树木营养贮藏蛋白质的细胞学、生物化学和生物学功能的研究
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摘要
氮是植物生长发育不可缺少的一种大量矿质元素，对植物氮代谢的研究一直受到高度重视。近年来，关于氮素吸收、同化、转运和种子氮贮藏的研究已取得较大进展，但是，营养器官中氮贮藏的研究仍然是植物氮代谢研究的一个薄弱环节。
营养器官中的季节性氮贮藏是树木氮代谢的显著特征，在树木的生长发育及树木对逆境的反应方面起重要作用。因此，关于氮贮藏的知识也是果树和林木栽培技术发展的基础。
二十世纪八十年代后期，树木氮贮藏研究获得了重大进展：在树木的营养器官中发现一类专门的贮藏蛋白质，称之为营养贮藏蛋白质。
至今，对树木营养贮藏蛋白质的细胞学、生物化学性质和生物学功能了解得还很不够。对树木营养贮藏蛋白质的认识借鉴种子，往往把营养贮藏蛋白质的重要性与种子中的贮藏蛋白质相类比。一般认为，在根系当年从土壤中吸收和同化的氮不能满足树木当年生长发育的需要时，营养器官中贮藏的氮起着一种“缓冲作用”。这种认识严重制约着树木营养贮藏蛋白质的研究。
本文首先综述了植物N代谢的差异、树木营养器官N贮藏的早期研究工作和植物营养贮藏蛋白质的研究进展。采用光镜和电镜技术、PAGE、SDS-PAGE和免疫印迹技术、电泳凝胶过碘酸—Schiff试剂染色、间接免疫荧光和电镜免疫细胞化学定位技术以及cDNA克隆技术，较深入地研究了大叶桃花心木和巴西橡胶树的营养贮藏蛋白质的细胞学、生物化学性质和生物学功能。在此基础上，较广泛地研究了营养贮藏蛋白质在18科64属70种热带树木和15科28属30种2变种温带落叶树木中的分布，较系统地研究了营养贮藏蛋白质在蔷薇科(12属17种3变种)、豆科(9属10种1变种)和楝科(12属16种2变种)中的分
布、超微结构及其分离鉴定和糠科树木营养贮藏蛋白质的免疫相关性。
在大叶桃花心木的皮层、次生韧皮部和木质部的普通薄壁细胞的
中央大液泡里存在蛋白质内含物。这些细胞的周缘细胞质中通常含有
淀粉粒。在接近落叶期和无叶期，液泡蛋白质在植株中大量积累，主
要分布在次生韧皮部的射线薄壁组织细胞和韧皮薄壁细胞中，并且末
端小枝的液泡蛋白质比树干和大根的丰富。末端小枝液泡蛋白质的消
失与新梢生长过程是同步的，表明这些蛋白质被新梢生长所消耗。落
叶前环剥一年生枝条基部，萌发的新梢生长被抑制，环剥伤口下方液
泡蛋白质的消失受到阻碍。树干和根中液泡蛋白质的明显减少发生在
新梢形成之后，这些蛋白质的完全消失发生在相应器官形成层活动最
活跃的时期。在新梢叶片成熟后l个月，新梢茎的树皮组织就开始积
累液泡蛋白质，此时正值高温和长日照时期。
采用SDS干AGE技术，从大叶桃花心木末端小枝的树皮组织中分离
到两种才对含量高的蛋白质，分子量约 18 kDa和 ZI kDa。用 21 kDa
蛋白质的抗血清进行兔疫印迹分析表明，18 kDa蛋白质和 21 kDa蛋白
质有相同的抗原决定簇。免疫荧光和电镜免疫细胞化学定位证明，它
们是贮藏蛋白质细胞的液泡蛋白质内含物的主要成分。在树木年生长
周期的后期，18 kDa和 ZI kDa蛋白质分布于枝条、树干、大根和小
根，大量积累在这些器官的次生韧皮部和末端小枝及小根的次生木质
部中。在树木新的年生长周期中，随着新梢的发育，积累在末端小枝
中的贮藏蛋白质开始被消耗／新梢叶片成熟时，末端小枝中的 18 kDa
和 21 kDa蛋白质己完全消失，而树干和大根中的这两种蛋白质的含量
相对稳定，与落叶期相比几乎没有变化。树干和大根中的贮藏蛋白质
的动用与各自的维管形成层活动密切相关，它们的大量消失发生在维
管形成层活动的高峰期。到IC。月初，高部位树干的树皮中己开始积累
贮藏蛋白质而低部位树干的树皮组织中的贮藏蛋白质己减少到低水
平。新梢叶片成熟后的 1周，在新梢茎的树皮组织中就可检测到 18 kDa
和 21 kDa两种蛋白质。新梢叶片成熟后的 2周，这两种蛋白质明显增
多。此时，上一年形成的末端小枝的树皮组织中己检测不到 18 kDa和
ZI kDa两种蛋白质，树干和根中的贮藏蛋白质正在消失。这些结果表
明，营养贮藏蛋白质的积累是一种程序性合成，积累的蛋白质供下一
个生长发育阶段利用。结果，当年的新梢生长和维管形成层的活动优
先利用原来积累的贮藏蛋白质而不是当年新同化的N。这种机制为树木
正常的生长发育提供了一个相对稳定的内环境。树木的茎干在整个年
3
生长周期中都存在相当数量的营养贮藏蛋白质，这对于树木抵抗偶然
的N素胁迫具有重要的生
People always attach importance to the nitrogen metabolosm in plant for nitrogen, a major mineral element, is necessary for plant growth and development. However, we still have little knowledge of the nitrogen storage in vegetative organs although more progress in understanding the nitrogen absorption, assimilation, transportation and the nitrogen storage in seeds have been achieved in recent years.
The nitrogen metabolism of woody plants is characterized by storing nitrogen compounds in their vegetative organs. The stored nitrogen compounds play an important role in the tree growth and in the resistance of tree to environmental stress. In addition, the knowledge of nitrogen storage is a basis of developing the cultivation techniques of woods and fruit trees.
At the late of 1980s, available evidence indicated that there presented a kind of proteins specific for nitrogen storage in vegetative organs of woody plants, and ,thus, the proteins were defined as vegetative storage proteins.
To date, little is known about the cytology, biochemical properties and biological roles of vegetative storage proteins in woody plants. As a result, people have to draw on the experience of seed storage proteins to understand the vegetative storage proteins and always equate their importance with that of seed storage proteins. It is commonly accepted that the nitrogen stored in vegetative organs buffers the nitrogen necessity if current supply of nitrogen is not available for the growth of woody plants. This understand of stored nitrogen compounds restricted seriously the progress in the investigation of vegetative storage proteins.
In the dissertation, we studied more extensively the cytology, biochemical properties and biological roles of vegetative storage proteins in Swietenia macrophylla and in Hevea brasiliensis by light- and electron microscopy, SDS-PAGE, PAGE, immuno-blotting, indirect immunohistochemical localization and colloidal gold labelling and cDNA clone techniques. To examine the distribution of vegetative storage proteins in woody plants, we sampled at random 70 tree species of tropical hardwoods
in 64 genera of 18 families and 30 tree species and 1 variety of temperate deciduous hardwoods in 28 genera of 15 families. We further investigated more systematically the distribution, ultrastructure of vegetative storage proteins in Rosaceae (17 species and 3 varieties in 12 genera), Leguminosae (10 species and 1 variety in 9 genera) and Meliaceae (16 species and 2 varieties in 12 genera), identified many vegetative storage proteins and tested their irnmuno-relatedness in Meliaceae and in other families.
The protein-storing cells in Swietenia macrophylla were found to be Populus-type, i.e. ordinary parenchyma cells containing both vacuolar protein inclusions and starch grains. The protein inclusions in a granular appearance accumulated in the central vacuole and starch grains generally in the cytoplasm of ordinary parenchyma cells. The storage protein accumulated in large quantity in the vegetative organs of Swietenia macrophylla at the period near leaf fall and in the leaf-fall period with terminal branchlets much richer in the storge protein than trunks and roots. The protein accumulated mainly in secondary phloem parenchyma cells and secondary phloem ray cells. The degradation of storage protein in terminal branchlets of Swietenia macrophylla synchronized with new shoot growth after leaf-absent period, suggesting that the protein was utilized to support the growth of new shoots. When the diminishing of the storage protein below the girdled site was blocked, the new shoot growth was also restrained. The storage protein in the trunks and roots diminished dramatically after the new shoot had fully developed and the protein had disappeared completely when the cambia of these organs activated vigorously. In the annual growth cycle of Swietenia macrophylla, the storage protein accumulated firstly in the new shoot rather than in the other organs. The protein accumulation began one month after the leaves of new
营养器官中的季节性氮贮藏是树木氮代谢的显著特征，在树木的生长发育及树木对逆境的反应方面起重要作用。因此，关于氮贮藏的知识也是果树和林木栽培技术发展的基础。
二十世纪八十年代后期，树木氮贮藏研究获得了重大进展：在树木的营养器官中发现一类专门的贮藏蛋白质，称之为营养贮藏蛋白质。
至今，对树木营养贮藏蛋白质的细胞学、生物化学性质和生物学功能了解得还很不够。对树木营养贮藏蛋白质的认识借鉴种子，往往把营养贮藏蛋白质的重要性与种子中的贮藏蛋白质相类比。一般认为，在根系当年从土壤中吸收和同化的氮不能满足树木当年生长发育的需要时，营养器官中贮藏的氮起着一种“缓冲作用”。这种认识严重制约着树木营养贮藏蛋白质的研究。
本文首先综述了植物N代谢的差异、树木营养器官N贮藏的早期研究工作和植物营养贮藏蛋白质的研究进展。采用光镜和电镜技术、PAGE、SDS-PAGE和免疫印迹技术、电泳凝胶过碘酸—Schiff试剂染色、间接免疫荧光和电镜免疫细胞化学定位技术以及cDNA克隆技术，较深入地研究了大叶桃花心木和巴西橡胶树的营养贮藏蛋白质的细胞学、生物化学性质和生物学功能。在此基础上，较广泛地研究了营养贮藏蛋白质在18科64属70种热带树木和15科28属30种2变种温带落叶树木中的分布，较系统地研究了营养贮藏蛋白质在蔷薇科(12属17种3变种)、豆科(9属10种1变种)和楝科(12属16种2变种)中的分
布、超微结构及其分离鉴定和糠科树木营养贮藏蛋白质的免疫相关性。
在大叶桃花心木的皮层、次生韧皮部和木质部的普通薄壁细胞的
中央大液泡里存在蛋白质内含物。这些细胞的周缘细胞质中通常含有
淀粉粒。在接近落叶期和无叶期，液泡蛋白质在植株中大量积累，主
要分布在次生韧皮部的射线薄壁组织细胞和韧皮薄壁细胞中，并且末
端小枝的液泡蛋白质比树干和大根的丰富。末端小枝液泡蛋白质的消
失与新梢生长过程是同步的，表明这些蛋白质被新梢生长所消耗。落
叶前环剥一年生枝条基部，萌发的新梢生长被抑制，环剥伤口下方液
泡蛋白质的消失受到阻碍。树干和根中液泡蛋白质的明显减少发生在
新梢形成之后，这些蛋白质的完全消失发生在相应器官形成层活动最
活跃的时期。在新梢叶片成熟后l个月，新梢茎的树皮组织就开始积
累液泡蛋白质，此时正值高温和长日照时期。
采用SDS干AGE技术，从大叶桃花心木末端小枝的树皮组织中分离
到两种才对含量高的蛋白质，分子量约 18 kDa和 ZI kDa。用 21 kDa
蛋白质的抗血清进行兔疫印迹分析表明，18 kDa蛋白质和 21 kDa蛋白
质有相同的抗原决定簇。免疫荧光和电镜免疫细胞化学定位证明，它
们是贮藏蛋白质细胞的液泡蛋白质内含物的主要成分。在树木年生长
周期的后期，18 kDa和 ZI kDa蛋白质分布于枝条、树干、大根和小
根，大量积累在这些器官的次生韧皮部和末端小枝及小根的次生木质
部中。在树木新的年生长周期中，随着新梢的发育，积累在末端小枝
中的贮藏蛋白质开始被消耗／新梢叶片成熟时，末端小枝中的 18 kDa
和 21 kDa蛋白质己完全消失，而树干和大根中的这两种蛋白质的含量
相对稳定，与落叶期相比几乎没有变化。树干和大根中的贮藏蛋白质
的动用与各自的维管形成层活动密切相关，它们的大量消失发生在维
管形成层活动的高峰期。到IC。月初，高部位树干的树皮中己开始积累
贮藏蛋白质而低部位树干的树皮组织中的贮藏蛋白质己减少到低水
平。新梢叶片成熟后的 1周，在新梢茎的树皮组织中就可检测到 18 kDa
和 21 kDa两种蛋白质。新梢叶片成熟后的 2周，这两种蛋白质明显增
多。此时，上一年形成的末端小枝的树皮组织中己检测不到 18 kDa和
ZI kDa两种蛋白质，树干和根中的贮藏蛋白质正在消失。这些结果表
明，营养贮藏蛋白质的积累是一种程序性合成，积累的蛋白质供下一
个生长发育阶段利用。结果，当年的新梢生长和维管形成层的活动优
先利用原来积累的贮藏蛋白质而不是当年新同化的N。这种机制为树木
正常的生长发育提供了一个相对稳定的内环境。树木的茎干在整个年
3
生长周期中都存在相当数量的营养贮藏蛋白质，这对于树木抵抗偶然
的N素胁迫具有重要的生
People always attach importance to the nitrogen metabolosm in plant for nitrogen, a major mineral element, is necessary for plant growth and development. However, we still have little knowledge of the nitrogen storage in vegetative organs although more progress in understanding the nitrogen absorption, assimilation, transportation and the nitrogen storage in seeds have been achieved in recent years.
The nitrogen metabolism of woody plants is characterized by storing nitrogen compounds in their vegetative organs. The stored nitrogen compounds play an important role in the tree growth and in the resistance of tree to environmental stress. In addition, the knowledge of nitrogen storage is a basis of developing the cultivation techniques of woods and fruit trees.
At the late of 1980s, available evidence indicated that there presented a kind of proteins specific for nitrogen storage in vegetative organs of woody plants, and ,thus, the proteins were defined as vegetative storage proteins.
To date, little is known about the cytology, biochemical properties and biological roles of vegetative storage proteins in woody plants. As a result, people have to draw on the experience of seed storage proteins to understand the vegetative storage proteins and always equate their importance with that of seed storage proteins. It is commonly accepted that the nitrogen stored in vegetative organs buffers the nitrogen necessity if current supply of nitrogen is not available for the growth of woody plants. This understand of stored nitrogen compounds restricted seriously the progress in the investigation of vegetative storage proteins.
In the dissertation, we studied more extensively the cytology, biochemical properties and biological roles of vegetative storage proteins in Swietenia macrophylla and in Hevea brasiliensis by light- and electron microscopy, SDS-PAGE, PAGE, immuno-blotting, indirect immunohistochemical localization and colloidal gold labelling and cDNA clone techniques. To examine the distribution of vegetative storage proteins in woody plants, we sampled at random 70 tree species of tropical hardwoods
in 64 genera of 18 families and 30 tree species and 1 variety of temperate deciduous hardwoods in 28 genera of 15 families. We further investigated more systematically the distribution, ultrastructure of vegetative storage proteins in Rosaceae (17 species and 3 varieties in 12 genera), Leguminosae (10 species and 1 variety in 9 genera) and Meliaceae (16 species and 2 varieties in 12 genera), identified many vegetative storage proteins and tested their irnmuno-relatedness in Meliaceae and in other families.
The protein-storing cells in Swietenia macrophylla were found to be Populus-type, i.e. ordinary parenchyma cells containing both vacuolar protein inclusions and starch grains. The protein inclusions in a granular appearance accumulated in the central vacuole and starch grains generally in the cytoplasm of ordinary parenchyma cells. The storage protein accumulated in large quantity in the vegetative organs of Swietenia macrophylla at the period near leaf fall and in the leaf-fall period with terminal branchlets much richer in the storge protein than trunks and roots. The protein accumulated mainly in secondary phloem parenchyma cells and secondary phloem ray cells. The degradation of storage protein in terminal branchlets of Swietenia macrophylla synchronized with new shoot growth after leaf-absent period, suggesting that the protein was utilized to support the growth of new shoots. When the diminishing of the storage protein below the girdled site was blocked, the new shoot growth was also restrained. The storage protein in the trunks and roots diminished dramatically after the new shoot had fully developed and the protein had disappeared completely when the cambia of these organs activated vigorously. In the annual growth cycle of Swietenia macrophylla, the storage protein accumulated firstly in the new shoot rather than in the other organs. The protein accumulation began one month after the leaves of new
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