盐生植物耐盐结构特性及植物晶体生物学特征与功能研究
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摘要
采用光学显微技术和石蜡切片相结合的研究方法及其利用晶体的折光方法,对埃及红海沿岸的沙漠地区极端盐渍环境(东经34 o 12′-35 o 27′o,北纬25 o 31′-27 o 22′)的大洋州滨藜(Atriplex nummularia Lindl.)、节状假木贼(Anabasis articulate (Forssk.) Moq.)、藜(Chenopodium album L.)等12种植物,中国吉林松嫩平原一般盐渍环境(东经123o31′-124 o 10′,北纬44 o 30′-44 o 45′)的藜(Chenopodium album L.)、碱地肤(Kochia sieversiana (Pallas) C.A. Mey.)、紫花地丁(Viola yedoensis Makino.)等6种植物,中国吉林长春中生环境(东经125o11′-125 o 32′,北纬43 o 45′-44 o 12′)的碱地肤(Kochia sieversiana (Pallas) C.A. Mey.)、灰绿藜(Chenopodium glaucum L.)、藜(Chenopodium album L.)等19种植物进行植物颉颃盐逆境演化结构比较研究。
结果发现,晶体是植物颉颃盐逆境演化特征,与抗盐成正相关。数理统计量化分析验证了晶体、粘液细胞、贮水组织、发达的角质层等与植物颉颃盐渍逆境关系密切:极端盐渍环境中的大洋洲滨藜皮层具有2-5层连续的晶体,一般盐渍环境中含有晶体最多的滨藜茎皮层中最多只形成接近1层连续的含晶细胞环;极端盐渍环境中的波斯骆驼刺茎表皮下有一层连续的大型粘液细胞层,一般盐渍环境中的植物没发现粘液细胞;极端盐渍环境中的四蕊猪毛菜茎具有发达的贮水组织,一般盐渍环境中的植物没发现贮水组织;极端盐渍环境中生长的中国柽柳茎表皮细胞角质层厚度达6.02μm,一般盐渍环境中碱地肤茎表皮角质层最厚为1.98μm。实验结果显示出极端盐渍环境植物都具有颉颃盐逆境的特殊演化结构。
盐生植物颉颃逆境的演化结构可分为五大类型、两小类型。五大类型为:大多植物具有盐腺、晶体、粘液细胞、贮水结构和发达的同化组织;两小类型为:具有发达的输导组织和机械组织。
研究中发现一种未见报道的可能与物质运输有关的新型复合结构,即:“光芒转运小体”,其结构和功能还有待于进一步研究。
实验结果表明,植物晶体是极端环境盐生植物的典型特征;晶体的存在和含量,为极端盐渍环境、一般盐渍环境和中生环境植物呈递减的趋势;晶体以草酸钙晶体为主,多以大型晶簇的形式存在于薄壁细胞的液泡中;一个细胞一般只形成一个植物晶体。在营养器官中茎是晶体聚集的首要部位,其次是叶,最后是根;根一般不存在植物晶体;植物晶体在盐生植物中的分布具有非均一性,呈典型的多级别区域化形式分布;耐盐植物晶体的分布遵循外向性和功能最优化原则;植物晶体的含量与环境的盐浓度、胁迫时间和生长时间成正相关;以植物晶为耐盐特性的盐生植物,植物晶体区域化的级别越高其耐盐能力越强;改良的天然抗氧化剂类似物显著提高了盐生植物的耐盐水平,但对植物晶体等组织水平的耐盐结构影响不显著。
目前有关晶体的观察只是停留在透射光学显微镜和扫描电镜的观察之中,但是这两种方法都掩盖了晶体的折光特点,本论文首次改革了光学显微镜的晶体观察方法,首次使晶体在细胞内呈现出很强的折光性。
晶体是植物颉颃盐逆境的一种有效的鉴定方法。利用晶体在植物体中存在与否,可缩短抗盐植物的鉴定时间,提高筛选抗盐植物的准确率。
因为具有晶体的植物能有效的改良盐渍生态环境,是生物治理盐渍环境的有效方法。应用具有植物晶体的植物改良盐碱地,既省工、省钱又省力,因此,具有晶体的抗盐植物的开发利用意义重大。
The combination of research methods i.e. optical microscopy, paraffin section and crystal refraction method was carried out to comparative study the evolutive structure of plants antagonism salt adversity. These plants are, Atriplex nummularia Lindl., Anabasis articulate (Forssk.) Moq., Chenopodium album L. etc. 12 kinds of plants on the extreme saline nvironment of the Red Sea coast in the Egypt's desert areas(East longitude 34 o 12′-35 o 27′o,North latitude 25 o 31′-27 o 22′).Chenopodium album L., Kochia sieversiana (Pallas) C.A. Mey., Viola yedoensis Makino., etc. 6 kinds of plants in general saline environment of Songnen plain in Jilin of China(East longitude 123o31′-124 o 10′,North latitude 44 o 30′-44 o 45′). Kochia sieversiana (Pallas) C.A. Mey., Chenopodium glaucum L., Chenopodium album L., etc. 19 kinds of plants in common environment of Changchun in Jilin of China (East longitude 125o11′-125 o 32′,North latitude 43 o 45′-44 o 12′)
It was found that plant crystals are the evolution characteristics of plants antagonism salt, and it is positive correlative with resistance salt. Statistics quantitative analysis verify the crystal, mucous cells, storage, developed corneum were closely relative with plant antagonism saline adversity:There are 2-5 layers of consecutive crystals in the cortex of Atriplex nummularia Lindl. in the extreme saline environment, but in the general saline environment, there is 1 layer of consecutive crystals in the cortex of Chenopodium album L.. There is a layer of consecutive mucus cells under the epidermal of stem of Alhagi maurarum Medic. in the extreme saline environment, on the contrary, in the general saline environment, there is no. There is developed water storage tissues in the stem of Salsola tetrandra Forssk. in the extreme saline environment, on the contrary, in the general saline environment, there is no. The thickness of epidermic cutin cell in the stem of Tamarix chinensis Lour. in the extreme saline environment was up to 6.02μm, but in the general saline environment, the thickness of Kochia sieversiana (Pallas) C.A. Mey. was 1.98μm. The results show that there are special evolution structures in the plants of extreme saline environment to resist the salt adversity.
The structures with typical salt-tolerant features of plant can be divided into five major types and two minor types. The five major types: the salt gland is most, and mainly contains large quantities of plant crystals, a lot of mucus cells, a large number of water storage cells and organizations, developed assimilation organizations. The two minor types are: developed transporting organizations and developed machinery organization.
In addition, a new type of composite structure was found. It was named“light transit body”according to the morphological characteristics and possible function. It maybe implement the function of transport materials. Its structure and function will be further studied.
The results showed that plant crystal is the typical characteristics in extreme environment of halophyte; The existence and content of plant crystal decreased according to the extreme salinity environment, the general salinity environment and the normal environment; Plant crystals, most are oxalate calcium crystals, were mainly in the form of large clusters of crystal in the thin-walled cells; A cell can normally form only one plant crystal. In the vegetative organs, the stem is the most important part of gathering crystals, followed by leaf, and finally the root; Generally, there is no crystal in root; the presence of massive crystal cells in halophytes was a representative structural feature and the salt in plants displayed multi-level compartmentation; The distribution of plant crystals follows the principle of typical outward and optimize function. The content of plant crystal was positively related with the content of environmental salt, the time of salt stress and the age of plant; Halophytes, their characteristics were salt resistance, the higher the level of multi-level compartmentation, the stronger the ability of salt-resistance.
At present, observing the crystals stayed in the level of transmission optical microscope and scanning electron microscopy observation, but these two methods both conceal the refraction characteristics of crystal. This paper for the first time reformed the observing crystal method with optical microscope, so that the crystals in the cells showed strong refraction. The thesis studied in-depth on the chemical structure of crystal. Modified the nalogues of natural anti-oxidants increase salt-tolerant level of halophytes significantly, but it has no significant effect on salt-tolerant structure of plant crystals or other organization.
Crystal is an effective method of identifying the plant antagonism salt stress. According to the crystals exist in the plant or not, the time to identify salt-tolerant plants can be shorten, and the accuracy to choose salt-tolerant plants can be improved.
Because crystal plants can effectively improve the salinity ecological environment, is an effective way to biologically control saline environment.
Application with the crystal plant to improve saline-alkali land can save money and effort, so utilizing the salt-resistant plant with crystals is of great significance.
结果发现,晶体是植物颉颃盐逆境演化特征,与抗盐成正相关。数理统计量化分析验证了晶体、粘液细胞、贮水组织、发达的角质层等与植物颉颃盐渍逆境关系密切:极端盐渍环境中的大洋洲滨藜皮层具有2-5层连续的晶体,一般盐渍环境中含有晶体最多的滨藜茎皮层中最多只形成接近1层连续的含晶细胞环;极端盐渍环境中的波斯骆驼刺茎表皮下有一层连续的大型粘液细胞层,一般盐渍环境中的植物没发现粘液细胞;极端盐渍环境中的四蕊猪毛菜茎具有发达的贮水组织,一般盐渍环境中的植物没发现贮水组织;极端盐渍环境中生长的中国柽柳茎表皮细胞角质层厚度达6.02μm,一般盐渍环境中碱地肤茎表皮角质层最厚为1.98μm。实验结果显示出极端盐渍环境植物都具有颉颃盐逆境的特殊演化结构。
盐生植物颉颃逆境的演化结构可分为五大类型、两小类型。五大类型为:大多植物具有盐腺、晶体、粘液细胞、贮水结构和发达的同化组织;两小类型为:具有发达的输导组织和机械组织。
研究中发现一种未见报道的可能与物质运输有关的新型复合结构,即:“光芒转运小体”,其结构和功能还有待于进一步研究。
实验结果表明,植物晶体是极端环境盐生植物的典型特征;晶体的存在和含量,为极端盐渍环境、一般盐渍环境和中生环境植物呈递减的趋势;晶体以草酸钙晶体为主,多以大型晶簇的形式存在于薄壁细胞的液泡中;一个细胞一般只形成一个植物晶体。在营养器官中茎是晶体聚集的首要部位,其次是叶,最后是根;根一般不存在植物晶体;植物晶体在盐生植物中的分布具有非均一性,呈典型的多级别区域化形式分布;耐盐植物晶体的分布遵循外向性和功能最优化原则;植物晶体的含量与环境的盐浓度、胁迫时间和生长时间成正相关;以植物晶为耐盐特性的盐生植物,植物晶体区域化的级别越高其耐盐能力越强;改良的天然抗氧化剂类似物显著提高了盐生植物的耐盐水平,但对植物晶体等组织水平的耐盐结构影响不显著。
目前有关晶体的观察只是停留在透射光学显微镜和扫描电镜的观察之中,但是这两种方法都掩盖了晶体的折光特点,本论文首次改革了光学显微镜的晶体观察方法,首次使晶体在细胞内呈现出很强的折光性。
晶体是植物颉颃盐逆境的一种有效的鉴定方法。利用晶体在植物体中存在与否,可缩短抗盐植物的鉴定时间,提高筛选抗盐植物的准确率。
因为具有晶体的植物能有效的改良盐渍生态环境,是生物治理盐渍环境的有效方法。应用具有植物晶体的植物改良盐碱地,既省工、省钱又省力,因此,具有晶体的抗盐植物的开发利用意义重大。
The combination of research methods i.e. optical microscopy, paraffin section and crystal refraction method was carried out to comparative study the evolutive structure of plants antagonism salt adversity. These plants are, Atriplex nummularia Lindl., Anabasis articulate (Forssk.) Moq., Chenopodium album L. etc. 12 kinds of plants on the extreme saline nvironment of the Red Sea coast in the Egypt's desert areas(East longitude 34 o 12′-35 o 27′o,North latitude 25 o 31′-27 o 22′).Chenopodium album L., Kochia sieversiana (Pallas) C.A. Mey., Viola yedoensis Makino., etc. 6 kinds of plants in general saline environment of Songnen plain in Jilin of China(East longitude 123o31′-124 o 10′,North latitude 44 o 30′-44 o 45′). Kochia sieversiana (Pallas) C.A. Mey., Chenopodium glaucum L., Chenopodium album L., etc. 19 kinds of plants in common environment of Changchun in Jilin of China (East longitude 125o11′-125 o 32′,North latitude 43 o 45′-44 o 12′)
It was found that plant crystals are the evolution characteristics of plants antagonism salt, and it is positive correlative with resistance salt. Statistics quantitative analysis verify the crystal, mucous cells, storage, developed corneum were closely relative with plant antagonism saline adversity:There are 2-5 layers of consecutive crystals in the cortex of Atriplex nummularia Lindl. in the extreme saline environment, but in the general saline environment, there is 1 layer of consecutive crystals in the cortex of Chenopodium album L.. There is a layer of consecutive mucus cells under the epidermal of stem of Alhagi maurarum Medic. in the extreme saline environment, on the contrary, in the general saline environment, there is no. There is developed water storage tissues in the stem of Salsola tetrandra Forssk. in the extreme saline environment, on the contrary, in the general saline environment, there is no. The thickness of epidermic cutin cell in the stem of Tamarix chinensis Lour. in the extreme saline environment was up to 6.02μm, but in the general saline environment, the thickness of Kochia sieversiana (Pallas) C.A. Mey. was 1.98μm. The results show that there are special evolution structures in the plants of extreme saline environment to resist the salt adversity.
The structures with typical salt-tolerant features of plant can be divided into five major types and two minor types. The five major types: the salt gland is most, and mainly contains large quantities of plant crystals, a lot of mucus cells, a large number of water storage cells and organizations, developed assimilation organizations. The two minor types are: developed transporting organizations and developed machinery organization.
In addition, a new type of composite structure was found. It was named“light transit body”according to the morphological characteristics and possible function. It maybe implement the function of transport materials. Its structure and function will be further studied.
The results showed that plant crystal is the typical characteristics in extreme environment of halophyte; The existence and content of plant crystal decreased according to the extreme salinity environment, the general salinity environment and the normal environment; Plant crystals, most are oxalate calcium crystals, were mainly in the form of large clusters of crystal in the thin-walled cells; A cell can normally form only one plant crystal. In the vegetative organs, the stem is the most important part of gathering crystals, followed by leaf, and finally the root; Generally, there is no crystal in root; the presence of massive crystal cells in halophytes was a representative structural feature and the salt in plants displayed multi-level compartmentation; The distribution of plant crystals follows the principle of typical outward and optimize function. The content of plant crystal was positively related with the content of environmental salt, the time of salt stress and the age of plant; Halophytes, their characteristics were salt resistance, the higher the level of multi-level compartmentation, the stronger the ability of salt-resistance.
At present, observing the crystals stayed in the level of transmission optical microscope and scanning electron microscopy observation, but these two methods both conceal the refraction characteristics of crystal. This paper for the first time reformed the observing crystal method with optical microscope, so that the crystals in the cells showed strong refraction. The thesis studied in-depth on the chemical structure of crystal. Modified the nalogues of natural anti-oxidants increase salt-tolerant level of halophytes significantly, but it has no significant effect on salt-tolerant structure of plant crystals or other organization.
Crystal is an effective method of identifying the plant antagonism salt stress. According to the crystals exist in the plant or not, the time to identify salt-tolerant plants can be shorten, and the accuracy to choose salt-tolerant plants can be improved.
Because crystal plants can effectively improve the salinity ecological environment, is an effective way to biologically control saline environment.
Application with the crystal plant to improve saline-alkali land can save money and effort, so utilizing the salt-resistant plant with crystals is of great significance.
引文
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