【作者】 马玉心；

【导师】 蔡体久；

【作者基本信息】 东北林业大学 ， 生态学， 2007， 博士

【摘要】 目前传粉生态学已经成为居群生物学和进化生物学中的热门领域。该领域主要研究与传粉有关的生物学特性及其规律。传粉生态学是植物有性生殖研究的核心内容。传粉系统由花粉、柱头和传粉媒介3个部分组成和维系，被认为是被子植物多样化最重要的一个因素。传粉是种子植物受精的必然阶段。花粉的运动在很大程度上限定了植物个体间的基因流和群体的交配方式，从而影响后代的遗传组成和适合度。本文主要从形态解剖、传粉、胚胎发育、种群格局等几个方面阐述兴安鹿蹄草的生殖及传粉生态学的规律。形态解剖学的实验方法主要采用传统的石腊切片法；传粉生态的研究方法主要依靠野外调查、石蜡切片、电镜扫描等方法进行；胚胎发育研究主要以石蜡切片法为主；种群格局主要以样地调查，有样方法、最近临体法、均匀度法等。通过解剖学研究揭示出营养、繁殖器官的结构，进而说明与传粉生态的关系。通过传粉生态的研究说明兴安鹿蹄草繁育系统的规律、传粉昆虫及其传粉行为、开花动态、柱头动态、协同进化、雌雄异位、访花频率及花粉流等规律；通过胚胎发育研究说明兴安鹿蹄草的不育机制及雌雄配子体发育的特征；通过种群格局的研究阐明兴安鹿蹄草的空间格局分布规律，并尝试了均匀度理论在种群格局测度中的应用，说明不同种群格局与种群开花率的关系。具体研究结果如下：兴安鹿蹄草的茎属于根状茎，茎具有初生与次生结构，初生结构表现出维管束散生，次生结构中木质部具有2～3个年轮，叶片为等面叶，无栅栏组织与海面组织的分化，又表现出旱生性质即角质层发达叶脉隆起，机械组织发达等，从叶脉的解剖结构可以推断兴安鹿蹄草为C4植物。花粉为复合花粉，柱头为分裂型，具有柱头腺体，其类型为结构腺体。种子梭形，种皮分离而透明，胚胎发育不完全，胚乳包围胚，花柱为圆形，花柱道五角形，中轴胎座，胎座发达。兴安鹿蹄草自花不育，也无无融合生殖。自然结实率很低，只有10％，主要采取异花传粉的生殖对策，传粉昆虫主要为杂棘蝇、东北驼花天牛、花蚤、蜜蜂科的一些种类，东北驼花天牛主要取食花粉，杂棘蝇主要舔食柱头腺体分泌物，蜜蜂科的种类如熊蜂也主要吃花粉。其中东北驼花天牛、杂棘蝇在长期的进化与自然选择过程中，与兴安鹿蹄草形成了协同进化的关系。东北驼花天牛在一日内的访花频率高峰期在中午。杂棘蝇、蜜蜂科的种类一日内访花有两次高峰。由于受环境温度的影响，昆虫的访花时间段各不相同，蜜蜂科种类偏向于温度比较高的时段，所以其访花时段是花即将凋落的时候。东北驼花天牛喜欢温凉的气候，所以其访花的时段为开花的头几日。杂棘蝇活动的高峰正值兴安鹿蹄草的开花盛期。东北驼花天牛的访花距离较短，访花路线为“之”字形，或折线形，杂棘蝇的访花距离较长。东北驼花天牛与杂棘蝇的花粉流截然不同，前者花粉流向为放射状，即离标记植株越近的植株得到的标记花粉越多，离标记植株越远的植株，则得到的标记花粉越少。后者花粉流向比较分散，被标记的花粉在样地中各处都有分布。兴安鹿蹄草存在动态雌雄异位，包括花柱与雄蕊的位置变化；柱头的动态变化。前者具体过程包括6个阶段，意义在于避免雌雄互相干扰；后者具体过程包括5个时期，意义在于增强柱头对花粉的接授。兴安鹿蹄草的胚胎发育特征表现为胚珠倒生，薄珠心，双层珠被。造胞细胞直接发育成大孢子母细胞。胚囊的发育属于蓼型，反足细胞多数，短命。在大孢子形成、胚囊形成、胚胎形成时期都发生不同程度的败育现象，败育机制为多个大孢子母细胞分裂不同步；雌配子体发育不正常与受精作用不能正常进行有关系；胚乳发育不全也是造成胚胎后期败育的主要原因：环境因素对雌配子体的发育有重要影响。种子形成时胚胎发育不完全。种子具有由种皮所形成的长长的翼。花粉囊发育的特征表现为，药室的表皮增厚，代替药室内壁起到保护作用。中层2—3层细胞，在花粉成熟时中层消失。绒毡层为变型绒毡层，小孢子进行减数分裂时绒毡层细胞解体。小孢子母细胞发育为小孢子的过程为离心式发育，小孢子母细胞在同一药室内发育是不同步的。边缘部分造胞细胞退化消失。雄配子体发育为二细胞型。兴安鹿蹄草的胚胎发育不全，属于生育期短的短命植物类型，其败育的主要原因在于气候因素。在小尺度范围内分别利用样方法、最近临体法、均匀度法测度了兴安鹿蹄草的种群分布格局，三种方法的测度结果均说明兴安鹿蹄草的种群空间格局类型为随机分布，三种方法中均匀度法是首次应用于种群空间格局的测度，经检验，此种方法更能准确反映种群空间格局类型，因为它既不受种群样方大小的限制，也与种群密度无关，其计算结果优于样方法与最近临体法。由于其计算过程不复杂，在使用上比点格局取样法更具有实用性。经统计分析发现兴安鹿蹄草的种群聚集度指数与透光度成负相关，与土壤pH值大小成负相关，与土壤通气度大小及土壤容量成正相关。种群空间格局与种群的开花率及繁殖生态学密切相关。传粉生态学已经越来越受到人们的重视，传粉生态学的研究结果对揭示植物的繁殖规律具有重要意义，本文围绕传粉生态研究为核心，从不同角度阐述了其生殖生物学规律，并首次报道了兴安鹿蹄草不育性的规律，研究结果将对进一步搞清兴安鹿蹄草的生殖生态学规律提供基础资料。更多还原

【Abstract】 Recently Pollination Ecology had became a popular field in Population Biology and Evolutionary Biology, researching of biological characters and its laws in relation to pollination. Study on Pollination Biology is the core subject of Plant Sexual Reproduction. That Pollination system consisted of three parts including pollen, stigma and pollination medium that is the most important factor attributed to the diversity of Angiosperm. Since pollination is the necessary stage of fertilization among Spermatophyte, the movement of pollen restricted the individual gene floating and mating modes between colonies. Consequently, the movement of pollen affected hereditary combination and Adaptation.The rules of Reproduction and pollination ecology of Pyrola dahurica(H. Andr.) was mainly expatiated from shape dissection, pollination, Embryonic Development, colony pattern and other aspects in this thesis. The experiment of Shape anatomy was employed by the method of traditional paraffine-slicing while studying on pollination biology reckoned on fieldwork, paraffine-slicing, electron microscope scanning and other ways.At the same time embryology is mainly dependent on paraffine-slicing while Population pattern is researched through field investigation, quadrat method, nearest-neighbor method, evenness method and others. Being revealed structure of Vegetative and propagative organs by Anatomical method is for getting further to explain relationship between them and pollination ecology. It was made out that P. dahurica, rules of reproduction system, pollination insects and their pollination behaviour, blossom dynamic, stigma dynamic, co-evolution, herkogamy of Visiting frequency and pollen flow through the research of pollination biology of P. dahurica. Infertility mechnism and developmental characteristics of female and male gametophytes was illuminated through the research of Embryology of P. dahurica. And the law of spatial distribution patterns of P. dahurica was accquired following the population pattern researched. Trying on applying the evenness theory to measurement of the population pattern, we indicated the relationship between population pattern and the population fecundity. The Specific research results as followed:The stem of Pyrola dahurica(H. Andr.) Kom was rhizome, the stem was provided with the primary structure and secondary structure, vascular bundle was scattered in the primary structure, the Xylem had 2～3 annual ring. leaves was equilateral, no differentiation of sponge tissue and palisade tissue, had the characteristics of growing in drought area of developed corneum and mechanical tissue, upheavaling vein. It could be concluded that P. dahurica was a sort of Carbon-4 plant from anatomical structure of its leaf vein; Its pollen was combined, style with structural gland was the type of dissociation, The Condia was tetrahedron shape, each pollen cultured in the medium would germinate a pollen tube, while pollen in the stigma would grow into 1～2 germ tubes and where it germinated was uncertain. Seeds took on spindle shape, its testa was Separated and transparent, its embryo was ateliosis, endosperm besieged embryo, style was round, stylar canal was pentagon, advanced spermaphore was axile placenta.Pyrola dahurica(H. Andr.) Kom was self-sterile and apomixis traits. The seed setting percentage was very low(only10％), P. dahurica adopt cross-pollination as reproductive countermeasure, which pollination insects mainly included phaonia hybrida(Schnabl)lapping the secretion of its stigma, Piidonia puziloi(Soisky) ingesting its pollen, phaonia hybrida (Schnabl), and a few species of honeybee family, such as Bombus, eating its pollen.Co-evolution relation came into being in long-term evolution and natural selection among P. puziloi、phaonia hybrida(Schnabl) and P. dahurica. It was at noon that the peak of visiting frequence within one day of P. puziloi occurred, the peak of visiting frequence of phaonia hybrida(Schnabl) and a few species of honeybee family occurred twice one day. The time period of visiting flower was different owing to effect of the temperature of environment, a few species of honeybee family prefered to visiting at higher temperature when the flower was about to wither.P. puziloi favours warm and cool weather, therefore, Its visiting time was concluded before flowering. The active peak of phaonia hybrida(Schnabl) was just the flowering bloom period of P. dahurica. visiting route of P. puziloi took on "zigzag shape" and "broken line form", and its distance of calling on flower was shorter than that of phaonia hybrida(Schnabl). The pollen flow of P. puziloi and phaonia hybrida(Sehnabl) were completely different, the former pollen flow direction was mainly radial which the most of pollen was accquired due to its near distance from the marker plant, and the latter one was dispersing which the few of pollen was accquired due to its far distance from the marker plant. The accomplished pollen flow direction by the latter was more dispersing than the marked one in the sample area.Movement herkogamy was one of characteristics of Pyrola dahurica(H. Andr.) Kom including the change of their position on style and stamen and dynamic change of style by itself. The former consist of 6 stages in order to avoid the interference between male and female; the latter included 5 phase in order to enhance style’s acception to the pollen.Embryology characteristics of Pyrola dahurica(H. Andr.) Kom was anatropous, thin nucellus, biteginous, building cell directly developed into megaspore mother cell, The development of embryo sac belongs to polygonum type with majority of ephemeral antipodal cells, During the period of Spore, embryo sac and embryo formation abortion occurred in various degree, which total percentage was 90％. Abortion was mainly because of the division asynchrony of most of megaspore mother cell, Developmental abnormality of female gametophyte and spermatization failure, otherwise, the abortion was mainly caused of agenesis of endosperm in the late stage of embryo. Environment was important impact factor on the development of female gametophyte. Seeds embryo only develops partially in the process of seed formation, seed was in possess of a very long wing originated from testa.The developmental characteristics of the anther sac was the thickened Epidermis of anther chamber instead of the chamber inner-wall’s protection. The 2-3 layer interlayer disappeared when the pollen matured. The tapetum originated from cytoclastic tapetum cell when the disformed microspore of tapetum type makes meiosis. The development of microspore mother cell was asynchronous in the same chamber. Spore mother cell in the peripheral part degenerated and die away, male gametophytes developed into 2-cell. The embryonic development of P. dahurica did not all belong to the ephemeral plant of short growing period which abortion mainly depended on climatic factor.Distribution patterns of population of P. dahurica was measured respectively by quadrat method, nearest-neighbor method and evenness method in small scale measurement, the measurement result indicated that distribution patterns of population of P. dahurica belonged to random distribution by the three method, among which the evenness method was first applied to the measurement of spatial distribution patterns of population, it was accurater to measure the type of spatial pattern than quadrat method and nearest-neighbor method because of its neither size restriction of populations nor quadrat and being irrelative with population density with more excellent results in comparision with two others, it was more of practicability than Point pattern analysis in practice because of its no complicated computational process. It was indicated that index of patchiness of P. dahurica population was inverse ratio to transmittance and PH value of soil, proportional to total porosity of the soils and bulk density of soil. Therefore, It was found out that the spatial pattern of population was closely related to flowering percentage and breeding ecology.Pollination ecology has been paid more and more attention to, the research result of Pollination ecology was of importance for discovering the propagation law of plant. Pollination ecology of P. dahurica was regarded as the studying core and the reproductive biology of P. dahurica was elaborated at a different angles in this article. Because this was the first article reported about the sterility law of P. dahurica, The research results was expected to lay the foundations for further studying on Breeding Ecology of P. dahurica.更多还原