金银忍冬（Lonicera maackii (Rupr.) Maxim.）和葱皮忍冬（Lonicera ferdinandii Franch.）均可在哈尔滨市等高寒地区露地越冬，并且通过观察发现两种忍冬在哈尔滨市的开花物候及果实物候明显不同，所以我们推测除抗寒力不同导致该现象外，两种忍冬花芽分化、大小孢子及雌雄配子体和胚的发育不同也可能导致了开花和果实物候的不同。经查阅文献发现葱皮忍冬和金银忍冬在高寒地区的花芽分化、大小孢子及雌雄配子体和胚的发育均未见报道，所以本文以葱皮忍冬和金银忍冬为实验材料，探讨两种忍冬在哈尔滨地区的花芽分化、大小孢子及雌雄配子体和胚发育的异同，有助于理解两种忍冬生长发育的基本规律，揭示两种忍冬开花及坐果的生物学原理。该论文采用石蜡切片技术观察两种忍冬花芽分化、大小孢子及雌雄配子体和胚的发育进程，并采用高效液相色谱法、蒽酮比色法、考马斯亮蓝-G250法等方法测定发育过程相关生理指标。主要研究结果如下：

（1）两种忍冬的芽均为混合芽，均于3月初开始萌动，3月末展叶，10月中旬落叶，展叶期至落叶期均持续约165 d。葱皮忍冬和金银忍冬的花期历时分别为15 d和23 d，葱皮忍冬花期较短。两种忍冬果实均呈现“青果期”、“变色期”和“红果期”；葱皮忍冬和金银忍冬果期历时分别为170 d和195 d，金银忍冬果期较长。葱皮忍冬果实内含有2-13粒种子，金银忍冬果实内含有1-5粒种子；葱皮忍冬种子的横纵径、厚度及千粒重分别为0.474 cm、0.670 cm、0.126 cm、11.610 g，金银忍冬分别为0.287 cm、0.445 cm、0.080 cm、4.857 g，葱皮忍冬均大于金银忍冬。葱皮与金银忍冬的发芽率分别为74.0%和83.0%，发芽势分别为56.0%和71.0%。

（2）两种忍冬花芽分化均分为顶端分生组织转化期、花萼原基形成期、花瓣原基形成期、雄蕊原基形成期和雌蕊原基形成期5个时期。葱皮忍冬花芽分化历时约40 d，为连续分化；金银忍冬历时约120 d，分化期间存在停滞现象。

（3）葱皮忍冬大孢子及雌配子体发育晚于小孢子及雄配子体，大孢子及雌配体子发育历时29 d，小孢子及雄配子体发育历时39 d。金银忍冬大孢子及雌配子体发育历时46 d，小孢子及雄配子体发育历时39 d。两种忍冬雄配子体发育均经历孢原细胞→造孢细胞→花粉囊壁→小孢子母细胞→小孢子，最后形成雄配子体；雌配子体发育均由胚珠原基→倒生胚→大孢子母细胞→功能大孢子→胚囊，最后形成雌配子体。

（4）两种忍冬胚的发育均经历了合子分裂、球形胚、心形胚、鱼雷形胚和成熟胚5个阶段。葱皮忍冬胚发育历时23 d；金银忍冬历时24 d，金银忍冬胚的发育较葱皮忍冬早。两种忍冬胚乳发育类型均为核型，且胚乳的发育早于胚的发育。

（5）在花芽分化过程中可溶性糖、淀粉、SOD等12项生理指标含量均发生显著变化，IAA、ZT等生理指标变化表明花瓣原基和雄蕊原基为两种忍冬花芽分化过程的关键发育期。在大小孢子及雌雄配子体发育中11项生理指标均发生显著变化；超氧化物歧化酶（SOD）、精胺（Spm）等指标变化表明花粉囊壁和花粉母细胞时期为葱皮忍冬小孢子及雄配子体的关键发育期，大孢子母细胞和功能大孢子时期为大孢子及雌配子体发育的关键发育期；花粉母细胞和四分体时期为金银忍冬小孢子及雄配子体的关键发育期，倒生胚时期为大孢子及雌配子体发育的关键发育期。胚的发育中Put、GA₃等多数生理指标均发生显著变化，表明心形胚和鱼雷胚时期为葱皮忍冬胚发育的关键发育期，心形胚为金银忍冬胚发育的关键发育期。

本研究首次分析两种忍冬不同发育阶段生理指标变化规律，在此基础上确定两种忍冬各发育阶段的关键发育期；首次解析了葱皮忍冬花芽分化、大小孢子及雌雄配子体和胚发育的过程及历时。

Both Lonicera maackii (Rupr.) Maxim. and Lonicera ferdinandii Franch. can overwinter in the open ground in the alpine areas such as Harbin City, etc., and we found that the flowering and fruiting phenology of the two species were obviously different in Harbin City, so we speculate that the differences in cold resistance may also lead to the differences in flowering and fruiting phenology of the two species. Therefore, we speculate that in addition to the differences in cold resistance, the differences in bud differentiation, size of spores, and development of male and female gametophytes and embryos between the two species may also lead to the differences in flowering and fruiting phenology. After checking the literature, we found that the differentiation of flower buds, the development of large and small spores, male and female gametophytes and embryos of Lonicera ferdinandii Franch. and Lonicera maackii (Rupr.) Maxim. have not been reported in the alpine region. Therefore, this paper takes Lonicera ferdinandii Franch. and Lonicera maackii (Rupr.) Maxim. as the experimental materials, and explores the similarities and differences of the development of flower buds, the development of large and small spores, male and female gametophytes and embryos of these two kinds of lonicerasias in Harbin, which can help to understand the basic law of the development of these two kinds of lonicerasias, and reveal the biological principles of flowering and fruit set of the two kinds of lonicerasias. The thesis used paraffin sections. In this thesis, the development of flower bud differentiation, large and small spores, male and female gametophytes and embryos of Lonicera ferdinandii Franch. and Lonicera maackii (Rupr.) Maxim.were observed by paraffin sections, and the physiological indexes of the developmental process were measured by high performance liquid chromatography, anthracene ketone colorimetry, and Caulmers Blue-G250 method.The main results are as follows：

(1) Two species of Lonicera had mixed buds and began budding in early March. They developed leaves in late March and became deciduous in mid-October. The period from leaf development to deciduous lasted about 165 days. The flowering period of Lonicera ferdinandii Franch was 15 days. In comparison, the flowering period of Lonicera maackii (Rupr.) Maxim lasted 23 days. It is worth noting that the flowering period of Lonicera ferdinandii Franch was shorter than that of Lonicera maackii (Rupr.) Maxim. The growth process of fruit in both Lonicera species is divided into three stages: the green fruit stage, the colour change stage, and the red fruit stage. Lonicera ferdinandii Franch bears fruit for approximately 170 days,, from early June to late November. The fruiting period of Lonicera maackii (Rupr.) Maxim lasts from late April to mid-November, which is approximately 195 days. It is worth noting that the fruiting period of Lonicera maackii (Rupr.) Maxim is longer than that of other species of Lonicera. The fruits of Lonicera maackii (Rupr.) Maxim have a thin skin and contain 1-5 seeds in the case of honeysuckle and 2-13 seeds in the case of Lonicera ferdinandii Franch. The dimensions and weight of 1,000 seeds of Lonicera ferdinandii Franch were 0.474 cm (transverse diameter), 0.670 cm (longitudinal diameter), 0.126 cm (thickness), and 11.610 g (weight). The corresponding dimensions and weight of 1,000 seeds of Lonicera maackii (Rupr.) Maxim were 0.287 cm (transverse diameter), 0.445 cm (longitudinal diameter), 0.080 cm (thickness), and 4.857 g (weight). The seeds of Lonicera maackii (Rupr.) Maxim were smaller than those of Lonicera ferdinandii Franch. The germination percentages for Lonicera ferdinandii Franch and Lonicera maackii (Rupr.) Maxim were 74.0% and 83.0%, respectively. The germination potential were 56.0% and 71.0%, respectively.

(2) The flower bud differentiation of both Lonicera was divided into five periods: apical meristem transformation, calyx primordium formation, petal primordium formation, stamen primordium formation and gynoecium primordium formation. The differentiation of flower buds in Lonicera ferdinandii Franch occurred over a period of approximately 40 days and was continuous. In contrast, the differentiation of flower buds in Lonicera maackii (Rupr.) Maxim occurred over a period of approximately 120 days, with a period of stagnation during the differentiation period.

(3) Lonicera ferdinandii Franch macrospores and female gametophytes developed later than microspores and male gametophytes. Microspores and male gametophytes developed over a 39-day period , while macrospores and female gametophytes developed over a 29-day period; The development of Lonicera maackii （Rupr.） Maxim spores, male and female gametophytes lasted 39 days; Two species of Lonicera undergo a similar process for the development of their gametophytes. The male gametophyte goes through sporogonium, sporopoietic cells, pollen sac wall, microsporangium, microspores, and finally becomes a male gametophyte. On the other hand, the female gametophyte develops from ovule primordium, inverted embryo, macrosporangium, functioning macrospores, and embryo sac, and finally becomes a female gametophyte.

(4) The embryos of both Lonicera species underwent five stages of development: syncytium division, spherical embryo, heart-shaped embryo, torpedo-shaped embryo, and mature embryo. Embryo development of Lonicera ferdinandii Franch lasted for 23 days. The experiment showed that the development of Lonicera maackii (Rupr.) Maxim embryos lasted for 24 days. The development of the embryo of Lonicera maackii (Rupr.) Maxim occurs earlier than that of Lonicera ferdinandii Franch.

(5) During the process of flower bud differentiation, 12 physiological indexes, including soluble sugar, starch, and SOD, underwent significant changes. The changes in physiological indexes such as IAA and ZT indicated that the petal primordium and stamen primordium were the key developmental stages in the flower bud differentiation process of the two species of Lonicera. Physiological parameters, other than growth hormone (IAA), underwent significant changes during the development of megasporogenesis, as well as male and female gametophytes. The levels of SOD and Spm suggest that the development of Lonicera ferdinandii Franch microspores and male gametophytes is critical during the period of pollen sac wall and pollen mother cell. Similarly, the development of macrospores and female gametophytes is critical during the period of macrosporangium and functional macrospore. The period of pollen mother cells and tetrads is crucial for the development of microspores and male gametophytes in Lonicera maackii (Rupr.) Maxim. Similarly, the period of inverted embryos is critical for the development of macrospores and female gametophytes. All physiological indices changed significantly during embryo development. Most indices, such as Put and GA₃, indicated that the heart-shaped and torpedo embryo stages were critical for the development of the Lonicera ferdinandii Franch embryo, while only the heart-shaped embryo stage was critical for the development of the Lonicera maackii (Rupr.) Maxim embryo.

This study is the first time to analyse the physiological indexes of two species of Lonicera at different developmental stages to determine their key developmental periods. For the first time, we have analysed the process and duration of flower bud differentiation, large and small spores, and the development of male and female gametophytes and embryos in Lonicera ferdinandii Franch.

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