10]尹林克.新疆珍稀濒危植物[M].乌鲁木齐:新疆科学技术出版社,2006.
[11] Scobie A R, Wilcock C C. Limited mate availability decreases reproductive success of fragmented populations of Linnaea borealis a rare clonal self-incompatible plant[J]. Annals of Botany, 2009, 1103:835-846.
[12] Cruden RW. Pollen-Ovule ratios:a conservative indication of breeding system in flowering plants[J]. Evolution, 1997, 31:32-46.
[13]黄云兰,安秀峰,师东,等.新疆荒漠植物耳叶补血草的爆发式开花式样与传粉模式[J].生物多样,2012,20(3):368-375.
[14]吴凌云,黄双全.虫媒传粉植物荞麦的生物学特性与研究进展[J].生物多样,2018,26(4):396–405.
[15]张大勇.植物生活史进化与繁殖生态学[M].北京:科学出版社,2004.
[16]白伟宁,张大勇.雌雄异熟. In:(张大勇)ed.植物生活史进化与繁殖生态学[M/OL].北京:科学出版社,2004:284-301.
[17] Back AJ, Kron P, Stewart SC. Phenological regulation of opportunities for within-inflorescence geitonogamy in the clonal species, Iris versicolor(Iridaceae)[J]. American Journal of Botany, 1996, 83:1033-1040.
[18] Vange V. Breeding system and inbreeding depression in the clonal plant species Knautia arvensis(Dipsacaceae):Implications for survival in abandoned grassland[J]. Biological Conservation, 2002, 108:59-67.
[19]翟雅芯,郭艳萍,张爱勤.一种新的花柱多态现象白花丹科植物细裂补血草[J].新疆大学学报(自然科学版),2014,31(2):134-138+143.
[20] Snow AA, Grove KF. Protandry, a neuter phase, and unisexual umbels in a hermaphroditic, neotropical vine(Bomarea acutifolia, Alstroemeriaceae)[J]. American Journal of Botany, 1995, 82:741-744.
[21] Harder LD, Wilson WG. Floral evolution and male reproductive success:optimal dispensing schedules for pollen dispersal by animal-pollinated plants[J]. Evolutionary Ecology, 1994, 8:542-559.
[22]陆婷,谭敦炎.动物传粉植物花粉呈现时序的进化意义[J].生物多样,2007,15(6):673-679.
[23] FANG Q, HUANG SQ. Plant-pollinator interactions in a biodiverse meadow are rather stable and tight for 3 consecutive years[J]. Integrative Zoology, 2016, 11:199–206.
[24] Zhang A Q, Xiong Y Z, Huang S Q. Maintenance of self-incompatibility in peripheral populations of a circumboreal woodland subshrub[J]. AoB Plants, ID:plu063, 2014.
[25]黄双全.了解生态网络需要监测植物与传粉者的相互作用[J].生物多样,2018,26(5):429-432.
[26] Kudo G,Suzuki S. Flowering phenology of alpine plant communities along a gradient of snowmelt timing[J].Polar Bioscience,1999, 1:100-113.
[27]李照广,尹弯,钟问,等.新疆夏尔希里自然保护区山地草原带蝴蝶资源调查[J].新疆大学学报(自然科学版),2017,34(2):218-222,252.
[28] Kevan PG. Inset pollination of high arctic flowers[J]. Journal of Ecology, 1972, 60:831-847.
[29] Inouye DW, Pyke GH. Pollination in snowy mountains of Australia:comparisions with montane Colorado USA[J]. Australian Journal of Botany, 1988, 81:1091-1095.
[30] Totland. Pollination in alpine Norway:flower phenology, insect visitors, and visitation rates in two plant communities[J].Canadian Journal of Botany, 1993, 71:1072–1079.
[2] Wilcock C C, Jenningse S B. Partner limitation and restoration of sexual reproduction in the clonal dwarf shrub Linnaea borealis L.(Caprifoliaceae)[J]. Protoplasma, 1999, 208:76-86.
[3] Araki K, Shimatani K, Ohara M. Floral distribution clonal structure and their effects on pollination success in a selfincompatible Convallaria keiskei population in northern Japan[J]. Plant Ecology, 2007, 189:175-186.
[4]董鸣.克隆植物生态学[M].北京:科学出版社,2011.
[5] Vallejo-Mar′?n M, Dorken ME, Barrett SCH. The ecological and evolutionary consequences of clonality for plant mating[J].Annual Review of Ecology, Evolution and Systematics, 2010, 41:193-213.
[6] Kudoh H, Dhibaike H, Whigham DF, et al. Genet structure and determinants of clonal structure in a temperate de-ciduous woodland herb Uvularia perfoloata[J]. Journal of Ecology, 1999, 87:244-257.
[7] Charpentier A. Consequences of clonal growth for plant mating[J]. Ecol, 2002, 15:521–30.
[8] Albert T, Raspe O, Jacquemart AL. Influence of clonal growth on selfing rate in Vacciniu myrtillus L.[J]. Plant Biology,2008, 10:643–49.
[9] Neiland M R M, Wilcock C C. Reproductive ecology and conservation of rare plant species in Scotland[J]. Botany Social Scotland News, 1997, 68:18-19.
[10]尹林克.新疆珍稀濒危植物[M].乌鲁木齐:新疆科学技术出版社,2006.
[11] Scobie A R, Wilcock C C. Limited mate availability decreases reproductive success of fragmented populations of Linnaea borealis a rare clonal self-incompatible plant[J]. Annals of Botany, 2009, 1103:835-846.
[12] Cruden RW. Pollen-Ovule ratios:a conservative indication of breeding system in flowering plants[J]. Evolution, 1997, 31:32-46.
[13]黄云兰,安秀峰,师东,等.新疆荒漠植物耳叶补血草的爆发式开花式样与传粉模式[J].生物多样,2012,20(3):368-375.
[14]吴凌云,黄双全.虫媒传粉植物荞麦的生物学特性与研究进展[J].生物多样,2018,26(4):396–405.
[15]张大勇.植物生活史进化与繁殖生态学[M].北京:科学出版社,2004.
[16]白伟宁,张大勇.雌雄异熟. In:(张大勇)ed.植物生活史进化与繁殖生态学[M/OL].北京:科学出版社,2004:284-301.
[17] Back AJ, Kron P, Stewart SC. Phenological regulation of opportunities for within-inflorescence geitonogamy in the clonal species, Iris versicolor(Iridaceae)[J]. American Journal of Botany, 1996, 83:1033-1040.
[18] Vange V. Breeding system and inbreeding depression in the clonal plant species Knautia arvensis(Dipsacaceae):Implications for survival in abandoned grassland[J]. Biological Conservation, 2002, 108:59-67.
[19]翟雅芯,郭艳萍,张爱勤.一种新的花柱多态现象白花丹科植物细裂补血草[J].新疆大学学报(自然科学版),2014,31(2):134-138+143.
[20] Snow AA, Grove KF. Protandry, a neuter phase, and unisexual umbels in a hermaphroditic, neotropical vine(Bomarea acutifolia, Alstroemeriaceae)[J]. American Journal of Botany, 1995, 82:741-744.
[21] Harder LD, Wilson WG. Floral evolution and male reproductive success:optimal dispensing schedules for pollen dispersal by animal-pollinated plants[J]. Evolutionary Ecology, 1994, 8:542-559.
[22]陆婷,谭敦炎.动物传粉植物花粉呈现时序的进化意义[J].生物多样,2007,15(6):673-679.
[23] FANG Q, HUANG SQ. Plant-pollinator interactions in a biodiverse meadow are rather stable and tight for 3 consecutive years[J]. Integrative Zoology, 2016, 11:199–206.
[24] Zhang A Q, Xiong Y Z, Huang S Q. Maintenance of self-incompatibility in peripheral populations of a circumboreal woodland subshrub[J]. AoB Plants, ID:plu063, 2014.
[25]黄双全.了解生态网络需要监测植物与传粉者的相互作用[J].生物多样,2018,26(5):429-432.
[26] Kudo G,Suzuki S. Flowering phenology of alpine plant communities along a gradient of snowmelt timing[J].Polar Bioscience,1999, 1:100-113.
[27]李照广,尹弯,钟问,等.新疆夏尔希里自然保护区山地草原带蝴蝶资源调查[J].新疆大学学报(自然科学版),2017,34(2):218-222,252.
[28] Kevan PG. Inset pollination of high arctic flowers[J]. Journal of Ecology, 1972, 60:831-847.
[29] Inouye DW, Pyke GH. Pollination in snowy mountains of Australia:comparisions with montane Colorado USA[J]. Australian Journal of Botany, 1988, 81:1091-1095.
[30] Totland. Pollination in alpine Norway:flower phenology, insect visitors, and visitation rates in two plant communities[J].Canadian Journal of Botany, 1993, 71:1072–1079.