植物繁育系统研究的最新进展和评述

 植物繁育系统是当今进化生物学研究中最为活跃的领域。繁育系统是指代表所有影响后代遗传组成的有性特征的总和,主要包括花形态特征、花的开放式样、花各部位的寿命、传粉者种类和频率、自交亲和程度和交配系统,其中交配系统是核心。我们重点综述了植物 有性繁育系统研究中1）传粉模式的多样性,2）从历史发生角度利用系统发育方法检验花性状的演化和繁育系统的生态转变过程,3）近交衰退对植物生殖史的影响及其机制和4）混和交配系统的时空动态、维持机理以及进化趋势的最新研究结果。总结了繁育系统在研究花适应性、物种形成机制和濒危植物     

中国特有种——沙芥繁育系统和传粉生物学研究

运用定位观察、杂交指数、花粉-胚珠比、套袋试验和重力玻片法等方法,对沙芥自然居群和人工栽培种群的开花日动态、繁育系统和传粉习性进行了研究.结果显示:沙芥在自然居群和人工栽培种群开花日高峰分别在19:00和15:00～18:00.沙芥繁育系统为异花授粉植物.沙芥传粉媒介中风的飘移能力非常微弱;沙芥自然居群的访花昆虫有19种,分属于6个目;人工栽培种群的访花昆虫有11种,分属于6个目.意大利蜜蜂、熊蜂、黑带食蚜蝇和拟蜂食蚜蝇是两个居群共有的主要传粉昆虫,在自然居群熊蜂每天的访花频率呈双峰曲线,其中访花最高峰与开花日高峰基本重叠.     

火把花的繁殖生物学研究

火把花（Colquhounia coccinea?）表现出典型的鸟媒综合征,因此具有作为引鸟景观植物的开发潜力。以自然、人工生境的火把花居群为研究对象,通过观察和试验对其开花物候、花部综合特征、访花动物及其行为、繁育系统、种子萌发特性进行研究,以明确火把花的繁殖特性及对不同访花动物的吸引潜力。结果显示,火把花的整体花期持续约3个月,单花花期为9.6±0.6 d;花蜜较丰富且稀薄,具有较短的花冠管和己糖为主的花蜜糖组成;访花动物主要是中华蜜蜂和多种食蜜鸟类,尤其是短喙的泛化鸟类,且在非自然生境中仍然能吸引鸟类访花;完全自交亲和但需要传粉者才能完成授粉,不存在花粉限制;不同授粉处理种子的发芽能力无显著差异;中华蜜蜂能有效传粉,鸟类的传粉作用需进一步验证。综上所述,火把花可供观赏的时间很长,具有明显的吸引鸟类访花的能力,容易通过有性繁殖途径快速获得大量幼苗。     

长喙毛茛泽泻（泽泻科）的交配系统研究

对濒凶植物长喙毛茛泽泻的交配系统作了综合研究。结果理：该种的花粉粒和柱头较平滑,花内分泌细胞存在但欠发达,开花前花粉萌发率较高而开花１ｈ后花粉萌发能力急剧衰退,花粉／胚珠比较低,花部综合特征与其花前期闭花授粉的交配习性相适应。该种的自然居群及实验居群中均未观察到花间的虫媒传粉,仅有极小型昆虫起花内传粉作用,亦未检出风媒花粉流。     

安徽特有植物安徽羽叶报春的繁殖生物学研究

本文分别从物候、繁育结构和传粉等方面研究了国家三级保护植物、安徽特有种--安徽羽叶报春(Primula merrilliana)的繁殖生物学特征。研究结果表明: (1) 安徽羽叶报春是典型的形态和功能意义上的花柱二型的二年生植物, 其始苗期为9月, 一直持续到第二年6月底或7月初死亡, 各居群长柱花植株比短柱花植株开花略早3–5 d; (2) 安徽羽叶报春的散孔型花粉显示其在报春花属中有重要的系统意义和进化意义。同时, 短柱花花粉量比长柱花少, 但其花粉直径是后者的1.78倍左右; (3) 长花柱的雌蕊长度是短花柱的1.64倍, 而且其柱头上的乳头状突起比短花柱上的长而稀; (4) 3个研究居群的P/O比都不相同, 我们发现花粉量比胚珠数量有更大的变异, 这可能与生境有关; (5) 安徽羽叶报春为异花授粉植物, 蓟马属(Thrips)昆虫为其重要的传粉者, 其同型花授粉可育性远比异型花授粉的高; (6) 其长、短柱型种子没有明显的形态差异, 多属不等边七面体, 少为其他多面体。据研究, 在小居群中两种花柱类型植株的比例失衡, 生境的破坏和伴生种的消失可能是导致安徽羽叶报春濒危的重要原因。     

藜异型性种子后代植株盐响应生理机制

种子异型性是植物适应异质生境的重要策略,异型性对后代植株的影响值得深入研究。以具有种子异型性的荒漠植物藜为材料,测定了异型种子后代植株在不同浓度(0,50,300 mmol/L)Na Cl胁迫下的表型及与耐盐相关的生理指标变化,探讨了异型种子后代植株之间对盐胁迫生理响应机制的差异。结果显示,褐色种子后代植株在各浓度盐胁迫下的表型(株高、分枝数)均优于黑色种子。对其生理指标测定结果显示,褐色种子植株除电导率显著低于黑色种子植株外,氧化损伤指标(氧自由基、丙二醛含量)及抗氧化酶(SOD、CAT、POX)活性、抗氧化剂(Car、As A)含量、渗调物质(可溶性糖、脯氨酸、甜菜碱)含量等在两者之间无明显差异。研究结果暗示,藜异型性种子后代植株的早期生长表型在盐胁迫下所产生的差异可能是由种子胚的大小及萌发快慢差异所导致,而这种差异并未引起盐胁迫下两种植株抗氧化系统响应的差异。     

水生被子植物的繁育系统与进化

对植物繁育系统的多样化及其作用模式和机制的研究,是理解植物各类群进化的一个重要基础。这一观点,已被从事植物进化生物学研究的国内外学者普遍接受。本世纪三十年代基因流概念的引人,促使人们在进化研究中,对植物繁育系统的重要性进行了重新评价。在过去的半个多世纪,植物进化生物学也因之而面貌一新,并得到了飞速的发展。时至今日,对植物进化过程中与繁育系统密切相关的一些重要机制,已有了较为深刻的认识（如有性过程中的遗传重组与植物的变异、进而与居群进化之间的关系等）。遗憾的是,这些认识几乎都是建立在对陆生植物研究…     

珍稀濒危植物陕西羽叶报春的开花及传粉生物学研究

陕西羽叶报春为中国珍稀濒危植物,研究其开花物候、传粉者及访花行为、花粉活力、柱头可授性、繁育系统,以明确陕西羽叶报春的传粉特性及制濒机制。以勉县陕西羽叶报春野生居群为研究对象,通过野外观察和人工授粉实验首次对陕西羽叶报春的传粉生物学进行研究。结果表明:(1)陕西羽叶报春种群花期为2-4月中旬,单花花期一般为14~17 d;单花开花进程中花部形态有6个明显发育变化:全部被萼片包裹、花苞伸出并为深粉色、花朵打开为淡粉色、花冠为白色、花朵枯萎、花冠脱落花萼宿存。(2)陕西羽叶报春为异花授粉植物,蜜蜂属为其主要传粉者。(3)柱头在花粉未开裂前就具有可授性,可授性逐渐增强;花药在花瓣打开后的第3天才开始开裂、散粉,在第4天活力最高,为98.18%,此后逐渐减弱。(4)繁育系统检测显示,长、短柱型花的花粉/胚珠比分别为902.26和831.48,杂交指数为4;套袋实验显示,陕西羽叶报春异型交配可育,同型交配不可育。由此表明,陕西羽叶报春的繁育系统为异交型,自交不亲和,需要传粉者。综上所述,陕西羽叶报春传粉者种类少、自交不亲和、胚珠败育现象可能是导致陕西羽叶报春濒危的重要原因。     

银杏传粉生物学研究进展

银杏(Ginkgo biloba L.)是最古老裸子植物之一,其传粉受精过程表现出许多特殊的原始性状和特征,在种子植物的系统演化上具有重要研究价值;同时,银杏种实经济价值高,但栽培上由于授粉受精不良而导致落花落果现象严重,因此了解银杏传粉生物学特性对于其种实的优质高产具有实际意义.本文从银杏雌雄株开花物候学、雌雄花形态结构特征、花粉和胚珠发育进程、花粉和胚珠生物学特性、传粉机制及花粉在胚珠内萌发生长进程等方面,对银杏传粉生物学相关研究进行综述,为银杏的系统演化、授粉受精和种实发育研究提供参考.     

青藏高原东缘块茎堇菜鳞茎分配的个体大小依赖性

块茎堇菜(Viola tuberifera)为青藏高原特有两型闭锁花植物,属多年生草本,具独特的混和交配系统,既可通过早春开放花异花受精和夏季地上地下闭锁花自花受精有性繁殖,还可通过秋季新鳞茎无性繁殖产生后代。高山环境下,异花受精常因花粉限制而无法正常进行,自花受精和克隆繁殖成为保障植物种群正常繁衍的不二之选,而克隆繁殖更能在植株资源消耗最小的情况下保障子代的存活。该文以青藏高原东缘高寒草甸的混合繁育植物块茎堇菜为研究对象,探索其生长期内鳞茎分配的个体大小依赖性,以及植株如何权衡鳞茎的资源分配以适应个体大小的变化。结果表明：块茎堇菜生活史阶段的鳞茎分配具有个体大小依赖性,鳞茎分配与个体大小呈极显著负幂指数相关关系(P<0.01),个体越大,鳞茎分配越小;反之,个体越小,鳞茎分配越高。即块茎堇菜对鳞茎的资源投入受个体大小的制约,通过鳞茎分配比例的高低响应植株自身资源状况的变化,保障在高寒环境下植物种群的生存和繁衍。该研究结果为高山植物克隆繁殖的生活史进化提供了依据。     

Patterns of flower morphology and structural changes during interconversion between chasmogamous and cleistogamous flowers in Viola philippica北大核心CSCD

Aims: Viola philippica is a species with a typical chasmogamous-cleistogamous (CH-CL) mixed breeding system. It provides a flower model system to investigate floral organs development under different photoperiods. Morphological changes of intermediate cleistogamous (inCL) flowers have been observed, the trends in variation of changes from CH flowers to CL flowers or from CL flowers to CH flowers have been analyzed, the localized effects of poorly developed stamens and petals in CL and inCL flowers have been identified. This research provided morphology and structural changes with implication for the evolutionary significance of the dimorphic flower formation for further study in dimorphic flower development. Methods: We used methods of anatomy and structural analysis to observe the morphological structures of flowers under different photoperiods. Important findings: Photoperiod played an important role in the development of CH and CL flowers in V. philippica. Under short-day light and intermediate-day light, both CH and inCL flowers developed simultaneously. Most of the floral buds were CH flowers under a photoperiod of short-day light, but most of the floral buds were inCL flowers under mid-day light. Complete CL flowers formed under long-day lights. However, there were a series of transitional types in the number and morphology of stamens and petals among inCL flowers, including five stamens with three petals related to CH flowers and two stamens with one petal related to CL flowers. The former type was dominant under short-day light conditions, and the latter type was dominant under mid-day light. Further more, there were localized effects in stamen and petal development for CL and inCL flowers. The development of ventral lower petal (corresponding to the lower petal with spur of CH flower) and the adjacent two stamens in inCL flowers were best, and the back petal was similar to that of CL flowers, an organ primordium structure. The adjacent stamens with the back petals tended to be poorly developed. In extreme cases, these stamens in inCL flowers had no pollen sac, only a membranous appendage or even a primordium structure. When the plants with CL or CH flowers were placed under short-day light or long-day light, the newly induced flowers all showed a series of inCL flower types, finally the CL flowers transformed into CH flowers, and the CH flowers transformed into CL flowers. This result indicates the gradual effects of different photoperiods on dimorphic flowers development of V. philippica. A long photoperiod could inhibit the development of partial stamens and petals, and a short photoperiod could prevent the suppression of long-day light and promote the development of stamens and petals.     

Patterns of flower morphology and structural changes during interconversion between chasmogamous and cleistogamous flowers in Viola philippica

Aims Viola philippica is a species with a typical chasmogamous-cleistogamous (CH-CL) mixed breeding system. It provides a flower model system to investigate floral organs development under different photoperiods. Morphological changes of intermediate cleistogamous (inCL) flowers have been observed, the trends in variation of changes from CH flowers to CL flowers or from CL flowers to CH flowers have been analyzed, the localized effects of poorly developed stamens and petals in CL and inCL flowers have been identified. This research provided morphology and structural changes with implication for the evolutionary significance of the dimorphic flower formation for further study in dimorphic flower development.Methods We used methods of anatomy and structural analysis to observe the morphological structures of flowers under different photoperiods.Important findings Photoperiod played an important role in the development of CH and CL flowers in V. philippica. Under short-day light and intermediate-day light, both CH and inCL flowers developed simultaneously. Most of the floral buds were CH flowers under a photoperiod of short-day light, but most of the floral buds were inCL flowers under mid-day light. Complete CL flowers formed under long-day lights. However, there were a series of transitional types in the number and morphology of stamens and petals among inCL flowers, including five stamens with three petals related to CH flowers and two stamens with one petal related to CL flowers. The former type was dominant under short-day light conditions, and the latter type was dominant under mid-day light. Further more, there were localized effects in stamen and petal development for CL and inCL flowers. The development of ventral lower petal (corresponding to the lower petal with spur of CH flower) and the adjacent two stamens in inCL flowers were best, and the back petal was similar to that of CL flowers, an organ primordium structure. The adjacent stamens with the back petals tended to be poorly developed. In extreme cases, these stamens in inCL flowers had no pollen sac, only a membranous appendage or even a primordium structure. When the plants with CL or CH flowers were placed under short-day light or long-day light, the newly induced flowers all showed a series of inCL flower types, finally the CL flowers transformed into CH flowers, and the CH flowers transformed into CL flowers. This result indicates the gradual effects of different photoperiods on dimorphic flowers development of V. philippica. A long photoperiod could inhibit the development of partial stamens and petals, and a short photoperiod could prevent the suppression of long-day light and promote the development of stamens and petals.     

不同生境对蝴蝶花花部与果实特征的影响

通过对毛竹林(Bamboo forest,BF)与林缘旷地(Open area of forest edge,OAFE)两类生境蝴蝶花自然种群花部与果实(种子)特征及降雨干扰影响的研究,探讨不同生境中蝴蝶花花部特征适应性及有性各组分的差异。结果表明:(1)竹林生境相对于林缘旷地生境,蝴蝶花单花花冠的长、宽较大,子房、花部(除花柄)及单花总生物量较小,比花柄长较大;两类生境蝴蝶花花柄生物量与子房生物量呈正相关(P0.05),协方差分析表明,两类生境该直线回归的总体斜率间(F=18.420,P0.001)及总体截距间(F=3791.7,P0.001)均具有显著差异,竹林生境花柄生物量随子房生物量增大而增大的程度更强。竹林生境的蝴蝶花侧花花冠长与宽最高,竹林生境顶花次之,林缘旷地顶花与林缘旷地侧花最低;花部(除花柄)与全花重都表现为林缘旷地侧花最高,林缘旷地顶花次之,竹林侧花与竹林顶花最低。比花柄长随竹林侧花与竹林顶花-林缘旷地侧花-林缘旷地顶花依次降低;竹林顶花的花柄比率最高,竹林侧花与林缘旷地侧花最低。(2)林缘旷地生境中蝴蝶花的每花序花数、花序结果百分率、单花结果百分率、每结果花序果实数、每结果花序果实重与种子重及花期内每花序掉落花数都高于竹林生境。(3)林缘旷地生境大雨干扰的4个时段花朵掉落数显著高于竹林环境(P0.01)。不同生境花部形态结构特征表明其自身的生境适应性,林缘旷地生境蝴蝶花为抵御干扰及为获得有性繁殖成功,有性组分的投入更高。     

高山植物花形态特征对花温度积累的影响

高山生态系统的低温环境不利于植物繁殖器官的发育和传粉过程,因此能改善花内或花序温度的结构对植物繁殖具有积极意义。该研究利用红外测温仪测量了高寒草甸中64种植物的花温度和叶温度以计算花温度积累,并检测了系统发育、花大小、花对称方式和花大小对花温度积累的影响。结果表明:所测量的64种植物种类,花部温度均高于叶部温度。花温度积累的系统发育信号较弱,表明物种间花温度积累的差异可能主要与不同物种对温度的需求有关。花温度积累与花大小存在显著的正相关关系,且两侧对称花的温度积累比辐射对称花更为迅速,但花对称方式和花颜色对花温度积累并没有显著的影响。该研究结果表明花温度积累在不同植物间存在很大的差异,但这些差异对植物繁殖的实际贡献和意义仍需要进一步验证。     

Evolution of floral dimorphism in a cleistogamous annual, <Emphasis Type="Italic">Impatiens noli-tangere</Emphasis> L. occurring under different environmental conditions

The resource allocation models for the evolution of a mixed production of cleistogamous and chasmogamous flowers were tested in four natural populations of Impatiens noli-tangere L. When we incorporated the estimated parameter values into our basic model, under the assumption that the environmental conditions did not change through the reproductive season, the model predicted the obligate chasmogamy in all four populations. However, including the fluctuation of the fertility of chasmogamous flowers in the model, the seasonal switching of chasmogamous and cleistogamous flower production becomes an evolutionary stable strategy, which explains the reproductive behavior of the individuals in two populations. Only after the relationship between the geitonogamy of chasmogamous flowers and the number of chasmogamous flower production was considered in the model did the simultaneous production of both of the dimorphic flowers become an evolutionary stable strategy. Further, the complete model clarifies the reproductive behavior of the individuals in the other two populations.     

Relationship between floral tube length and nectar robbing in Duranta erecta L. (Verbenaceae)

Although nectar robbing is a common phenomenon in plant species with tubular flowers or flowers with nectar spurs, the potential effect of this illegitimate interaction on plant reproductive success has not received the deserved attention. In the present study, we analysed the functional relationship between flower morphology and nectar robbing, and examined the reproductive consequences of the interaction in a population of Duranta erecta (Verbenaceae) on the island of Cuba. The results show that nectar robbing is conducted by the carpenter bees Xylocopa cubaecola and affects up to 44% of flowers in the studied population. However, not all the flowers have the same probability of being robbed. The chance of flowers being robbed increases with flower length and flower diameter. Moreover, nectar robbing significantly decreases the chance that flowers will set fruit. Also, the impact of nectar robbing on the probability of flowers to set fruits is dependent on the plant. We suggest that nectar robbing may represent an opposite selective force that balances the selection for longer corollas often imposed by pollinators specializing in visiting tubular flowers. Such a relationship with nectar robbers would have obvious implications for the evolution of tubular or closed flowers. This preliminary finding deserves further research in light of the ecological and evolutionary consequences of nectar robbing in tubular flowers.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 392–398.     

Why do flowers close at night? Experiments with the Lesser celandine Ficaria verna Huds (Ranunculaceae)

Although the mechanisms of flower closure have been widely studied, the evolutionary processes underlying this phenomenon are largely unknown. Open flowers may face a trade‐off between successful pollination/pollen transfer on the one hand and destruction of reproductive organs caused by exogenous factors (e.g. rain) on the other. The cost of flower opening was investigated by treating flowers of the Lesser celandine both in the field and under laboratory conditions. No differences were found in anthesis and reproductive success between treated and untreated flowers in any of the experiments. However, treated flowers in the field experiment suffered from significantly higher predation rates, in all probability caused by slugs and roe deer. It is suggested that flower closure may decrease the rate of being eaten by some herbivores, particularly in early‐spring plants that flower at times of insufficient food availability.     

植物繁殖生态学的若干重要问题

发现模式,并根据过程解释模式,是科学研究的主要目标.在植物繁殖生态学领域内,人们已经总结了一些重要的模式,包括普遍的雌雄同体、分离的花果期、较低的结实率、大量的同时开花、胁迫生境更易出现的异交、生活史性状与交配系统的相互作用、花序内的花分化,以及花内的雄蕊分化(异型雄蕊)等.对这些模式如何从机理上予以解释,还需要植物繁殖生态学家的艰苦努力.本文着重强调了植物许多重要繁殖特征都可以通过植株、花序以及花各层次上的资源分配而得到很好的解释,尤其是可以在很大程度上协调其他不同侧重点研究得到的矛盾结论.在今后逐渐强调多因素、多尺度对花部特征与传粉系统的适应与进化的研究中,从直接决定花部特征与交配系统的资源分配出发,将是促进人们准确理解与预测花部特征适应与进化的一个重要研究方向.     

甜荞pin型花与thrum型花雌雄蕊发育的比较研究

为从形态上研究甜荞(Fagopyrum esculentum)二型花发育的差异性,该文运用石蜡切片技术对甜荞pin型花和thrum型花大、小孢子发生和雌、雄配子体的发育过程进行观察比较.结果表明:(1)甜荞2种花的雌蕊、雄蕊发育的细胞学特征有很高的相似性,具直生胚珠、双珠被、厚珠心;大孢子四分体直线型排列,合点端为功能...     

荒漠孑遗植物裸果木的开花物候特征

2014年和2015年连续2年对哈密和乌恰裸果木(Gymnocarpos przewalskii)自然种群的开花物候特征进行野外定点观测,并运用相对开花振幅、开花强度和开花同步性等参数对其开花物候特征进行分析。结果表明,2个种群每年均开1次花,4月下旬到5月上旬进入始花期,开花进程基本相似,均呈单峰曲线,具有较高的开花同步性,呈现"集中大量"开花模式;在个体水平上,始花期、终花期在种群和年份间略有差异;单花持续时间在年际间是稳定的,但在种群间有所不同,即乌恰种群的单花持续时间略长于哈密种群;不同裸果木种群的开花数与结籽数间均呈极显著正相关,开花同步性指数与相对开花强度间呈显著负相关;乌恰种群的始花期与开花数间呈负相关。裸果木个体开花振幅、开花同步指数及开花模式在种群和年份间的相似性很大程度上是由其在长期的地理变迁中形成的遗传特性决定的,而始花期和开花持续时间等开花物候特征的差异主要是由环境和气象因子(海拔、气温、降水及光照)差异造成。研究结果对于探讨该属植物的繁殖生物学特性及其保护具有重要意义。