山核桃雌配子体发育及其生殖过程研究
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摘要
山核桃(Carya carthayensis)是我国特有的优质干果和木本油料树种,主产于浙皖交界的天目山区。为进一步揭示山核桃的生殖奥秘,验证无融合生殖现象的存在,探讨其多胚发生来源与无融合生殖发生机制,提供山核桃育种的科学依据。本文从野外控制授粉和胚胎学角度,利用石蜡切片、荧光染色、扫描电子显微镜等技术手段,研究了山核桃雌配子体发育以及从传粉、花粉管萌发与生长、受精到胚胎发育的整个生殖过程。主要研究结论如下:
1、山核桃胚珠为直生胚珠,单珠被,厚珠心型。雌配子体为八核七细胞蓼形胚囊。雌配子体发育进程与雌花柱头颜色状态存在一定的联系。整个雌配子体发育进程时间较短。
2、山核桃在雌花柱头颜色微红-近紫黑色近10天时间内授粉均有坐果,但鲜红色状态下授粉坐果率最高,之后依次为紫红色、微红色、近紫黑色。套袋不授粉也有少量坐果,2,4-D处理坐果有所提高为4%。山核桃×薄壳山核桃的果实和种子明显增大,果皮色泽显绿,存在花粉直感现象,不授粉和激素处理收获果实和种子偏小。自然授粉下山核桃多胚苗比例在9.04%(统计586株),多胚苗能正常成苗。
3、采用13种培养基进行离体萌发试验均未见有花粉萌发,TTC法和醋酸洋红法可用于花粉活力粗略测定。原位萌发试验发现授粉后8h山核桃花粉在柱头上开始萌发,花粉管能正常生长。胚胎学显示5月中旬山核桃成熟胚囊阶段亦或受精后卵器有明显退化迹象,胚乳则正常发生。之后随着胚乳发育,胚珠膨大,珠心组织除珠心喙未被吸收外其余仅保留了表皮层细胞。多数珠心喙细胞体积增大,液泡化程度不一,近胚囊端部分细胞解体过程中,一部分细胞分裂形成细胞形状小、结构致密的珠心胚原始体。之后原始细胞旺盛分裂,6月底形成1个或多个块状细胞团,细胞团继续分裂伸长,呈现棒状,随着顶端增大,依次经历球型胚、心型胚、鱼雷胚、子叶胚阶段。7月下旬,子叶胚大小约2~3mm,此时山核桃内果皮开始变硬,胚乳逐渐被吸收,胚则快速发育,子叶裂瓣顶端分生组织快速生长充满子房腔,胚发育成熟。山核桃胚来源于珠心细胞,属于无融合生殖中不定胚生殖范畴。
Hickory (Carya cathayensis) is China’s unique high-quality dry fruit tree and woody oil plant. It mainly distributes at Tianmu mountain system matched between Zhejiang province and Anhui province. In order to reveal reproduction secrets of hickory, verify the existence of apomixis phenomenon, explore the origin of polyembryony and the occurrence mechanism in hickory, guide the breeding work, female gametophyte development and reproductive process from pollination, germination and growth of pollen tubes, fertilization to embryonic development were studied on the view of controlled pollination and embryology by means of paraffin section, fluorescence staining and scanning electron microscopy. Main results are as follows:
1. The ovule was unitegmic, orthotropous and crassinucellate in Hickory. Female gametophyte was polygonal embryo sac with 8 nucleis. There was corresponding relation between the developmental process of female gametophyte and the color status of stigmas. The whole developmental process of the female gametophyte was with very short time.
2. Female flowers pollinated among the time nearly 10 days from light red to light dark purple of the stigma color had fruit setting. The fruit setting rate was highest when female flowers were pollinated in bright red stage, followed by that when the stigma of female flower was in purplish red, light red and light dark purple. Fruits also were harvested under the treatment of bagging and without pollination and smearing the stigma with 2,4-D could promote the fruit setting with the fruit setting rate at 4%. Harvest fruits and seeds of Carya cathayensis×Carya illinoensis enlarged obviously and the exocarp color was more green compared with the control which indicated that there was metaxenia phenomenon. Harvest fruits and seeds under the treatment of bagging and without pollination and smearing with 2,4-D were relatively small. The proportion of the polyembryonic seedling in hickory was 9.04%( statistic 586 seedlings) in natural conditions and polyembryonic seedlings could develop and grow well.
3. There were no germinated pollens when pollen vigor testing was conducted using 13 different culture mediums. TTC and acetic red dyeing could be used to determine pollen viability inaccurately. Hickory pollen began germinating after pollinated 8 hours and the pollen tube grew normally through in situ germinating experiment. There were degradation signs of the egg apparatus and the endosperm developed normally at the mature embryo sac stage or after fertilization at the middle May. With the endosperm developing, the ovule enlarged and nucellar tissue were absorbed only remaining epidermis cells expect the nucellar beak region. Most nucellar cells in the nucellar beak region had increased volume with different degree of vacuolization. During the process of the disintegration of some nucellar cells near the embryo sac, some cells divided and formed nucellar embryo initials with small shaped cells and compact structure. One or more cell clusters formed at the micropylar end with the division of initials at the late June. Cell clusters developed and elongated and presented as rod-shape at first. The embryo experienced bulboid type embryo, heart embryo, torpedo embryo and cotyledon embryo in sequence. Endocarp began to shrink with the embryo 2~3mm at the late July, from then on endosperm was absorbed gradually and embryo developed rapidly. The growth and filling of the embryo depended on the development of meristem at the lower margin of the cotyledon lobes.The embryo in hickory origined from nucellar cells and it belonged to adventitious embryony reproduction one type of apomixis.
1、山核桃胚珠为直生胚珠,单珠被,厚珠心型。雌配子体为八核七细胞蓼形胚囊。雌配子体发育进程与雌花柱头颜色状态存在一定的联系。整个雌配子体发育进程时间较短。
2、山核桃在雌花柱头颜色微红-近紫黑色近10天时间内授粉均有坐果,但鲜红色状态下授粉坐果率最高,之后依次为紫红色、微红色、近紫黑色。套袋不授粉也有少量坐果,2,4-D处理坐果有所提高为4%。山核桃×薄壳山核桃的果实和种子明显增大,果皮色泽显绿,存在花粉直感现象,不授粉和激素处理收获果实和种子偏小。自然授粉下山核桃多胚苗比例在9.04%(统计586株),多胚苗能正常成苗。
3、采用13种培养基进行离体萌发试验均未见有花粉萌发,TTC法和醋酸洋红法可用于花粉活力粗略测定。原位萌发试验发现授粉后8h山核桃花粉在柱头上开始萌发,花粉管能正常生长。胚胎学显示5月中旬山核桃成熟胚囊阶段亦或受精后卵器有明显退化迹象,胚乳则正常发生。之后随着胚乳发育,胚珠膨大,珠心组织除珠心喙未被吸收外其余仅保留了表皮层细胞。多数珠心喙细胞体积增大,液泡化程度不一,近胚囊端部分细胞解体过程中,一部分细胞分裂形成细胞形状小、结构致密的珠心胚原始体。之后原始细胞旺盛分裂,6月底形成1个或多个块状细胞团,细胞团继续分裂伸长,呈现棒状,随着顶端增大,依次经历球型胚、心型胚、鱼雷胚、子叶胚阶段。7月下旬,子叶胚大小约2~3mm,此时山核桃内果皮开始变硬,胚乳逐渐被吸收,胚则快速发育,子叶裂瓣顶端分生组织快速生长充满子房腔,胚发育成熟。山核桃胚来源于珠心细胞,属于无融合生殖中不定胚生殖范畴。
Hickory (Carya cathayensis) is China’s unique high-quality dry fruit tree and woody oil plant. It mainly distributes at Tianmu mountain system matched between Zhejiang province and Anhui province. In order to reveal reproduction secrets of hickory, verify the existence of apomixis phenomenon, explore the origin of polyembryony and the occurrence mechanism in hickory, guide the breeding work, female gametophyte development and reproductive process from pollination, germination and growth of pollen tubes, fertilization to embryonic development were studied on the view of controlled pollination and embryology by means of paraffin section, fluorescence staining and scanning electron microscopy. Main results are as follows:
1. The ovule was unitegmic, orthotropous and crassinucellate in Hickory. Female gametophyte was polygonal embryo sac with 8 nucleis. There was corresponding relation between the developmental process of female gametophyte and the color status of stigmas. The whole developmental process of the female gametophyte was with very short time.
2. Female flowers pollinated among the time nearly 10 days from light red to light dark purple of the stigma color had fruit setting. The fruit setting rate was highest when female flowers were pollinated in bright red stage, followed by that when the stigma of female flower was in purplish red, light red and light dark purple. Fruits also were harvested under the treatment of bagging and without pollination and smearing the stigma with 2,4-D could promote the fruit setting with the fruit setting rate at 4%. Harvest fruits and seeds of Carya cathayensis×Carya illinoensis enlarged obviously and the exocarp color was more green compared with the control which indicated that there was metaxenia phenomenon. Harvest fruits and seeds under the treatment of bagging and without pollination and smearing with 2,4-D were relatively small. The proportion of the polyembryonic seedling in hickory was 9.04%( statistic 586 seedlings) in natural conditions and polyembryonic seedlings could develop and grow well.
3. There were no germinated pollens when pollen vigor testing was conducted using 13 different culture mediums. TTC and acetic red dyeing could be used to determine pollen viability inaccurately. Hickory pollen began germinating after pollinated 8 hours and the pollen tube grew normally through in situ germinating experiment. There were degradation signs of the egg apparatus and the endosperm developed normally at the mature embryo sac stage or after fertilization at the middle May. With the endosperm developing, the ovule enlarged and nucellar tissue were absorbed only remaining epidermis cells expect the nucellar beak region. Most nucellar cells in the nucellar beak region had increased volume with different degree of vacuolization. During the process of the disintegration of some nucellar cells near the embryo sac, some cells divided and formed nucellar embryo initials with small shaped cells and compact structure. One or more cell clusters formed at the micropylar end with the division of initials at the late June. Cell clusters developed and elongated and presented as rod-shape at first. The embryo experienced bulboid type embryo, heart embryo, torpedo embryo and cotyledon embryo in sequence. Endocarp began to shrink with the embryo 2~3mm at the late July, from then on endosperm was absorbed gradually and embryo developed rapidly. The growth and filling of the embryo depended on the development of meristem at the lower margin of the cotyledon lobes.The embryo in hickory origined from nucellar cells and it belonged to adventitious embryony reproduction one type of apomixis.
引文
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