五种C4荒漠植物光合器官中含晶细胞的比较分析

为了探讨荒漠植物适应干旱环境的机理,选择光合器官发生很大变化的5种C4荒漠植物进行了解剖结构的对比研究。结果表明,这5种植物中含晶细胞的数量、大小、形态和分布位置等存在差异。白梭梭（Haloxylon persicum）和梭梭（H.ammodendron）的同化枝普遍具有含晶细胞;沙拐枣（Calligonum mongolicum）的含晶细胞很少,一般只分布在贮水组织或靠近栅栏组织处;木本猪毛菜（Salsola arbuscula）的含晶细胞也不多,主要分布在栅栏组织和表皮细胞之间;猪毛菜（S.collina）的含晶细胞更少,仅在贮水组织中偶尔可见晶簇。比较梭梭、白梭梭和沙拐枣同化枝不同部位的解剖结构发现,梭梭同化枝基部含晶细胞最多,中部次之,顶部最少;白梭梭同化枝项部的含晶细胞数量较多,中部及基部较少;沙拐枣同化枝顶部与基部的粘液细胞较多,中部较少,基部几乎没有栅栏组织,而其维管组织较为发达。综合晶体的酸碱溶解性及硝酸银组化分析结果,并参照能谱仪的分析结果得知,梭梭、白梭梭、沙拐枣和木本猪毛菜的叶片或同化枝中所含晶体的主要成分为草酸钙。通过比较解剖结构发现,梭梭和白梭梭的同化枝中含晶细胞最多,其它3种植物的同化器官中含晶细胞较少,而沙拐枣同化枝中有粘液细胞存在。     

荒漠植物梭梭和沙拐枣的花环结构及C4光合特征

 为了探讨梭梭（Haloxylon ammodendron）和沙拐枣（Calligonum mongolicum）适应高温强光荒漠环境的光合作用机构及特征,通过对其同化枝的解剖结构观察,δ13C值分析,以及气体交换测定表明：二者均具有花环结构(Kranz anatomy),肉细胞（Mesophyll cell）呈栅栏状,其内侧是维管束鞘细胞（Bundle sheath cell）,小维管束与维管束鞘细胞相接。在栅栏组织和贮水组织中,梭梭具有形状巨大的含晶细胞;沙拐枣具有大量的粘液细胞。梭梭和沙拐枣同化枝的δ13C值分别为-14.3‰和-14.8‰,在不同生长季节和土壤水分条件下,二者的δ13C值变化在-14‰到-16‰之间。梭梭和沙拐枣的CO2补偿点分别为2 μmol•mol－1和4 μmol•mol－1,光饱和点分别为1 660和1 756 μmol•m－2•s－1,表观光合量子效率分别为0.044和0.057 mol CO2•mol－1 photons。这表明,广泛分布于我国荒漠地区的木本植物梭梭和沙拐枣为C4植物,其光合途径不随生长季节和水分条件的变化而改变。     

不同梭梭种群同化枝的解剖结构特征及其与生态因子的关系分析

该研究于中国西北干旱区按自然降水梯度从东向西选择6个梭梭种群,取其当年生同化枝作为试验材料,采用石蜡切片法,应用光学显微技术测定同化枝9项解剖结构指标,分析不同生境下梭梭同化枝解剖结构的变异特点以及与生态因子的关系,为干旱区生态环境保护和梭梭资源的保护与利用提供理论依据。结果表明:(1)干旱荒漠环境的梭梭同化枝解剖结构具有连续的栅栏组织、花环结构、含晶细胞和贮水组织等结构特征,但其表皮细胞、角质层厚度、气孔位置和气下室与旱生植物结构有差异,主要表现为表皮细胞和角质层较薄,气孔半下陷且气下室不发达。(2)梭梭同化枝各种群间的9项解剖结构指标都存在显著差异,其中变异系数较大的是维管束个数/直径和角质层厚度,变异系数分别为22.78%和15.20%;相关分析表明:同化枝直径、栅栏细胞切向长、贮水组织厚度和维管柱直径均与经度、纬度、海拔、一月均温、七月均温、年均温、年降水量和年相对湿度等8个生态因子有不同程度的相关性,其中与经度、海拔、七月均温和年平均相对湿度显著相关。(3)6个梭梭种群通过聚类分析可分为两大类,分析结果与所观察到的生境类型、地理分布的划分基本一致;随着自然降水量自东向西减少,梭梭的抗旱性相应逐渐增强。     

两种盐生植物解剖结构的生态适应性

结合对研究区域土壤水分和盐分的分析,对3种不同生境下梭梭和多枝柽柳两种荒漠盐生植物光合器官解剖结构的研究表明:梭梭(Haloxylon ammodendron)依靠当年生绿色同化枝进行光合作用,且同化枝具有发达的贮水组织,是典型的超旱生稀盐盐生植物;同化枝的栅栏组织富含叶绿体,是C_4高光效植物,提高了植物的光合作用效率,进一步增强了梭梭对荒漠干旱、盐渍化环境的适应能力.多枝柽柳(Tamarix ramosissima)同化枝及叶的表皮上均具下陷的气孔和泌盐腺,是旱生泌盐盐生植物;叶为全栅型,且同化枝具有发达的维管柱,占同化枝直径的60%以上,此外,其同化枝及叶的表皮细胞外凸形成乳状突,是一种环境胁迫指示结构.这些特征均能说明梭梭和多枝柽柳具有很好的抗旱、耐盐碱能力,且作为荒漠环境的优势树种,它们对荒漠生态系统的恢复和重建有积极意义.     

C4荒漠植物猪毛菜与木本猪毛菜的叶片解剖结构及光合生理特征

 为了比较C₄荒漠植物猪毛菜(Salsola collina)和木本猪毛菜(S. arbuscula)的抗旱结构和适应环境的光合作用特征, 在二者混生的群落中, 选择代表性植株, 采集叶片进行叶片解剖结构分析, 在自然条件下测定了二者叶片的气体交换参数。研究结果表明：猪毛菜叶片具表皮毛, 具有更发达的薄壁贮水组织;木本猪毛菜叶片具有更厚的角质层, 表皮下有1层下皮细胞, 其栅栏组织细胞较长, 排列更紧密。猪毛菜的净光合速率明显高于木本猪毛菜, 日平均值分别为21.5和15.7 μmol CO₂·m^–2·s^–1。猪毛菜的蒸腾速率也明显高于木本猪毛菜, 日平均值分别为14.9和10.2 mmol·m^–2·s^–1。猪毛菜和木本猪毛菜的水分利用效率的日平均值分别为1.39和1.53 μmol CO₂·mmol^–1 H₂O, 特别是在14:00时分别为1.61和2.30 μmol CO₂·mmol^–1 H₂O, 木本猪毛菜高出猪毛菜约42%。猪毛菜的光补偿点低于木本猪毛菜, 而光饱和点和光量子效率较高, 具有更低的CO₂补偿点。这表明：二者的旱生结构不同, 木本猪毛菜具有更显著的荒漠植物特征;在适于二者混生的环境下, 猪毛菜比木本猪毛菜的光合能力更强, 而木本猪毛菜的水分利用效率更高。     

新疆10种藜科植物叶片和同化枝的旱生和盐生结构的研究

 本文应用扫描电镜和光学显微镜对生长在新疆荒漠地区10种藜科植物中亚滨藜(Atriplex centralasiatica),心叶驼绒藜(Ceratoides ewersmanniana),驼绒藜(Ceratoides latens),盐节木(Halocnemum strobilaceum),盐穗木(Halostachys caspica),梭梭(Haloxylon ammodendron),圆叶盐爪爪(Kalidium schrenkianum),绒藜(Londesia eriantha),费尔干猪毛菜(Salsola ferganica),浆果猪毛菜(Salsola foliosa)的叶和同化枝进行了形态解剖学研究。结果表明,它们是通过以下结构来适应旱生和盐生环境的：叶片及角质膜厚,气孔器下陷,具表皮毛;栅栏组织发达,多为等面叶;部分植物叶片退化成鳞片状,而由同化枝执行光合功能;多数植物叶片和同化枝内部具有粘液和含晶细胞,贮水组织发达。根据盐分是否排出体外,又划分出聚盐和泌盐植物。在泌盐植物中,盐腺具有单细胞和多细胞及分泌孔类型,并对其聚盐和泌盐机理作了初步探讨。     

C4荒漠植物猪毛菜与木本猪毛菜的叶片解剖结构及光合生理特征

为了比较C₄荒漠植物猪毛菜(Salsola collina)和木本猪毛菜(S. arbuscula)的抗旱结构和适应环境的光合作用特征, 在二者混生的群落中, 选择代表性植株, 采集叶片进行叶片解剖结构分析, 在自然条件下测定了二者叶片的气体交换参数。研究结果表明：猪毛菜叶片具表皮毛, 具有更发达的薄壁贮水组织;木本猪毛菜叶片具有更厚的角质层, 表皮下有1层下皮细胞, 其栅栏组织细胞较长, 排列更紧密。猪毛菜的净光合速率明显高于木本猪毛菜, 日平均值分别为21.5和15.7 μmol CO₂·m^-2·s^-1。猪毛菜的蒸腾速率也明显高于木本猪毛菜, 日平均值分别为14.9和10.2 mmol·m^-2·s^-1。猪毛菜和木本猪毛菜的水分利用效率的日平均值分别为1.39和1.53 μmol CO₂·mmol^-1H₂O, 特别是在14:00时分别为1.61和2.30 μmol CO₂·mmol^-1H₂O, 木本猪毛菜高出猪毛菜约42%。猪毛菜的光补偿点低于木本猪毛菜, 而光饱和点和光量子效率较高, 具有更低的CO₂补偿点。这表明：二者的旱生结构不同, 木本猪毛菜具有更显著的荒漠植物特征;在适于二者混生的环境下, 猪毛菜比木本猪毛菜的光合能力更强, 而木本猪毛菜的水分利用效率更高。     

不同年代梭梭同化枝解剖结构特征及其对土壤条件的响应

叶片是植物与外界环境进行能量、水气交换的主要器官,对环境变化较敏感且可塑性较大。该研究以民勤不同年代梭梭同化枝为研究对象,采用石蜡切片法,分析不同年代梭梭同化枝解剖结构变异特点及其与土壤条件的关系,旨在探讨梭梭同化枝在干旱环境的生态适应机制。结果表明:(1)除角质层和导管孔径,不同年代梭梭同化枝解剖结构指标差异显著(P<0.05),变异系数变化范围为5.19%~21.53%。(2)随着生长年限的增加,梭梭同化枝直径变小,其他解剖结构指标占半径比例呈先增加后降低趋势。(3)土壤养分含量随年代的增加表现为不断降低的趋势,土壤含水率则因地点不同表现出较大差异。(4)角质层、栅栏组织与速效磷含量呈显著负相关(P<0.05);维管柱直径与电导率呈显著负相关(P<0.05)。研究认为,梭梭同化枝解剖结构与土壤条件存在密切关系,通过改变叶片结构更好地适应干旱荒漠环境,以期为该区梭梭资源的保护与利用提供理论依据。     

沙拐枣属植物同化枝和子叶的解剖学研究

采用石蜡切片技术,对相同生境下的戈壁沙拐枣、柴达木沙拐枣和英吉沙沙拐枣的子叶,及中国特有种柴达木沙拐枣、塔里木沙拐枣、阿拉善沙拐枣和英吉沙沙拐枣同化枝的横切面解剖结构进行观察研究.结果显示:(1)3种沙拐枣子叶均为全栅等面叶;表皮为单层细胞,上下表皮均分布气孔,栅栏组织以内为一圈连续排列的维管束鞘细胞;中脉和小叶脉排列近似M型,包埋于薄壁细胞中.(2)塔里木沙拐枣同化枝的保护组织、栅栏组织较发达,英吉沙沙拐枣同化枝机械组织和输导组织较为发达,阿拉善沙拐枣同化枝的栅栏组织较为发达,相比较柴达木沙拐枣同化枝的保护组织、机械组织和输导组织最不发达.(3)柴达木沙拐枣和英吉沙沙拐枣中子叶与同化枝的横切面各组织分布相似,主要差异为横切面形状、维管束排列方式和机械组织发达程度.研究表明,子叶与同化枝的解剖结构是该属植物在干旱环境下长期适应的结果,解剖结构的差异体现了种间、器官间对相同环境的不同适应策略.     

我国荒漠不同生态类型植物的旱生结构

本文研究了我国荒漠15种不同类型植物的解剖结构。结果表明,超旱生和旱生植物具有典型的旱生结构,尤以超旱生植物明显,例如沙冬青、梭梭。 旱生结构的基本特征是：叶片及角质层厚,具表皮毛、腺毛、瘤状或乳状突起,气孔下陷,栅栏组织及贮水组织发达,构成等面叶及肉质叶,具含晶细胞和粘液细胞;维管束鞘、维管束帽及纤维细胞的存在,增强了植株的坚固性。中生植物叶片及角质层较薄：气孔平置或拱起,栅栏组织及海绵组织同等发育,胞间隙大,为背腹叶,这是典型的中生结构。     

两种梭梭根部输导组织坚韧度及其抗旱性

以梭梭和白梭梭一年生盆栽幼苗为试材,测定60％（对照）、40％和20％的土壤相对含水量（sRwc）处理20d后两种梭梭同化枝的电导率和含水量,地上和地下部水势,根部木质素、纤维素、半纤维素含量,根肉质化程度和根长度。结果表明：两种梭梭同化枝含水量随着SRWC的下降均保持较高的水平;SRWC为40％和20％时,两种同化枝电导率的变化不显著,且均保持较低的值;两种梭梭地下部与地上部水势差值随土壤含水量的降低而增大;SRWC为40％的土壤条件促进两种梭梭的根系生长,20％的SRWC条件下仍保持与对照一样的水平;不同SRWC条件下,梭梭和白梭梭根部的木质素、纤维素、半纤维素含量的变化幅度均较小,且保持很高的水平,总含量分别为46．9％-53．3％和50．6％-57．6％。由此推断,在干旱胁迫下两种梭梭的根系依赖于较强的根部榆导组织坚韧度,往土壤深层扎根找水,适应干旱环境。     

Anatomical Observation of Assimilating Branches of Nine Xerophytes in Gansu

The nine xerophytes grown in Gansu, i.e. Haloxylon persicum Bge., H. ammodendron Bge., Calligonum junceum Litw., C. arborescens Litw., C. leucocladum Bge., C. mongolicum Turcz., C. caput-medusae Schrenk, Hedysarum scoparium Fisch. et Mey. and Limonium aureum Hill., all, especially Haloxylon persicum, possessed distinct structural features of xeromorphism. Generally their leaves were extremely degenerated or exist as basal leaves. Photosynthesis was carried out by the young green twigs. The epidermis of which had thickened cell walls and were overlaid with thick cuticle. Underneath the epidermis it was the hypodermis. Inner to the hypodermis, the well developed palisade layer was shown to have cells containing abundant chloroplasts. Water-storing tissues with large cells were seen central to the palisade tissue. In tran- sverse section, the ratio of the cortex width and stem diameter was greater than that of mesophytes. In contrast, the ratio of cortex width and root diameter was smaller. Crystal cells or mucilage cells probably of xeromorphic charateristics were generally present in these xerophyes.     

Comparative Studies on Relation of Photosynthesis to Water Status of Two Species of Haloxylon Under Controlled Environments

The net photosynthetic rate,transpiration rate and water content of assimilating shoots of Haloscyion ammodendron and H. persicum were measured under different radiation intensities, temperatures changes and different water status. Their water potential under different water status was also measured. The experimental results showed. (1) The dependence of photosynthetic rate and transpiration rate on radiation intensit in the two species of Haloscylon. The increase of net photosynthetic rate and transpiration rate of H. arnrnodendron depending on radiation intensity was lower than that of H. persicurn. Two species of Haloxylon also maintained high transpiration rate under dark. (2) The average transpiration rate of H. persicurn was lower than H. arnrnodendron,showing that H. persicurn utilized water more efficiently than H. arnrnodendron. (3)The optimal temperature of photosynthesis of H. arnrnodendron was 24–32℃ and the optimal temperature of water utilization of H. arnrnodendron was 23. 5–25℃. (4) The water potential of H. amrnodendron was lower than –3.5 MPa whereas that of H. persicum was lower than –2.8MPa indicating that the two species of Haloscylon were under severe water stress ,and would lead to permanent wilting of their assimilating shoots.     

Effect of salinity on seed germination,ion content and photosynthesis of cotyledons in halophytes or xerophyte growing in Central Asia

Aims We investigated the impact of salinity on seed germination, chlorophyll content, chloroplast structure and photosynthesis of the green embryos in desiccated seeds of the xerophyte Haloxylon persicum, xero-halophyte Haloxylon ammodendron and euhalophyte Suaeda physophora.Methods Seeds of H. persicum, H. ammodendron and S. physophora were collected from natural environment in Fukang, Xinjiang province. Pretreatment with 700 mM NaCl was carried out to stimulate the natural 'seed priming'; we analyzed the joint effect of salinity and different species on germination physiology and cotyledonal structure and photosynthetic function changes during germination and recovery stage.Important findings We found that seeds did not suffer ion toxicity for the two halophytes H. ammodendron and S. physophora, as evidenced by the high final germination after ungerminated seeds pretreated with 700 mM NaCl were transferred to distilled water, but the final germination of the xerophyte H. persicum was significantly lower than that of control. The Na + concentration in embryos increased under salinity for all species, while K + concentration decreased by salinity only for H. persicum and H. ammodendron, i.e. the concentration of K + in embryos of H. persicum and H. ammodendron decreased by 36% and 46%, respectively. For all species, whether dry intact seeds or cotyledons of dry seeds imbibed in deionized water and NaCl solution, had high chlorophyll content. Treatment with NaCl also caused chloroplast thylakoids to swell and chlorophyll content to decrease in seeds of H. persicum, but no significant change was observed in the more salt-tolerant species S. physophora and H. ammodendron. Fluorescence measurement showed that 700 mM NaCl decreased the Fv/Fm ratio of cotyledons in seeds for all species, especially for H. persicum and H. ammodendron. Photosynthetic oxygen releasing was detected from the seeds that were moistened with distilled water and 700 mM NaCl for 6 or 24 h and from the seeds that were initially moistened with 700 mM NaCl in darkness for 10 days, then transferred to distilled water for another 6 and 24 h. The results indicated that the chlorophyll in cotyledon of desiccated seed had photosynthetic function in early germination stage, even under high-saline condition. In addition, the photosynthesis of chlorophyll in the embryonic cotyledons of desiccated seeds during germination was similar to that in leaves of young seedlings for all species. In conclusion, the chloroplasts of the two halophytes were more salt resistant compared with the xerophyte H. persicum. The photosynthetic function of chlorophyll in cotyledons of mature seeds may be ecologically important for seedling development in early stage for plants growing in extremely saline or arid environments.     

民勤荒漠区主要灌木叶片C、N、P化学计量特征的季节变化

为探究不同灌木叶片C、N、P化学计量特征季节变化规律,揭示荒漠植物对环境的适应策略,以民勤荒漠区4种主要灌木梭梭、沙拐枣、唐古特白刺、柠条锦鸡儿为研究对象,分析不同荒漠植物在生长季内叶片的C、N、P含量及其计量比变化特征。结果表明:(1)沙拐枣、柠条锦鸡儿叶片C含量显著高于唐古特白刺、梭梭(P<0.05),且唐古特白刺显著高于梭梭,沙拐枣与柠条锦鸡儿差异不显著;唐古特白刺叶片N含量显著高于其他3种植物叶片;唐古特白刺叶片P含量最高,并显著高于柠条锦鸡儿,但两者均与梭梭和沙拐枣差异不显著。(2)4种荒漠植物叶片C、N、P含量及其计量比各指标在生长季节内的变异系数表现为:P(28.34%)>C∶P(24.70%)>N∶P(19.07%)>N(17.49%)>C∶N(16.89%)>C(2.91%)。(3)C含量与N、P含量呈不显著正相关关系;除沙拐枣,其他叶片N含量与P含量呈显著正相关关系,4种荒漠植物叶片N∶P值的变化主要由P含量变化决定。(4)植物叶片C、N、C∶N、C∶P和N∶P含量的变异主要受植物种类影响,植物叶片P含量的变异主要受生长季节影响。研究发现,民勤荒漠4种灌木植物叶片C、N、P含量及C∶N、C∶P和N∶P在生长季内因物种而不同,它们在生长季内变异系数在植物种之间也存在差异。