海马齿盐适应细胞形态、结构、蛋白变化及耐盐基因功能初步研究
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摘要
海马齿(Sesuvium portulacastrum L.)是一种生长在热带、亚热带盐滩涂地的肉质草本植物,在淡水栽培和海水栽培条件下均能生长良好,并完成生活史。
本研究以淡水栽培和海水栽培3个月的海马齿植物为材料,从叶和茎的显微形态结构变化、叶肉细胞超微结构、蛋白质积累、离子微区分布、比较蛋白质组学等对海马齿植物的耐盐适应性进行了比较。并从海马齿植物的基因组中分离了三个与耐盐相关的新基因(STRG152)的同源基因,并对其功能在酵母菌和遗传模式植物拟南芥中进行了初步的分析。结果如下：
1.海马齿在海水栽培条件下,叶片增厚,叶表皮细胞层中薄壁细胞体积变大,且向外凸出形成泡状细胞。我们推测这些细胞主要是吸收水分,稀释被运输到叶片表皮层的盐浓度。
2.海水栽培的海马齿叶肉细胞质膜明显向内折叠,出现大量大小、形状各异的质膜突起,以及质膜片层；而淡水栽培的海马齿叶肉细胞质膜向内折叠不明显,质膜突起少见。
3.海水栽培的海马齿植物叶肉细胞叶绿体变小、数量增多；形状变短,由肾形、梭形或弓形变成椭圆形或一端膨大的不规则形状；叶绿体基粒片层结构清晰完整,垛叠程度增加,叶绿体没有受到明显伤害；叶绿体中淀粉粒数量增多,体积变大,淀粉粒表面出现皱褶,形状由长椭圆型变成短椭圆形或不规则形状,电子云密度变低；叶绿体上脂质体增多且体积变大。
4.海水栽培的海马齿叶肉细胞线粒体数目增加,体积变小,形状由淡水处理的棒状和球状变成椭圆状和球状,线粒体结构保持完整,线粒体嵴清晰,外膜模糊,受到伤害；而淡水栽培的海马齿叶肉细胞线粒体轮廓清晰,嵴清晰。
5.淡水栽培的海马齿植物茎中Na元素比叶中Na元素要高,而海水栽培的海马齿叶片中的Na元素比茎中Na元素要高；在盐胁迫下,Na元素在叶片的表皮、栅栏组织和贮水组织中均大量增加,各部分增加的比例基本相等；而海水栽培的海马齿茎中Na元素相对于淡水栽培的海马齿茎中的Na元素也有所增加,各部分增加的比例髓细胞相对于表皮和维管束系统要高。
6.海马齿植物,Na+的毒害,没有减少植物对K+离子的吸收,反而增加了它在海马齿茎中的含量,所以海马齿植物在盐胁迫条件下,植物不会因为缺K+而受到伤害。
7.盐胁迫条件能使氯元素含量在茎和叶Na元素增加的组织中增加较多；在茎的微管系统中基本保持不变,但在茎的髓细胞和叶的表皮细胞层中Cl元素增加较多。我们推测氯元素在海马齿中的作用为中和阳离子,增加渗透压使细胞吸水,促进海马齿叶片肉质化。
8.海水栽培的海马齿茎中的薄壁细胞颗粒状和膜状物质增多；蛋白质类物质在叶片中各组织中积累增加,但在茎中增加不是很明显。
9.淡水栽培和海水栽培的海马齿植物差异蛋白点以表达量正负2倍为标准,海水栽培的海马齿蛋白质表达有约13.3%的下调,有约15.7%的上调,通过MALD-TOF-MS和MALD-TOF-TOF-MS分析表达差异在±2.5倍的蛋白质点,共鉴定出73个蛋白质,这些蛋白质可以分成11类。(ⅰ)8个胁迫相关蛋白；(ⅱ)25个光合作用相关蛋白；(ⅲ)6个能量代谢相关蛋白；(ⅳ)10个碳代谢相关蛋白；(ⅴ)4个转录、翻译相关蛋白；(ⅵ)4个抗氧化酶类；(ⅶ)6个信号转导相关蛋白；(ⅷ)2个细胞骨架蛋白；(ⅸ)2个生长发育相关蛋白；(ⅹ)5个功能不清楚蛋白；(Ⅺ)1个核酸代谢相关蛋白。
10.从海马齿植物基因组中分离了3个STRG152同源基因,命名为SRTG152-Ⅰ, SRTG152-Ⅱ和SRTG152-Ⅲ并登陆(GenBank登录号为：FJ457924,FJ457925和FJ457926),对其中的一个基因SRTG152-Ⅰ在酵母菌和遗传模式植物拟南芥中进行了初步的功能分析,初步判断该基因与抗高浓度K+有关。
Sesuvium portulacastrum L. is a kind of succulent herb growing in tropical and subtropical intertidal land. It grows well and finishs life cycle when it is cultivated in both fresh and sea water. In this study, the seedlings of S. portulacastrum watered with fresh water and sea water for 3 months were used as the materials to study their adaptable characteristics in response to salt tolerances, which included morphological structure, mesophyll cell ultrastructure, protein accumulation, the ion micro-distriction and protein expression. In addition, we isolated three STRG152 homologous genes from S. portulacastrum and preliminarily analysis their function in yeast and Arabidopsis. The results are as follows:
1. When the plants of S. portulacastrum were watered with sea water, the leaves were thicken, and the leaf epidermal cells consisted of the bubble-like parenchyma cells that were enlarged and bulged from the surface. It is deduced that the main function of these cells was for water-absorbing from outside to reduce the cell osmotic pressure, which they are useful to reduce the salt concentration in plant vivo.
2. When the plants of S. portulacastrum were watered with the sea water, the plasma membrane of mesophyll cells were clearly folded inward, and formed many of different sizes and shapes of protrusions and lamellaes in their plasma membrane; However when the plants of S. portulacastrum were cultivated in the fresh water, the folding inward of plasma membrane in mesophyll cells was not so obvious.
3. When the plants of S. portulacastrum were watered with sea water, the chloroplasts in mesophyll cells were smaller with shorted shape, but the number of them was increased. The shape of chloroplasts was changed from shapes of kidney, spindle or arciform to ellipse or irregular. The grana lamella of chloroplasts was integrity and clear, and the stacking degree was increased. The chloroplasts had not been significantly hurt. The number of starch grains in chloroplasts was increased, and the size was enlarged. The surface of starch grains appeared wrinkly, and the shape was changed from oblong to short oval or irregular. Electron density was become lower. The number of liposomes on chloroplast was increased, and the size was enlarged.
4. When the plants of S. portulacastrum were watered with sea water, the mitochondria in mesophyll cells were smaller with shorted shape, but the number of them was increased. The shape was changed from the shapes of stick and sphere to ellipse and sphere. The construct of mitochondria was integrity. The mitochondrial cristaes were clear, but the mitochondria membrane was fuzzy and hurt. When the plants of S. portulacastrum were watered with fresh water, the mitochondria were line profile and the mitochondrial cristaes were clear.
5. The Na element in the stem was higher than that in leaf of S. portulacastrum when it was watered with fresh water, but Na element in the leaf was higher than that in the stem of S. portulacastrum when it was watered with sea water. Under salt stress, Na element sharply increased in the leaf tissues of epidermal cells, palisade tissue and water organization, and the increasing proportion among them was almost equal. Na element in stem of S. portulacastrum watered with sea water was slightly hgher than that of S. portulacastrum watered with sea water, and the increasing proportion among the stem tissues was higher in the pith cells than epidermal and vascular system.
6. The Na+ poison didn't decrease the absorption of K+ in S. portulacastrum vivo, in contrast to it, the K+ content was increased when the plants were watered with sea water, so they was not hurt because of lacking K+ under salt-stress.
7. Under salt stress, the Cl element was much increased in the parts of leaf and stem, in which Na element was increased. Cl element was almost the similar level in vascular system of stem, but the Cl element is higher in pith and epidermal systems when the plants were watered with sea water. It is deduced that Cl element plays a role in cation neutralization; osmotic pressure addition, and leaf succulence.
8. When the plants of S. portulacastrum were watered with sea water, the granular materials and membranous materials were increased in the stem parenchyma cells, and protein accumulationwas increased in all tissue of leaf, but not clearly increased in stem.
9. With plus or minus 2 fold protein expression as a standard,13.3% down-regulation protein dots, and 15.7% up-regulation dots were observed when the plants of S. portulacastrum were watered with sea water.73 protein dots with plus or minus 2.5 fold protein expressing have been analyzed by Using MALDI-TOF-MS and MALDI-TOF-TOF-MS. These protein dots can be divided into 11 categories. (ⅰ) 8 protein dots belong to stress responses and defense related proteins; (ⅱ) 25 protein dots belonge to photosynthesis related proteins; (ⅲ) 6 protein dots belong to energy metabolism related proteins; (ⅳ)10 protein dots belong to carbohydryate metabolism associated proteins; (ⅴ) 4 protein dots belong to transcription, translation and trafficking; (ⅵ) 4 protein dots belong to detoxifying and antioxidant enzymes; (ⅶ) 6 protein dots belong to signal transduction related proteins; (ⅷ) 2 protein dots belong to cytoskeleton dynamics; (ⅸ) 2 protein dots belong to growth and development related proteins; (ⅹ) 5 protein dots belong to function unknown and hypothetical proteins; (Ⅺ) 1 protein dot belongs to nucleotide acid metabolism protein.
10. We cloned tree homologous genes of STRG152 from S. portulacastrum andnamed SRTG152-Ⅰ, SRTG152-Ⅱand SRTG152-Ⅲ(GenBank accession:FJ457924, FJ457925 and FJ457926). We preliminarily analyzed the function of SRTG152-Ⅰgene in Yeast and Arabidopsis. SRTG152-Ⅰgene was preliminarily identified to relate high K+ tolerance.
本研究以淡水栽培和海水栽培3个月的海马齿植物为材料,从叶和茎的显微形态结构变化、叶肉细胞超微结构、蛋白质积累、离子微区分布、比较蛋白质组学等对海马齿植物的耐盐适应性进行了比较。并从海马齿植物的基因组中分离了三个与耐盐相关的新基因(STRG152)的同源基因,并对其功能在酵母菌和遗传模式植物拟南芥中进行了初步的分析。结果如下：
1.海马齿在海水栽培条件下,叶片增厚,叶表皮细胞层中薄壁细胞体积变大,且向外凸出形成泡状细胞。我们推测这些细胞主要是吸收水分,稀释被运输到叶片表皮层的盐浓度。
2.海水栽培的海马齿叶肉细胞质膜明显向内折叠,出现大量大小、形状各异的质膜突起,以及质膜片层；而淡水栽培的海马齿叶肉细胞质膜向内折叠不明显,质膜突起少见。
3.海水栽培的海马齿植物叶肉细胞叶绿体变小、数量增多；形状变短,由肾形、梭形或弓形变成椭圆形或一端膨大的不规则形状；叶绿体基粒片层结构清晰完整,垛叠程度增加,叶绿体没有受到明显伤害；叶绿体中淀粉粒数量增多,体积变大,淀粉粒表面出现皱褶,形状由长椭圆型变成短椭圆形或不规则形状,电子云密度变低；叶绿体上脂质体增多且体积变大。
4.海水栽培的海马齿叶肉细胞线粒体数目增加,体积变小,形状由淡水处理的棒状和球状变成椭圆状和球状,线粒体结构保持完整,线粒体嵴清晰,外膜模糊,受到伤害；而淡水栽培的海马齿叶肉细胞线粒体轮廓清晰,嵴清晰。
5.淡水栽培的海马齿植物茎中Na元素比叶中Na元素要高,而海水栽培的海马齿叶片中的Na元素比茎中Na元素要高；在盐胁迫下,Na元素在叶片的表皮、栅栏组织和贮水组织中均大量增加,各部分增加的比例基本相等；而海水栽培的海马齿茎中Na元素相对于淡水栽培的海马齿茎中的Na元素也有所增加,各部分增加的比例髓细胞相对于表皮和维管束系统要高。
6.海马齿植物,Na+的毒害,没有减少植物对K+离子的吸收,反而增加了它在海马齿茎中的含量,所以海马齿植物在盐胁迫条件下,植物不会因为缺K+而受到伤害。
7.盐胁迫条件能使氯元素含量在茎和叶Na元素增加的组织中增加较多；在茎的微管系统中基本保持不变,但在茎的髓细胞和叶的表皮细胞层中Cl元素增加较多。我们推测氯元素在海马齿中的作用为中和阳离子,增加渗透压使细胞吸水,促进海马齿叶片肉质化。
8.海水栽培的海马齿茎中的薄壁细胞颗粒状和膜状物质增多；蛋白质类物质在叶片中各组织中积累增加,但在茎中增加不是很明显。
9.淡水栽培和海水栽培的海马齿植物差异蛋白点以表达量正负2倍为标准,海水栽培的海马齿蛋白质表达有约13.3%的下调,有约15.7%的上调,通过MALD-TOF-MS和MALD-TOF-TOF-MS分析表达差异在±2.5倍的蛋白质点,共鉴定出73个蛋白质,这些蛋白质可以分成11类。(ⅰ)8个胁迫相关蛋白；(ⅱ)25个光合作用相关蛋白；(ⅲ)6个能量代谢相关蛋白；(ⅳ)10个碳代谢相关蛋白；(ⅴ)4个转录、翻译相关蛋白；(ⅵ)4个抗氧化酶类；(ⅶ)6个信号转导相关蛋白；(ⅷ)2个细胞骨架蛋白；(ⅸ)2个生长发育相关蛋白；(ⅹ)5个功能不清楚蛋白；(Ⅺ)1个核酸代谢相关蛋白。
10.从海马齿植物基因组中分离了3个STRG152同源基因,命名为SRTG152-Ⅰ, SRTG152-Ⅱ和SRTG152-Ⅲ并登陆(GenBank登录号为：FJ457924,FJ457925和FJ457926),对其中的一个基因SRTG152-Ⅰ在酵母菌和遗传模式植物拟南芥中进行了初步的功能分析,初步判断该基因与抗高浓度K+有关。
Sesuvium portulacastrum L. is a kind of succulent herb growing in tropical and subtropical intertidal land. It grows well and finishs life cycle when it is cultivated in both fresh and sea water. In this study, the seedlings of S. portulacastrum watered with fresh water and sea water for 3 months were used as the materials to study their adaptable characteristics in response to salt tolerances, which included morphological structure, mesophyll cell ultrastructure, protein accumulation, the ion micro-distriction and protein expression. In addition, we isolated three STRG152 homologous genes from S. portulacastrum and preliminarily analysis their function in yeast and Arabidopsis. The results are as follows:
1. When the plants of S. portulacastrum were watered with sea water, the leaves were thicken, and the leaf epidermal cells consisted of the bubble-like parenchyma cells that were enlarged and bulged from the surface. It is deduced that the main function of these cells was for water-absorbing from outside to reduce the cell osmotic pressure, which they are useful to reduce the salt concentration in plant vivo.
2. When the plants of S. portulacastrum were watered with the sea water, the plasma membrane of mesophyll cells were clearly folded inward, and formed many of different sizes and shapes of protrusions and lamellaes in their plasma membrane; However when the plants of S. portulacastrum were cultivated in the fresh water, the folding inward of plasma membrane in mesophyll cells was not so obvious.
3. When the plants of S. portulacastrum were watered with sea water, the chloroplasts in mesophyll cells were smaller with shorted shape, but the number of them was increased. The shape of chloroplasts was changed from shapes of kidney, spindle or arciform to ellipse or irregular. The grana lamella of chloroplasts was integrity and clear, and the stacking degree was increased. The chloroplasts had not been significantly hurt. The number of starch grains in chloroplasts was increased, and the size was enlarged. The surface of starch grains appeared wrinkly, and the shape was changed from oblong to short oval or irregular. Electron density was become lower. The number of liposomes on chloroplast was increased, and the size was enlarged.
4. When the plants of S. portulacastrum were watered with sea water, the mitochondria in mesophyll cells were smaller with shorted shape, but the number of them was increased. The shape was changed from the shapes of stick and sphere to ellipse and sphere. The construct of mitochondria was integrity. The mitochondrial cristaes were clear, but the mitochondria membrane was fuzzy and hurt. When the plants of S. portulacastrum were watered with fresh water, the mitochondria were line profile and the mitochondrial cristaes were clear.
5. The Na element in the stem was higher than that in leaf of S. portulacastrum when it was watered with fresh water, but Na element in the leaf was higher than that in the stem of S. portulacastrum when it was watered with sea water. Under salt stress, Na element sharply increased in the leaf tissues of epidermal cells, palisade tissue and water organization, and the increasing proportion among them was almost equal. Na element in stem of S. portulacastrum watered with sea water was slightly hgher than that of S. portulacastrum watered with sea water, and the increasing proportion among the stem tissues was higher in the pith cells than epidermal and vascular system.
6. The Na+ poison didn't decrease the absorption of K+ in S. portulacastrum vivo, in contrast to it, the K+ content was increased when the plants were watered with sea water, so they was not hurt because of lacking K+ under salt-stress.
7. Under salt stress, the Cl element was much increased in the parts of leaf and stem, in which Na element was increased. Cl element was almost the similar level in vascular system of stem, but the Cl element is higher in pith and epidermal systems when the plants were watered with sea water. It is deduced that Cl element plays a role in cation neutralization; osmotic pressure addition, and leaf succulence.
8. When the plants of S. portulacastrum were watered with sea water, the granular materials and membranous materials were increased in the stem parenchyma cells, and protein accumulationwas increased in all tissue of leaf, but not clearly increased in stem.
9. With plus or minus 2 fold protein expression as a standard,13.3% down-regulation protein dots, and 15.7% up-regulation dots were observed when the plants of S. portulacastrum were watered with sea water.73 protein dots with plus or minus 2.5 fold protein expressing have been analyzed by Using MALDI-TOF-MS and MALDI-TOF-TOF-MS. These protein dots can be divided into 11 categories. (ⅰ) 8 protein dots belong to stress responses and defense related proteins; (ⅱ) 25 protein dots belonge to photosynthesis related proteins; (ⅲ) 6 protein dots belong to energy metabolism related proteins; (ⅳ)10 protein dots belong to carbohydryate metabolism associated proteins; (ⅴ) 4 protein dots belong to transcription, translation and trafficking; (ⅵ) 4 protein dots belong to detoxifying and antioxidant enzymes; (ⅶ) 6 protein dots belong to signal transduction related proteins; (ⅷ) 2 protein dots belong to cytoskeleton dynamics; (ⅸ) 2 protein dots belong to growth and development related proteins; (ⅹ) 5 protein dots belong to function unknown and hypothetical proteins; (Ⅺ) 1 protein dot belongs to nucleotide acid metabolism protein.
10. We cloned tree homologous genes of STRG152 from S. portulacastrum andnamed SRTG152-Ⅰ, SRTG152-Ⅱand SRTG152-Ⅲ(GenBank accession:FJ457924, FJ457925 and FJ457926). We preliminarily analyzed the function of SRTG152-Ⅰgene in Yeast and Arabidopsis. SRTG152-Ⅰgene was preliminarily identified to relate high K+ tolerance.
引文
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