水稻胚乳发育中珠心凋亡与胚乳细胞构建的研究
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摘要
水稻是中国及亚洲地区主要的粮食作物之一，在粮食生产中占有举足轻重的地位。胚乳细胞的发育状况直接影响籽粒的产量和品质。本论文以籼稻品种：胜泰1号为材料，对水稻胚乳发育过程中胚乳细胞的构建，胚乳细胞中淀粉质体的发生发育以及胚乳发育中珠心细胞的细胞程序性死亡及它与胚乳发育的关系，进行结构和功能的研究。主要研究结果如下：
1．水稻胚乳细胞的构建 花后约两天，游离核胚乳开始细胞化。中央细胞壁产生钉状垂周内突，延伸入中央细胞腔。内质网线状排列，通过电子致密小泡相互连接；同时，小泡将内质网和壁内突融合，成为垂周壁，将合胞体的胚乳分隔成一个个向心开口的圆柱体。圆柱体内的核分裂，伴随产生成膜体，在成膜体的基础上融合小泡产生平周壁，形成一个个单核的细胞。平周壁的产生还有另一个来源，即通过垂周壁的游离末端分支相互融合而成。携带有壁物质的电子致密小泡可来源于高尔基体分沁的携带有壁物质的小泡；内质网自身囊腔膨大而缢缩形成的球状体；周围珠心细胞PCD降解后产生的，包含有降解物质的电子致密小泡，通过中央细胞壁内突而运输入中央细胞。还观察到细胞化过程中酸性磷酸酶(AcPase)的定位变化。在新形成的细胞壁上观察到强烈的AcPase的活性，而早已形成的细胞壁上几乎没有酶活。
2．胚乳细胞中淀粉质体的发生水稻淀粉粒属复粒淀粉，淀粉质体除来源于质体、淀粉质体外，还来源于线粒体。，花后约60小时，胚乳细胞中开始出现淀粉体。淀粉体可通过出芽和中间缢缩断裂两种方式进行增殖。用AcPase定位表明，完全淀粉化的胚乳细胞中，细胞核被淀粉质体挤压变形而解体，其余细胞器被吞噬或解体，直至整个胚乳细胞中只含有淀粉质体、蛋白质体和脂体，而表现为一种凋亡表形。不同发育时期稻米籽粒胚乳细胞中，淀粉粒大小的动态变化表明：花后3～12天是水稻胚乳细胞中淀粉粒的发育的高峰期，在该时段，淀
水稻胚乳发育中珠心凋亡和胚乳细胞构建的研究粉粒直径的增加几乎接近于直线增长。12天以后，淀粉粒的直径维持在一个接近恒定的水平。胚乳中央部分与周围部分的细胞中淀粉粒的形状有所不同。中央部分的多为多面体形，且排列紧密;而周围部分的淀粉粒多为圆球形，排列较为疏松 3.胚乳发育过程中，珠心细胞的细胞程序性死亡珠心细胞PCD始于大抱子时期。用超微细胞化学的方法，对珠心细胞PCD过程中，酸性磷酸酶(aeid phosphatase，Aepase)进行研究表明，随着pen进行的程度的不同，AcPase依次定位于核膜、核质、胞质、细胞质膜、液泡膜、内质网膜和线粒体膜，并伴随有典型的凋亡细胞表形。表现为:染色体固缩且周缘化，线粒体腔膨大，内质网槽库膨大，细胞质膜和细胞壁分离，出现大量吞噬泡，细胞器被降解或吞噬，最终产生凋亡小体，其中包含有降解的核质或胞质，胞壁破裂，凋亡小体释放。同时，还用线粒体标志酶一细胞色素氧化酶(cytochrome oxidase)对线粒体进行超微细胞化学的定位研究，证明了在PCD过程中确有线粒体晴模糊，腔膨大，线粒体膜破裂等线粒体解体现象的存在。观察表明珠心细胞的PCD降解产物对胚乳细胞构建的哺乳关系。
Rice(Oryza Sativa L.) is one of the main resources of cereal grown in China and even in the Asian. It plays a very important role in food supply. Endosperm is the most primary part of the rice grain that can be eaten, so the developmental pattern of endosperm influence the grain quantity and quantily directly. Starch is both the major component of the yield in the world's main crop plants and an important raw material for many industrial processes.
In this thesis, we report the structural and functional research about nucellar cells PCD, endosperm cells construction, and amyloplasts initiation and development during the development of the endosperm of the rice. The main research results were as followed.
1.Construction of the rice endosperm cells Cellularization of the free-nuclear endosperm was initiated about 2 days after pollination(DAP) with the form of the peg-like anticlinal ingrowths on the central cell wall. These ingrowths extended into the lumen of central cell and fused with the linear endoplasmic reticulum(ER) by many electron-dense vesicles. Periclinal walls were formed by two mechanisms: (1) formation of the phragmoplasts followed the kynokinesis, (2) fusion of the free-edges of the opposing anticlinal walls. It was speculated that the electron-dense vesicles had three resources. From the Golgi apparatus, ER or degenerated nucellar cells which were transported through the ingrowths of central cell wall. By the means of cytochemistry of ultrastructure, We observed the location change of the AcPase. During the proceeding of the cellularization, the new-formed cell walls had distinct aggradation of the AcPase, but on the cell walls which were formed a longer time ago, we could not observe the aggr
adation of the AcPase.
2. The resource and development of amyloplasts in the
endosperm cells The amyloplasts of rice were compound starch granules. We observed that the amyloplasts could be initiated not only from the plastids, proplastids and amyloplasts, but also from the mitochondria. They appeared in the endosperm cell after 60 hours after pollination, and propagated by the formation of gemma or by contraction in the middle before splitting. The ultrastructural location of AcPase suggested that the nucleus and other cell apparatus were degenerated in the mature endosperm cells, which were filled with amyloplasts and protein bodies. So they showed another kind of phenomena of PCD.
The compare of the starch granules of different stage of development showed that the 3-12 days after pollination was the fastigium of the development of the starch granules. During this period, the augmentation of the diameter of the starch granules was approximate the pattern of beeline. But after 12 DAP, the diameter of the starch granules maintained at a invariable level. The shape of the starch granules in the medial region of the endosperm was different from the ambient region of that. Most of the starch granules in the medial region of the endosperm was polyhedral, crystalline and compact arranged; but the starch granules in the ambient region were almost ball shape, loose arrangement.
3. The proceeding of Programmed Cell Death (PCD) of nucellar cells during the development of the endosperm PCD of nucellar cells initiated in 2 days after pollination (2DAP). By the means of ultrastructure cytochemistry, we researched the variation of AcPase (Acid phosphatase) in nucellar cells. During the proceeding of PCD, AcPase was located on the nuclear membrane, nucleus, cytoplasma membrane, ER membrane, Mitochondria membrane, and cytoplasm in respectively. At the same time, it accompanied with phenomena of apoptosis which included, chromatin condensation and margination, cytoplasm vacuolation, mitochondria swelling and alveolation. endoplasmic reticulum(ER) cisterna swelling, cytoplasma membrane and cell wall separation, cell apparatuses
degeneration, apoptosis bodies produce, which contained the degenerated material from cell apparatuses and cell nucleus. At last, the cell walls were broken, and apoptosi
1．水稻胚乳细胞的构建 花后约两天，游离核胚乳开始细胞化。中央细胞壁产生钉状垂周内突，延伸入中央细胞腔。内质网线状排列，通过电子致密小泡相互连接；同时，小泡将内质网和壁内突融合，成为垂周壁，将合胞体的胚乳分隔成一个个向心开口的圆柱体。圆柱体内的核分裂，伴随产生成膜体，在成膜体的基础上融合小泡产生平周壁，形成一个个单核的细胞。平周壁的产生还有另一个来源，即通过垂周壁的游离末端分支相互融合而成。携带有壁物质的电子致密小泡可来源于高尔基体分沁的携带有壁物质的小泡；内质网自身囊腔膨大而缢缩形成的球状体；周围珠心细胞PCD降解后产生的，包含有降解物质的电子致密小泡，通过中央细胞壁内突而运输入中央细胞。还观察到细胞化过程中酸性磷酸酶(AcPase)的定位变化。在新形成的细胞壁上观察到强烈的AcPase的活性，而早已形成的细胞壁上几乎没有酶活。
2．胚乳细胞中淀粉质体的发生水稻淀粉粒属复粒淀粉，淀粉质体除来源于质体、淀粉质体外，还来源于线粒体。，花后约60小时，胚乳细胞中开始出现淀粉体。淀粉体可通过出芽和中间缢缩断裂两种方式进行增殖。用AcPase定位表明，完全淀粉化的胚乳细胞中，细胞核被淀粉质体挤压变形而解体，其余细胞器被吞噬或解体，直至整个胚乳细胞中只含有淀粉质体、蛋白质体和脂体，而表现为一种凋亡表形。不同发育时期稻米籽粒胚乳细胞中，淀粉粒大小的动态变化表明：花后3～12天是水稻胚乳细胞中淀粉粒的发育的高峰期，在该时段，淀
水稻胚乳发育中珠心凋亡和胚乳细胞构建的研究粉粒直径的增加几乎接近于直线增长。12天以后，淀粉粒的直径维持在一个接近恒定的水平。胚乳中央部分与周围部分的细胞中淀粉粒的形状有所不同。中央部分的多为多面体形，且排列紧密;而周围部分的淀粉粒多为圆球形，排列较为疏松 3.胚乳发育过程中，珠心细胞的细胞程序性死亡珠心细胞PCD始于大抱子时期。用超微细胞化学的方法，对珠心细胞PCD过程中，酸性磷酸酶(aeid phosphatase，Aepase)进行研究表明，随着pen进行的程度的不同，AcPase依次定位于核膜、核质、胞质、细胞质膜、液泡膜、内质网膜和线粒体膜，并伴随有典型的凋亡细胞表形。表现为:染色体固缩且周缘化，线粒体腔膨大，内质网槽库膨大，细胞质膜和细胞壁分离，出现大量吞噬泡，细胞器被降解或吞噬，最终产生凋亡小体，其中包含有降解的核质或胞质，胞壁破裂，凋亡小体释放。同时，还用线粒体标志酶一细胞色素氧化酶(cytochrome oxidase)对线粒体进行超微细胞化学的定位研究，证明了在PCD过程中确有线粒体晴模糊，腔膨大，线粒体膜破裂等线粒体解体现象的存在。观察表明珠心细胞的PCD降解产物对胚乳细胞构建的哺乳关系。
Rice(Oryza Sativa L.) is one of the main resources of cereal grown in China and even in the Asian. It plays a very important role in food supply. Endosperm is the most primary part of the rice grain that can be eaten, so the developmental pattern of endosperm influence the grain quantity and quantily directly. Starch is both the major component of the yield in the world's main crop plants and an important raw material for many industrial processes.
In this thesis, we report the structural and functional research about nucellar cells PCD, endosperm cells construction, and amyloplasts initiation and development during the development of the endosperm of the rice. The main research results were as followed.
1.Construction of the rice endosperm cells Cellularization of the free-nuclear endosperm was initiated about 2 days after pollination(DAP) with the form of the peg-like anticlinal ingrowths on the central cell wall. These ingrowths extended into the lumen of central cell and fused with the linear endoplasmic reticulum(ER) by many electron-dense vesicles. Periclinal walls were formed by two mechanisms: (1) formation of the phragmoplasts followed the kynokinesis, (2) fusion of the free-edges of the opposing anticlinal walls. It was speculated that the electron-dense vesicles had three resources. From the Golgi apparatus, ER or degenerated nucellar cells which were transported through the ingrowths of central cell wall. By the means of cytochemistry of ultrastructure, We observed the location change of the AcPase. During the proceeding of the cellularization, the new-formed cell walls had distinct aggradation of the AcPase, but on the cell walls which were formed a longer time ago, we could not observe the aggr
adation of the AcPase.
2. The resource and development of amyloplasts in the
endosperm cells The amyloplasts of rice were compound starch granules. We observed that the amyloplasts could be initiated not only from the plastids, proplastids and amyloplasts, but also from the mitochondria. They appeared in the endosperm cell after 60 hours after pollination, and propagated by the formation of gemma or by contraction in the middle before splitting. The ultrastructural location of AcPase suggested that the nucleus and other cell apparatus were degenerated in the mature endosperm cells, which were filled with amyloplasts and protein bodies. So they showed another kind of phenomena of PCD.
The compare of the starch granules of different stage of development showed that the 3-12 days after pollination was the fastigium of the development of the starch granules. During this period, the augmentation of the diameter of the starch granules was approximate the pattern of beeline. But after 12 DAP, the diameter of the starch granules maintained at a invariable level. The shape of the starch granules in the medial region of the endosperm was different from the ambient region of that. Most of the starch granules in the medial region of the endosperm was polyhedral, crystalline and compact arranged; but the starch granules in the ambient region were almost ball shape, loose arrangement.
3. The proceeding of Programmed Cell Death (PCD) of nucellar cells during the development of the endosperm PCD of nucellar cells initiated in 2 days after pollination (2DAP). By the means of ultrastructure cytochemistry, we researched the variation of AcPase (Acid phosphatase) in nucellar cells. During the proceeding of PCD, AcPase was located on the nuclear membrane, nucleus, cytoplasma membrane, ER membrane, Mitochondria membrane, and cytoplasm in respectively. At the same time, it accompanied with phenomena of apoptosis which included, chromatin condensation and margination, cytoplasm vacuolation, mitochondria swelling and alveolation. endoplasmic reticulum(ER) cisterna swelling, cytoplasma membrane and cell wall separation, cell apparatuses
degeneration, apoptosis bodies produce, which contained the degenerated material from cell apparatuses and cell nucleus. At last, the cell walls were broken, and apoptosi
引文
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Folsom M W, Cass D D. Embryo sac development in soybean: the central cell and aspects of fertilization. Amer J Bot, 1992,79(12): 1407-1017
Gartner P J, Nagl W. Acid phosphstase activity in platids (plastolysomes) of senescing embryo-suspensor cells. Planta, 1980, 149:341-349
Gori P. The fine structure of the developing Euphorbia dulcis endosperm. Ann Bot, 1987, 60:563-569
Greenberg J T. Progranned cell death: a way of life for plants. Proc Natl Acad Sci USA, 1996, 93:12094-12097
Halac I N D. Fine structure of nucellarr cells during development of the embryo sac in Oenothera biennis. Ann Bot, 1980, 45:515-521
Jensen C G. Dynamic of spindle microtubule organization: kinetochore fiber microtubules of plant endosperm. J Cell Bio, 1982, 92:540-557
Kluck R M, Bossy-Wetzel E, Green D R, Newrneyer D D. The release of
cytochrome c from mitochondria: a primary site for Bcl-2 regulation of apoptosis. SCINENCE, 1997, 275:1132-1136
Knight M E, Harn C, Lilley C E R, Guan H, Singletary G W, Mu Forster C, Wasserman B P, Keeling P L. Molecular cloning of starch synthase I from maize(W64) endosperm and expression in Escherichia coli. Plant J, 1998, 14(5):613-622
Lammeren V, M A A. Structure and function of the microtubular cytoskeleton during endosperm development in wheat: an immunofluorescence study. Protoplasm, 1988, 146:18-27
Li P, Nijhawan D, Budihardjo I, Srinivasula S M, Ahmad M, Alnemeri E S, Wang X. Cytochome c and dATP-dependent formation of Apaf-1/caspase-9 complex initiates an apoptosis protaase cascade. Cell, 1997, 91:479-489
Liu X, Kim C N, Yang J, Jemmerson R, Wang X. Induction of apoptosis program in cell-free extracts: requirement for dATP and cytochrome c. Cell, 1996, 86:147-157
Matile P, Ginsburg S, Schellenberg M, Thomas H. Catobolites of chlorophyⅡ in senescing barley leaves are localized in the vacuoles of mesophyⅡ cell. Proc Natl Acad Sci USA, 1988, 85:9529-9532
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Ramachandran C, Raghavan V. Changes in nuclear DNA content of endosperm cells during grain development in rice (Oryza sativa). Ann Bot, 1989, 64(4):459-468
Reed J C. Cytochrome c: can't live with it——can't live without it. Cell, 1997, 91:559-562
Steller H. Mochanisms and genes of cellar suicide. SCINENCE, 1995, 276:1445-1462
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Thimann K V, Tetley R M, Krivak B M. Metabolism of oat leaves during senescence. Plant Physiol, 1977, 59:448-454
Webb M C, Gunning B E S. The microtubular cytoskeleton during development of the zygote, proembryo and free-nuclear endosperm in Arabidopsis thaliana(L.) Heynh. Palanta, 1991, 184(2):187-195
Williams G T. Molecular regulation of apoptosis: genetic control on cell death. CELL, 1993, 74:777-779
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Umemoto T, Nakamura Y, Ishikura N. Activity of starch synthase and the amylose content in rice endosperm. Phytochemistry, 1995, 40(6): 1613-1616
Zou H, Henzel W J, Liu X, Lutschg A, Wang X. Apaf-1, a human protein homologous to C. elegans CED-4, participates in cytochrome cdependent activation of caspase-3. Cell, 1997, 90:405-413
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Folsom M W. Cass D D. Embryo sac development in soybean: cellularization and egg apparatus expansion. CAN J BOT, 1990, 68:2135-2147
Folsom M W, Cass D D. Embryo sac development in soybean: the central cell and aspects of fertilization. Amer J Bot, 1992,79(12): 1407-1017
Gartner P J, Nagl W. Acid phosphstase activity in platids (plastolysomes) of senescing embryo-suspensor cells. Planta, 1980, 149:341-349
Gori P. The fine structure of the developing Euphorbia dulcis endosperm. Ann Bot, 1987, 60:563-569
Greenberg J T. Progranned cell death: a way of life for plants. Proc Natl Acad Sci USA, 1996, 93:12094-12097
Halac I N D. Fine structure of nucellarr cells during development of the embryo sac in Oenothera biennis. Ann Bot, 1980, 45:515-521
Jensen C G. Dynamic of spindle microtubule organization: kinetochore fiber microtubules of plant endosperm. J Cell Bio, 1982, 92:540-557
Kluck R M, Bossy-Wetzel E, Green D R, Newrneyer D D. The release of
cytochrome c from mitochondria: a primary site for Bcl-2 regulation of apoptosis. SCINENCE, 1997, 275:1132-1136
Knight M E, Harn C, Lilley C E R, Guan H, Singletary G W, Mu Forster C, Wasserman B P, Keeling P L. Molecular cloning of starch synthase I from maize(W64) endosperm and expression in Escherichia coli. Plant J, 1998, 14(5):613-622
Lammeren V, M A A. Structure and function of the microtubular cytoskeleton during endosperm development in wheat: an immunofluorescence study. Protoplasm, 1988, 146:18-27
Li P, Nijhawan D, Budihardjo I, Srinivasula S M, Ahmad M, Alnemeri E S, Wang X. Cytochome c and dATP-dependent formation of Apaf-1/caspase-9 complex initiates an apoptosis protaase cascade. Cell, 1997, 91:479-489
Liu X, Kim C N, Yang J, Jemmerson R, Wang X. Induction of apoptosis program in cell-free extracts: requirement for dATP and cytochrome c. Cell, 1996, 86:147-157
Matile P, Ginsburg S, Schellenberg M, Thomas H. Catobolites of chlorophyⅡ in senescing barley leaves are localized in the vacuoles of mesophyⅡ cell. Proc Natl Acad Sci USA, 1988, 85:9529-9532
Nakamura Y, Umemoto T, Takahata Y, Komae K, Amano E, Satoh H. Changes in structure of starch and enzyme activities affected by sugary mutations in developing rice endosperm. Possible role of starch debranching enzyme (R-enzyme) in amylopection biosynthesis. Physio Plantarum, 1996, 97(3):491-498
Otegui M, Maldonado S. Lina C, Lederkremer R M. Development of the Myrsine laetevirens (Myrsinaceae). I. Cellularization and deposition of cell-wall storage carbohydrates, Int J Plant Sci, 1999, 160(3):491-500
Ramachandran C, Raghavan V. Changes in nuclear DNA content of endosperm cells during grain development in rice (Oryza sativa). Ann Bot, 1989, 64(4):459-468
Reed J C. Cytochrome c: can't live with it——can't live without it. Cell, 1997, 91:559-562
Steller H. Mochanisms and genes of cellar suicide. SCINENCE, 1995, 276:1445-1462
Smith A M, Denyer K, Martin C. The synthesis of the starch granule. Annu Rev Plant Physiol Plant Mol Biol, 1997, 48:67-87
Thimann K V, Tetley R M, Krivak B M. Metabolism of oat leaves during senescence. Plant Physiol, 1977, 59:448-454
Webb M C, Gunning B E S. The microtubular cytoskeleton during development of the zygote, proembryo and free-nuclear endosperm in Arabidopsis thaliana(L.) Heynh. Palanta, 1991, 184(2):187-195
Williams G T. Molecular regulation of apoptosis: genetic control on cell death. CELL, 1993, 74:777-779
Wittenbach V A, Lin W, Hebert R R. Vacuolar localization of proteases and degradation of chloroplasts in mesophyⅡ protoplasts from senescing primary wheat leaves. Plant Physiol, 1982, 69:98-102
Umemoto T, Nakamura Y, Ishikura N. Activity of starch synthase and the amylose content in rice endosperm. Phytochemistry, 1995, 40(6): 1613-1616
Zou H, Henzel W J, Liu X, Lutschg A, Wang X. Apaf-1, a human protein homologous to C. elegans CED-4, participates in cytochrome cdependent activation of caspase-3. Cell, 1997, 90:405-413
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