被子植物翅果的多样性及演化
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摘要
翅果能够依靠风力进行传播,可能是被子植物快速散布和物种分化的一个重要因素。狭义的翅果是指果皮延伸成翅且不开裂的干果;广义的翅果则包括果皮、花被片或苞片形成果翅的所有果实。根据果翅形态及其生长方式的不同,广义的翅果可分为单侧翅果、周位翅果(圆翅果与蝶翅果)、棱翅果、披针翅果、翼状萼翅果、叶状苞翅果6种类型,其空中运动方式有自旋式(单侧翅果、翼状萼翅果)、波浪式(周位翅果、叶状苞翅果)、翻滚自旋式(周位翅果)、直升机式(披针翅果、翼状萼翅果)和滚筒式(棱翅果)。棱翅果与圆翅果在被子植物基部类群樟目就有发生,并同时出现在单子叶植物和双子叶植物中,可能是最早出现的翅果类型。翅果的演化过程呈现出果翅数量增加、果翅偏向单侧和果翅负荷(果实质量与果翅面积之比)降低的趋势,以利于适应较小的风并增加传播距离。果翅除了促进果实与种子的风力传播外,还具有物理防御、调节种子萌发和促进二次传播等作用。泛热带分布的金虎尾科有着极其丰富的翅果类型,与其多次跨洋长距离扩散密切相关,可以作为研究翅果适应与演化的一个模式类群。结合生态和演化-发育生物学方法,研究不同类型翅果在适应风力传播方面的差异、萼片或苞片发育成翅的分子与遗传机制、翅果不同类型的演化历史及其对被子植物物种多样性的影响等是今后值得探讨的重要问题。
Samara(winged fruit) can be dispersed easily by wind and may be a crucial factor for angiosperm spread and diversification. In a narrow sense, a samara is an indehiscent dry fruit with wing(s) developed from fruit pericarp, while in a broad sense samaras also include all winged fruits with wings developed from both pericarp and perianth or bracts. According to the wing shape and growth patterns of samaras, we divided samaras into six types, i.e. single-winged, lanceolate-winged, rib-winged, sepal-winged, bract-winged, and perigynous samaras. Perigynous samaras can be further classified into two forms, i.e. round-winged and butterfly-winged samaras. Accordingly, the aerodynamic behavior of samaras can be classified into five types, autogyro, rolling autogyro, undulator, helicopter, and tumbler. The rib-winged and round-winged samaras can be found in Laurales, a basal angiosperm, and may represent the primitive type of early samaras. In the derived clades, samaras evolved enlarged but unequal wings and decreased wing loading(the ratio of fruit weight to wing size), which is likely an adaptation to gentle wind and secondary dispersal through water or ground wind. The wings of some samaras(such as sepal-winged and bract-winged samaras) may have multiple functions including wind dispersal, physical defense for the seeds, and adjust seed germination strategy. The pantropical family Malpighiaceae is extraordinarily rich in samara types, which is likely related to its multiple inter-continent dispersal in history, which is known as "Malpighiaceae Route". Therefore, Malpighiaceae can be used as a model system for the studies on samara adaptation and evolution. We identified the following issues that deserve further examination in future studies using both ecological and evo-devo methods: 1) the adaption of different types of samaras in dispersal processes, 2) the molecular and developmental mechanism of sepal-and bract-wings, and 3) the evolution of samara types and their effects on angiosperm diversification.
Samara(winged fruit) can be dispersed easily by wind and may be a crucial factor for angiosperm spread and diversification. In a narrow sense, a samara is an indehiscent dry fruit with wing(s) developed from fruit pericarp, while in a broad sense samaras also include all winged fruits with wings developed from both pericarp and perianth or bracts. According to the wing shape and growth patterns of samaras, we divided samaras into six types, i.e. single-winged, lanceolate-winged, rib-winged, sepal-winged, bract-winged, and perigynous samaras. Perigynous samaras can be further classified into two forms, i.e. round-winged and butterfly-winged samaras. Accordingly, the aerodynamic behavior of samaras can be classified into five types, autogyro, rolling autogyro, undulator, helicopter, and tumbler. The rib-winged and round-winged samaras can be found in Laurales, a basal angiosperm, and may represent the primitive type of early samaras. In the derived clades, samaras evolved enlarged but unequal wings and decreased wing loading(the ratio of fruit weight to wing size), which is likely an adaptation to gentle wind and secondary dispersal through water or ground wind. The wings of some samaras(such as sepal-winged and bract-winged samaras) may have multiple functions including wind dispersal, physical defense for the seeds, and adjust seed germination strategy. The pantropical family Malpighiaceae is extraordinarily rich in samara types, which is likely related to its multiple inter-continent dispersal in history, which is known as "Malpighiaceae Route". Therefore, Malpighiaceae can be used as a model system for the studies on samara adaptation and evolution. We identified the following issues that deserve further examination in future studies using both ecological and evo-devo methods: 1) the adaption of different types of samaras in dispersal processes, 2) the molecular and developmental mechanism of sepal-and bract-wings, and 3) the evolution of samara types and their effects on angiosperm diversification.
引文
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Friis EM,Crane PR,Pedersen KR(2011).Early Flowers and Angiosperm Evolution.Cambridge University Press.Cambridge,UK.
Green DS(1980).The terminal velocity and dispersal of spinning samaras.American Journal of Botany,67,1218-1224.
Greene DF,Johnson EA(1990).The dispersal of winged fruits and seeds differing in autorotative behaviour.Canadian Journal of Botany,68,2693-2697.
Gunasekara N(2004).Phylogenetic and Molecular Dating Analyses of The Tropical Tree Family Dipterocarpaceae Based on Chloroplast mat K Nucleotide Sequence Data.Master degree dissertation,Concordia University,Montreal.
Harris JG,Harris MW(2001).Plant Identification Terminology:An Illustrated Glossary.Spring Lake Publishing,Spring Lake,Utah.
Herrera CM(1989).Seed dispersal by animals:A role in angiosperm diversification?The American Naturalist,133,309-322.
Hong DY,Barrie FR,Bell CD(2011).Valerianaceae.In:Wu ZY,Raven PH,Hong DY eds.Flora of China.Vol.19.Science Press&Missouri Botanical Garden Press,Beijing.
Huang YJ,Zhu H,Chen WY,Zhou ZK(2013).Intraspecific variation in samara morphology of Acer and its implication in fossil identification.Plant Diversity and Resources,35,295-302.[黄永江,朱海,陈文允,周浙昆(2013).槭属翅果种内形态变异性及对其化石鉴定的意义.植物分类与资源学报,35,295-302.]
Jurado E,Westoby M,Nelson D(1991).Diaspore weight,dispersal,growth form and perenniality of Central Australian plants.Journal of Ecology,79,811-830.
Lansdown RV(2006).The genus Callitriche(Callitrichaceae)in Asia.Novon,16,354-361.
Li AR,Bao BJ,Grabovskaya-Borodina AE,Hong SP,McNeill J,Mosyakin SL,Ohba H,Park CW(2003).Polygonaceae.In:Wu ZY,Raven PH,Hong DY eds.Flora of China.Vol.5.Science Press&Missouri Botanical Garden Press,Beijing.
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Li XW,Li J,Ashton PS(2007).Dipterocarpaceae.In:Wu ZY,Raven PH,Hong DY eds.Flora of China.Vol.13.Science Press&Missouri Botanical Garden Press,Beijing.
Lord J,Egan J,Clifford T,Jurado E,Leishman M,Williams D,Westoby M(1997).Larger seeds in tropical floras:Consistent patterns independent of growth form and dispersal mode.Journal of Biogeography,24,205-211.
Lu AM,Stone DE,Grauke LJ(1999).Juglandaceae.In:Wu ZY,Raven PH,Hong DY eds.Flora of China.Vol.4.Science Press&Missouri Botanical Garden Press,Beijing.
Ma JS,Zhang ZX,Liu QR,Peng H,Funston AM(2008).Celastraceae.In:Wu ZY,Raven PH,Hong DY eds.Flora of China.Vol.11.Science Press&Missouri Botanical Garden Press,Beijing.
Manchester SR,O’Leary EL(2010).Phylogenetic distribution and identification of fin-winged fruits.The Botanical Review,76,1-82.
Norberg RA(1973).Autorotation,self-stability,and structure of single-winged fruits and seeds(samaras)with comparative remarks on animal flight.Biological Review,48,561-596.
Peck CJ,Lersten NR(1991).Samara development of black maple(Acer saccharum ssp.nigrum)with emphasis on the wing.Canadian Journal of Botany,69,1349-1360.
Planchuelo G,Catalán P,Delgado JA(2016).Gone with the wind and the stream:Dispersal in the invasive species Ailanthus altissima.Acta Oecologica,73,31-37.
Qian ZN,Ren MX(2016).Floral evolution and pollination shifts of the“Malpighiaceae route”taxa,a classical model for biogeographical study.Biodiversity Science,24,95-101.[钱贞娜,任明迅(2016).“金虎尾路线”植物的花进化与传粉转变.生物多样性,24,95-101.]
Qin HN,Gilbert MG(2007).Elaeagnaceae.In:Wu ZY,Raven PH,Hong DY eds.Flora of China.Vol.13.Science Press&Missouri Botanical Garden Press,Beijing.
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