竹秆基本组织分化与功能研究
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摘要
利用细胞生物学研究方法,以毛竹(Phyllostachys edulis)为材料,对竹秆基本组织细胞的分化和发育进行了系统的研究。揭示了基本组织长、短细胞之间的差异,并对基本组织细胞分化发育的机理和长、短细胞生理功能进行了探讨。
根据光学解剖观察,将基本组织细胞发育过程分为原始细胞期、增殖期、伸长期和成熟期。长细胞在增殖期开始伸长,短细胞始终不伸长并一直保持活跃的生理代谢。且离短细胞越远的长细胞越长。这些都表明短细胞具有基本分生组织细胞特点,基本组织可能由其分化发育而形成。
偏光观察表明次生壁在伸长后期开始沉积,次生壁形成于细胞完全停止伸长之前,次生壁的沉积可能导致了细胞伸长减慢并逐渐停止。
基本组织长、短细胞次生壁结构具有明显的差异。长细胞的次生壁随着年龄的增加逐渐增厚,在前四年加厚较快,以后减慢。长细胞在相对休眠期主要沉积木质素,而生长期主要沉积纤维素,从而形成暗的窄层和宽的亮层交替分布的多层结构。木质素含量随着秆龄增加而增加。长细胞的细胞壁结构有利于加强竹秆的机械强度。短细胞仅在与长细胞连接处形成次生壁并木质化,而胞间隙处的短细胞壁始终保持初生壁状态,随着年龄的增加短细胞的壁不加厚,木质素含量也不增加。表明短细胞在发育过程中保持幼嫩细胞的特点,仅在局部形成次生壁。
成熟的长细胞部分细胞器已经降解,而短细胞在发育过程中一直具有较丰富的线粒体、内质网和运输小泡,表明短细胞始终保持较旺盛的生长代谢能力。
长细胞具有贮存功能,竹秆内淀粉粒主要贮存于长细胞内。而短细胞内没有淀粉粒,始终保持比较活跃的生理代谢。
成熟长细胞的ATP酶活性降低。但短细胞始终具有较高的ATP酶活性,尤其是质膜、运输小泡、纹孔和胞间隙中,表明短细胞与周围细胞间频繁的物质、信息交流。短细胞可能参与了细胞间的物质运输,主要通过共质体和质外体两种方式。长细胞壁上的过氧化物酶活性主要集中在次生壁窄层中,以第一年相对休眠期酶活性最高。随着秆龄的增加,长细胞壁上的酶活性降低。长细胞壁上的过氧化物酶参与了木质素的形成。而短细胞壁一直都具有较高酶活性,短细胞过氧化物酶并不完全对应于木质素的沉积部位。短细胞可能参与了活性氧的清除和长细胞次生壁的形成。
The development of ground tissue in bamboo(Phyllostachys edulis) culms were systematically investigated with methods and technologies of cell biology. The marked differences between the long cell and the short cell were elucidated. And the mechanisms of ground tissue differentiation and development and the physiological functions of the long cell and the short cell were discussed.
The results indicated that the development of ground tissue was classified into four stages based on the light microscopy observations: initiating stage, dividing stage, elongation stage and maturation stage. Some cells of the ground tissue began elongation at early dividing stage,but short cells had no elongation and remained active metabolism. And the longer cell was farther from the short cell. The above observations indicated that the short cell was meristematic as the gound meristem cell, and ground tissue might come of the short cell.
The observations by polarized light microscope showed that secondary walls formed at the late stage of elongation, indicating that the secondary walls began to be laid down while the long cells were still undergoing elongation, suggesting that it might act to cause the slow down and eventual cessation of cell elongation.
There were significant differences between the secondary walls of long cells and those of the short cells. The long cell walls underwent thickening with aging. In the former four years, long cell walls had a marked thickening, subsequently, the degree of thickening decreased gradually. The long cell walls formed gradually the polylamellate structure in alternate arrangement of broadly light lamellae with deposition of cellulose in growing period and narrowly dark lamellae, which mainly deposited lignin in relative dormancy period. And lignin in the long cell walls increased with age. It is significant to increase the mechanical strength of the bamboo culms.
But the secondary wall and lignificated regions of the short cells were only confined to the portion in contact with the long cell walls, while the walls at the corner region remained the state of primary wall in mature culms. And there was no thickening, the polylamellate structure and lignin increase with age in the secondary walls of the short cells. The short cell remained in the young state during the development except for the partial secondary wall.
During the development of the gound tissue, mature long cells lost some of the organelles, while the short cell remained plentiful mitochondria, endoplasmic reticulums and transfer vesicles indicating the active metabolism.
The long cell was a reservior where the energy is stored in the form of starch granules. Whereas the short cells remained no starch granule accumulated and had active physiological metabolism.
ATPase activity in mature long cells declined. On the other hand, the short cell remained high ATPase activity, especially in the plasma membrane, transfer vesicles, pits and intercellular spaces, indicating the short cells had active exchange and transport with other cells chiefly by symplastic transport and apolastic transport suggesting that the short cell functioned in transport.
Peroxidase activity concentrated in the narrow lamellae of the long cell walls,where lignin mainly deposited, especially in the first relative dormancy period. The peroxidase activity in the narrow lamellae of the long cell walls declined gradually with aging. So the peroxidase in the long cell walls took part in the synthesis and deposition of lignin. But the short cells differed from the long cells and always had peroxidase activity. Also, Peroxidase in the short cells perhaps took part in the synthesis of lignin in the long cell walls and the elimination of reactive oxygen.
根据光学解剖观察,将基本组织细胞发育过程分为原始细胞期、增殖期、伸长期和成熟期。长细胞在增殖期开始伸长,短细胞始终不伸长并一直保持活跃的生理代谢。且离短细胞越远的长细胞越长。这些都表明短细胞具有基本分生组织细胞特点,基本组织可能由其分化发育而形成。
偏光观察表明次生壁在伸长后期开始沉积,次生壁形成于细胞完全停止伸长之前,次生壁的沉积可能导致了细胞伸长减慢并逐渐停止。
基本组织长、短细胞次生壁结构具有明显的差异。长细胞的次生壁随着年龄的增加逐渐增厚,在前四年加厚较快,以后减慢。长细胞在相对休眠期主要沉积木质素,而生长期主要沉积纤维素,从而形成暗的窄层和宽的亮层交替分布的多层结构。木质素含量随着秆龄增加而增加。长细胞的细胞壁结构有利于加强竹秆的机械强度。短细胞仅在与长细胞连接处形成次生壁并木质化,而胞间隙处的短细胞壁始终保持初生壁状态,随着年龄的增加短细胞的壁不加厚,木质素含量也不增加。表明短细胞在发育过程中保持幼嫩细胞的特点,仅在局部形成次生壁。
成熟的长细胞部分细胞器已经降解,而短细胞在发育过程中一直具有较丰富的线粒体、内质网和运输小泡,表明短细胞始终保持较旺盛的生长代谢能力。
长细胞具有贮存功能,竹秆内淀粉粒主要贮存于长细胞内。而短细胞内没有淀粉粒,始终保持比较活跃的生理代谢。
成熟长细胞的ATP酶活性降低。但短细胞始终具有较高的ATP酶活性,尤其是质膜、运输小泡、纹孔和胞间隙中,表明短细胞与周围细胞间频繁的物质、信息交流。短细胞可能参与了细胞间的物质运输,主要通过共质体和质外体两种方式。长细胞壁上的过氧化物酶活性主要集中在次生壁窄层中,以第一年相对休眠期酶活性最高。随着秆龄的增加,长细胞壁上的酶活性降低。长细胞壁上的过氧化物酶参与了木质素的形成。而短细胞壁一直都具有较高酶活性,短细胞过氧化物酶并不完全对应于木质素的沉积部位。短细胞可能参与了活性氧的清除和长细胞次生壁的形成。
The development of ground tissue in bamboo(Phyllostachys edulis) culms were systematically investigated with methods and technologies of cell biology. The marked differences between the long cell and the short cell were elucidated. And the mechanisms of ground tissue differentiation and development and the physiological functions of the long cell and the short cell were discussed.
The results indicated that the development of ground tissue was classified into four stages based on the light microscopy observations: initiating stage, dividing stage, elongation stage and maturation stage. Some cells of the ground tissue began elongation at early dividing stage,but short cells had no elongation and remained active metabolism. And the longer cell was farther from the short cell. The above observations indicated that the short cell was meristematic as the gound meristem cell, and ground tissue might come of the short cell.
The observations by polarized light microscope showed that secondary walls formed at the late stage of elongation, indicating that the secondary walls began to be laid down while the long cells were still undergoing elongation, suggesting that it might act to cause the slow down and eventual cessation of cell elongation.
There were significant differences between the secondary walls of long cells and those of the short cells. The long cell walls underwent thickening with aging. In the former four years, long cell walls had a marked thickening, subsequently, the degree of thickening decreased gradually. The long cell walls formed gradually the polylamellate structure in alternate arrangement of broadly light lamellae with deposition of cellulose in growing period and narrowly dark lamellae, which mainly deposited lignin in relative dormancy period. And lignin in the long cell walls increased with age. It is significant to increase the mechanical strength of the bamboo culms.
But the secondary wall and lignificated regions of the short cells were only confined to the portion in contact with the long cell walls, while the walls at the corner region remained the state of primary wall in mature culms. And there was no thickening, the polylamellate structure and lignin increase with age in the secondary walls of the short cells. The short cell remained in the young state during the development except for the partial secondary wall.
During the development of the gound tissue, mature long cells lost some of the organelles, while the short cell remained plentiful mitochondria, endoplasmic reticulums and transfer vesicles indicating the active metabolism.
The long cell was a reservior where the energy is stored in the form of starch granules. Whereas the short cells remained no starch granule accumulated and had active physiological metabolism.
ATPase activity in mature long cells declined. On the other hand, the short cell remained high ATPase activity, especially in the plasma membrane, transfer vesicles, pits and intercellular spaces, indicating the short cells had active exchange and transport with other cells chiefly by symplastic transport and apolastic transport suggesting that the short cell functioned in transport.
Peroxidase activity concentrated in the narrow lamellae of the long cell walls,where lignin mainly deposited, especially in the first relative dormancy period. The peroxidase activity in the narrow lamellae of the long cell walls declined gradually with aging. So the peroxidase in the long cell walls took part in the synthesis and deposition of lignin. But the short cells differed from the long cells and always had peroxidase activity. Also, Peroxidase in the short cells perhaps took part in the synthesis of lignin in the long cell walls and the elimination of reactive oxygen.
引文
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