高粱胚乳细胞与母体组织发育关系的研究

doi:10.3864/j.issn.0578-1752.2014.17.002

摘要/Abstract

摘要： 【目的】探明高粱颖果发育过程中胚乳细胞与母体组织发育关系及其胞内淀粉体的发育特性。【方法】通过挂牌和记号笔点颖相结合的方法对供试高粱品种KS-304花后发育天数进行精确标记，详细跟踪观测了颖果的生长；采取Spurr树脂包埋、番红-甲基紫复染法制作半薄切片，光镜下详细观察了高粱颖果发育各过程中胚乳各部位以及种皮果皮结构的变化差异与联系；采用扫描电镜研究发育中后期及成熟时颖果断面不同部位细胞内淀粉体的形态；应用冰冻切片，荧光显微镜观察了成熟颖果横断面的结构。【结果】颖果发育分形成期、乳熟期、蜡熟期与完熟期，内胚乳发育分为游离核期、细胞化期、分化期、发育期及成熟期；其中颖果发育形成期与胚乳发育的前3个时期相对应，乳熟期对应发育期，而蜡熟期与完熟期对应胚乳的成熟期。颖果发育早期，珠心组织存留时间较长，珠心表皮细胞约在花后15 d消失。花后7 d，表层胚乳细胞开始积累脂质体，11 d时转为糊粉层细胞，成熟时糊粉层为1层，其细胞内除常规糊粉粒圆球体外，还含有少量单粒淀粉体，粒径约3 μm。亚糊粉层细胞位于糊粉层细胞与内胚乳细胞之间，兼糊粉层与内胚乳细胞特点，贮藏大量蛋白体，细胞内淀粉体构成复杂。内胚乳发育存在区域差异，中央近胚处的细胞物质积累滞后于周缘胚乳细胞，成熟时，前者淀粉体充实较为疏松，发育为粉质胚乳，后者充实紧密，淀粉体彼此受到挤压呈多面体，最终发育为角质胚乳。高粱胚乳细胞内存在于不同于其他谷物颖果胚乳淀粉体的“发生中心”结构，主要表现为淀粉粒在管状质体内生长，随着体积的增长，后期与“发生中心”分离而形成新的淀粉体。果皮发育前期增厚，中后期呈缓慢减薄的趋势，中果皮发育后期细胞内观察到淀粉体存在特殊的二次增长，构成从前期复粒淀粉体为主转为单粒；“种皮”来源于珠心、珠被细胞依次降解后的胞壁残留堆叠而成，中间含有透明角质层。【结论】白高粱KS-304胚乳发育与玉米颖果胚乳类似，最终形成角质与粉质胚乳；颖果胚乳淀粉体发育存在独特的“发生中心”；中果皮发育后期其细胞可能行使积累同化产物“库”的作用。

关键词: 高粱 , 胚乳 , 淀粉体 , 糊粉层 , 果皮 , 种皮

Abstract: 【Objective】The objective of this research was to clarify the relationship between endosperm and its maternal tissue during development process of the sorghum caryopsis. 【Method】Sorghum KS-304 was used as the experimental material with its caryopsis development days precisely recorded and its caryopsis development closely observed. Structural changes of endosperm cells and relation with their maternal tissue were observed through semi-thin sections under light microscopy by applying Spurr resin tissue embedding. The ultrastructure of cells from different tissues and the starch granules wherein in developing and full ripe caryopsis were observed under SEM. The relationships of testa and aleurone layer of full ripe caryopsis were studied through fluorescent microscopy using cryosectioning.【Result】Four stages could be found in caryopsis development, they were formation stage, milky stage, dough stage, and full maturity stage. Accordingly, endosperm development was divided into five stages, i.e. coenocyte stage, cellurization stage, differentiation stage, developmental stage and the final maturation stage, and the first three of which are equal to that of the formation stage of caryopsis; while the last maturation stage corresponds to the last two stages of caryopsis development. It took as long as 15 days for the nucellar epidermis to be fully degraded. The outer peripheral endosperm cells started to accumulate lipid bodies as early as 7 DPA (days after pollination) and, turned into aleurone cells in 11 DAP. Only 1 layer of aleurone cells formed in full ripe caryopsis. Besides commonly seen aleurone granules and globoids, some single amyloplasts were also found on ripe aleurone cells, circa. 3 μm in diameter. Growth and development of endosperm cells also varied according to different locations, amyloplasts in cells surrounding embryo tended to have a slow growth rate and were loosely packed than other areas, making them floury endosperm compared to others’ corneous endosperms when finally matured. Amyloplasts constitution in subaleurone cells were quite composite, in a manner that mingles both aleurone and starchy endosperm cells. Amyloplast formation in starchy endosperm was unique, which has an “occurrence center” during initiation. Starch “grow” inside a tube-like plastid, and when mature, falls apart, leaving the mature amyloplast with an uneven oval shape. Starch/amyloplasts in mesocarp may not be exhausted approaching mature stage, and on the contrary, there seems to be a secondary growth in both quantity and diameter.【Conclusion】 Development of starchy endosperm in sorghum KS-304 showed a similar pattern to the corn. Starch formation of amyloplasts in starchy endosperm followed a unique way and is independent of all other crop spiecies. Cells of mesocarp layer may act as an extra “sink” during later stages in caryopsis development.

Key words: sorghum , endosperm , amyloplast , aleurone , pericarp , testa

李栋梁，荆彦平，李小刚，顾蕴洁，王忠. 高粱胚乳细胞与母体组织发育关系的研究[J]. 中国农业科学, 2014, 47(17): 3336-3347.

LI Dong-liang, JING Yan-ping, LI Xiao-gang, GU Yun-jie, WANG Zhong. Investigation of Sorghum Endosperm Cell Development and the Relationship with Its Maternal Tissue[J]. Scientia Agricultura Sinica, 2014, 47(17): 3336-3347.

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参考文献

[1]王海莲, 管延安, 张华文, 杨延兵, 秦岭. 高梁基因组学研究进展. 基因组学与应用生物学, 2009, 28(3): 549-556.
Wang H L, Guan Y A, Zhang H W, Yang Y B, Qin L. Advances in the study on sorghum genomics. Genomics and Applied Biology, 2009, 28(3): 549-556. (in Chinese)
[2]Earp C F, McDonough C M, Rooney L W. Microscopy of pericarp development in the caryopsis of Sorghum bicolor (L.) Moench. Journal of Cereal Science, 2004, 39(1): 21-27.
[3]顾蕴洁, 王忠, 陈义芳, 陈刚, 黄海. 高粱胚乳发育的观察与研究. 南京师大学报: 自然科学版, 2002, 25(3): 99-108.
Gu Y J, Wang Z, Chen Y F, Chen G, Huang H. The observation and study on the development of sorghum endosperm. Journal of Nanjing Normal University: Natural Science Edition, 2002, 25(3): 99-108. (in Chinese)
[4]王忠, 顾蕴洁, 王慧慧, 郑彦坤, 王玲玲, 李栋梁. 关于高粱胚乳细胞发育的研究. 中国科技论文在线 2012. http://www.paper.edu.cn.
Wang Z, Gu Y J, Wang H H, Zheng Y K, Wang L L, Li D L. Investigation of the sorghum endosperm cell development. Sciencepaper online. 2012. http://www.paper.edu.cn. (in Chinese)
[5]Kladnik A, Chourey P S, Pring D R, Dermastia M. Development of the endosperm of Sorghum bicolor during the endoreduplication- associated growth phase. Journal of Cereal Science, 2006, 43(2): 209-215.
[6]汤述翥, 薛元龙. 杂交高粱KS-304开发利用研究初报. 江苏农业科学, 1998(6): 24-26.
Tang S Z, Xue Y L. Preliminary report on exploitation and utilization of sorghum hybrids KS-304. Jiangsu Agricutural Science, 1998(6): 24-26.(in Chinese)
[7]李筱倩, 孙龙生, 张艳云, 罗方妮. KS-304 杂交高粱对猪饲用价值的研究. 中国饲料, 1998, 4: 006.9-10.
Li X Q, Sun L S, Zhang Y Y, Luo F N. Studies on feeding values of sorghum hybrids KS-304 for swine. China Feed, 1998, 4:006: 9-10.(in Chinese)
[8]Freeman J A, Spurlock B O. A new epoxy embedment for electron microscopy. The Journal of Cell Biology, 1962, 13(3): 437-443.
[9]王忠, 顾蕴洁, 郑彦坤, 王慧慧. 水稻胚乳细胞发育的结构观察及其矿质元素分析. 中国水稻科学, 2012, 26(6): 693-705.
Wang Z, Gu Y J, Zheng Y K, Wang H H. Structure observation of rice endosperm cell development and its mineral element analysis. Chinese Journal of Rice Science, 2012, 26(6): 693-705. (in Chinese)
[10]李冉俐, 申家恒, 贾媛, 李伟, 王黎明. 高粱受精过程及其经历的时间. 作物学报, 2009, 35(12): 2234-2242.
Li R L, Shen J H, Jia Y, Li W, Wang L M. Fertilization process in sorghum and its performance time for each stage. Acta Agronomica Sinica, 2009, 35(12): 2234-2242. (in Chinese)
[11]Reddy R N, Narayana L L, Rao N G P. Apomixis and its utilisation in grain sorghum-II; embryology of F1 progeny of reciprocal crosses between R473 and 302. Proceedings of the Indian Academy of Sciences: Section B. Part 2, Plant Sciences, 1979, 88(6): 455-461.
[12]Bosnes M, Weideman F, Olsen O A. Endosperm differentiation in barley wild-type and sex mutants. The Plant Journal, 1992, 2(5): 661-674.
[13]Olsen O A. Nuclear endosperm development in cereals and Arabidopsis thaliana. The Plant Cell Online, 2004, 16(suppl. 1): S214-S227.
[14]Swigoňová Z, Lai J, Ma J, Ramakrishna W. Close split of sorghum and maize genome progenitors. Genome Research, 2004, 14(10a): 1916-1923.
[15]Evers T, Millar S. Cereal grain structure and development: Some implications for quality. Journal of Cereal Science, 2002, 36: 261-284.
[16]王忠, 李卫芳, 顾蕴洁, 石火英, 高煜珠. 水稻胚乳的发育及其养分输入的途径.作物学报,1995, 21(5):520-527
Wang Z, Li W F, Gu Y J, Shi H Y, Gao Y Z. Development of rice endosperm and the pathway of nutrients entering the endosperm. Acta Agronomica Sinica, 1995, 21(5): 520-527. (in Chinese)
[17]王忠, 顾蕴洁. 小麦胚乳发育及其养分输入途径. 作物学报, 1998, 24(5): 536-543.
Wang Z, Gu Y J. Development of wheat endosperm and the pathway of nutrients entering the endosperm. Acta Agronomica Sinica, 1998, 24(5): 536-543. (in Chinese)
[18]王忠, 顾蕴洁, 李卫芳, 练兴明, 陈刚. 关于大麦胚乳发育的研究. 中国农业科学, 1996, 29(6): 38-45.
Wang Z, Gu Y J, Li W F, Lian X M, Chen G. The study about development of barley endosperm. Scientia Agricultura Sinica, 1996, 29(6): 38-45. (in Chinese)
[19]Charlton W L, Keen C L, Merriman C. Endosperm development in Zea mays; implication of gametic imprinting and paternal excess in regulation of transfer layer development. Development, 1995, 121(9): 3089-3097.
[20]Robert P, Jamme F, Barron C, Bouchet B, Saulnier L. Change in wall composition of transfer and aleurone cells during wheat grain development. Planta, 2011, 233(2): 393-406.
[21]Artschwager E, McGuire R C. Cytology of reproduction in sorghum vulgare. Journal of Agricultural Research, 1949, 78: 659-673.
[22]Rooney L W, Miller F R, Mertin J V. Variation in the structure and kernel characteristics of sorghum//Proceedings of the International Symposium On Sorghum Grain Quality. ICRISAT Center, Patancheru, India, 1981: 143-162.
[23]Paterson A H, Bowers J E, Bruggmann R, Dubchak I. The sorghum bicolor genome and the diversification of grasses. Nature, 2009, 457(7229): 551-556.
[24]Mace E S, Tai S, Gilding E K, Li Y, Prentis P J. Whole-genome sequencing reveals untapped genetic potential in Africa’s indigenous cereal crop sorghum. Nature Communications, 2013, 4: 2320.
[25]Earp C F, McDonough C M, Awika J, Rooney L W. Testa development in the caryopsis of Sorghum bicolor (L.) Moench. Journal of Cereal Science, 2004, 39(2): 303-311.