锦鸡儿属植物系统关系及抗旱生理生态特性的研究
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摘要
锦鸡儿属(Caragana Fabr.)植物隶属于豆科(Leguminosae),蝶形花亚科(Papilionoideae)。全世界约有100余种,主要分布于亚洲和欧洲的干旱和半干旱地区。我国产62种,9个变种,12个变型。主产我国华北、东北、西北、西南各省市,并且主要集中分布在草原和荒漠区。该属植物抗旱、抗寒、耐瘠薄、繁殖性强,可防风固沙、保持水土,对我国干旱、半干旱地区生态建设具有重要的意义,同时还具有广泛的应用价值。本文利用ITS序列对锦鸡儿属植物进行了初步的分子系统学研究；选择不同抗旱性的6个代表种进行了光合特征、生理生态特征及抗氧化系统对干旱胁迫响应等的比较研究。
1.以锦鸡儿属11个系29种为代表材料,选择性扩增nrITS (?)序列并双向测序,结合黄耆亚族Astralinae (Adens) Benth其他6属7个代表种的nrITS序列进行最大简约性(MP)和最小进化(ME)的系统发育分析。结果显示：锦鸡儿属植物ITS序列长度为6l1bp-614bp；ITS-1长度为229bp-231bp；5.8S序列长度为163bp；ITS-2序列长度为216bp-220bp。与外类群排序后长度为655bp,共有170个可变位点,其中107个简约信息位点,信息位点达16.3%,因此在探讨锦鸡儿属内及近缘属间的系统学问题具有较高的参考价值,可以为属内及属间系统关系提供有力的分子证据：所研究的锦鸡儿植物被清楚地分为五个主要分支；Sect. tragacanthoides的种在MP和ME进化树中位置分散,其组的分类有待进一步商榷；卷叶锦鸡儿(C. ordosica,新种)虽然形态上与垫状锦鸡儿(C. tibetica)相似,但nrITS证据显示与它与荒漠锦鸡儿(C. roborovskyi)遗传学关系紧密；支持C. davazamcii, C. korshinskii, C. intermedia,和C. microphylla是不同的种的分类；锦鸡儿属在系统发育上不是一个单.系类群,与丽豆属(Calophaca Fish. exDC.)植物具有极为相近的亲缘关系。
2.为了更好地了解锦鸡儿属植物适应干旱环境的光合和生理生态学特性,我们选择了不同抗旱性的6个代表种：树锦鸡儿(C. arborescens,中生),小叶锦鸡儿(C. microphylla,半干旱种),以及荒漠锦鸡儿(C. roborovsyi),狭叶锦鸡儿(Cstenophylla),刺叶锦鸡儿(C. acanthophylla),和中亚锦鸡(?)(C. tragacanthoides)等4个旱生树种进行了光合日变化、叶绿素荧光日变化及抗氧化酶活性的比较研究。结果显示：C. microphylla在早晨具有最高的同化速率(A,9.93±0.05μmolm-2s-1),但正午下降了84%,而且午后仅恢复到最大值的54%。C.roborovskyi, C. stenophylla,C.acanthophylla,和C.tragacanthoides等旱生种在干旱环境中具有高同化速率(A)和较强的抗氧化能力,但是以较高的蒸腾速率(E)和较低的水分利用效率(WUE)为代价的。抵抗策略可能是旱生锦鸡儿植物适应高温、干旱等不利环境的主要机制；C.microphylla具有高光合的潜能,但在干旱、高温等不利环境下以降低气孔导度(gs)和A来增加抵抗力,节水策略可能是其适应干旱环境的主要机制。
3.为了进一步了解锦鸡儿植物的抗氧化系统对干旱环境的适应机制,我们对上述6中不同抗旱性的锦鸡儿植物在一个干旱-复水周期内抗氧化系统及超氧化岐化酶(SOD),过氧化物酶(POD),过氧化氢酶(CAT)的同工酶谱变化进行了研究。结果显示：SOD、POD、“抗坏血酸-谷胱甘肽循环”("ASA-GSH cycle")是旱生锦鸡儿植物抵御干旱,保护细胞正常的生理活动,适应干旱环境的物质基础；更多MnSOD和FeSOD同工酶的表达可能是旱生锦鸡儿植物适应干旱环境的根本原因。CAT在抵御干旱,保护细胞正常生理活动中作用有限。C.roborovskyi和C.tragacanthoides抗氧化能力较强,更适应十旱环境。脯氨酸可能在Carborescens和C.microphylla抵御干旱胁迫中起主要作用。
Caragana Fabr., belonging to the Papilionoideae (Leguminosae) and comprising about100species in the world, are mainly distributed in arid and semi-arid area of Asia and Europe. There are62species,9varieties and12geographic variants of this genus recorded in China. It mainly locates in the cold and drought areas in northwestern China. The species of Caragana play an important role in environmental protection, such as sand dune fixation and water conservation, in arid areas of China. They play important roles in ecological management and they have broad economic values. This paper presented the studies on the primary phylogenetic relationships of Caragana species and on the basis of results,6species represented different water resistance were chosen. The photosynthetic and ecophysiological traits with changes of some anti-oxidative physiological indices under a drying-rehydration cycle were studied. The main results were summarized as follows:
1. Internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA from29taxa of Caragana species and seven close relatives (all belong to Astralinae (Adens) Benth) were used for primary phylogenetic analysis. Length of the entire ITS region ranges from611to614bp in Caragana species sampled. The ITS-1region (229-231bp) was only slightly longer than ITS-2(216-220bp). No length variation was found in5.8S (163bp) region. The aligned sequences nrITS of Caragana are655bp, and170sites are variable, with107phylogenetically informative sites. The ITS sequences data are useful to resolve some relationships at lower taxonomic levels within Caragana. The species of Caragana we studied were distributed into five clades in general accordance with previous morphological and phytogeographical studies. The ITS data reveal that the species of Sect, tragacanthoides were dispersed in MP tree or ME tree. Although the morphology of C. ordosica is similar to C. tibetica, the nrITS results revealed an unexpectedly close relativeship between C. ordosica and C. roborovskyi. The ITS data also indicate C. davazamcii, C. korshinskii. C. intermedia, and C. microphylla are not the same species and Caragana Fabr. is not a monophyletic group with very close connection with Calophaca Fish. exDC.
2. To better understand the physiological characteristics in adapting to arid conditions employed by Caragana plants, the diurnal changes in photosynthetic traits and ecophysiological parameters in C. arborescens Lam.(mesophyte), C. microphylla Lam.(semiarid species), C. roborovskyi Kom., C. stenophylla Pojark., C. acanthophylla Pojark., and C. tragacanthoides Poir.(xerophytic species) were measured in Tenggeli desert. C. microphylla had the highest assimilation rate (A)(9.93±0.05μmolm-2s-1) in the morning but decreased over84%in the midday and resumed only54%in the afternoon. The four xerophytic species exhibited higher A, transpiration rate (E), stomatal conductance (gs), and lower water use efficient (WUE) than C. arborescens. In addition, the xerophytic species had more effective complex antioxidant systems. C. microphylla appears to increase drought and high temperature resistance by reducing stomata conductance and effectively osmotic adjustment, whereas the xerophytic species may use efficient antioxidative systems to adapt to the arid environment.
3. To better understand the possible mechanisms employed by the xerophytic Caragana species against drought stress, the activities of antioxidant enzymes as well as the levels of their osmotically active solutes were studied in six Caragana species grown under a drying-rehydration cycle. Our data demonstrated that the guaiacol peroxidase (POD), superoxide dismutase (SOD) with the "ascorbate-glutathione (AsA-GSH) cycle" may be the effective protector mechanism in xerophytic Caragana species, and that catalase (CAT) may have limited effects under severe stress. Different congener species likewise have differential antioxidant mechanisms to regulate their redox status. It was also determined that drought stress did not induce any new isoenzyme of SOD, POD, and CAT, but that the xerophytic Caragana species considerably have more MnSOD and FeSOD isoenzymes. On the basis of the data obtained, we could infer both C. roborovskyi and C. tragacanthoides plants have more effective complex antioxidant systems against oxidative damage. Free proline may be crucial in the resistance of C.arborescens and C. microphylla to drought stress
1.以锦鸡儿属11个系29种为代表材料,选择性扩增nrITS (?)序列并双向测序,结合黄耆亚族Astralinae (Adens) Benth其他6属7个代表种的nrITS序列进行最大简约性(MP)和最小进化(ME)的系统发育分析。结果显示：锦鸡儿属植物ITS序列长度为6l1bp-614bp；ITS-1长度为229bp-231bp；5.8S序列长度为163bp；ITS-2序列长度为216bp-220bp。与外类群排序后长度为655bp,共有170个可变位点,其中107个简约信息位点,信息位点达16.3%,因此在探讨锦鸡儿属内及近缘属间的系统学问题具有较高的参考价值,可以为属内及属间系统关系提供有力的分子证据：所研究的锦鸡儿植物被清楚地分为五个主要分支；Sect. tragacanthoides的种在MP和ME进化树中位置分散,其组的分类有待进一步商榷；卷叶锦鸡儿(C. ordosica,新种)虽然形态上与垫状锦鸡儿(C. tibetica)相似,但nrITS证据显示与它与荒漠锦鸡儿(C. roborovskyi)遗传学关系紧密；支持C. davazamcii, C. korshinskii, C. intermedia,和C. microphylla是不同的种的分类；锦鸡儿属在系统发育上不是一个单.系类群,与丽豆属(Calophaca Fish. exDC.)植物具有极为相近的亲缘关系。
2.为了更好地了解锦鸡儿属植物适应干旱环境的光合和生理生态学特性,我们选择了不同抗旱性的6个代表种：树锦鸡儿(C. arborescens,中生),小叶锦鸡儿(C. microphylla,半干旱种),以及荒漠锦鸡儿(C. roborovsyi),狭叶锦鸡儿(Cstenophylla),刺叶锦鸡儿(C. acanthophylla),和中亚锦鸡(?)(C. tragacanthoides)等4个旱生树种进行了光合日变化、叶绿素荧光日变化及抗氧化酶活性的比较研究。结果显示：C. microphylla在早晨具有最高的同化速率(A,9.93±0.05μmolm-2s-1),但正午下降了84%,而且午后仅恢复到最大值的54%。C.roborovskyi, C. stenophylla,C.acanthophylla,和C.tragacanthoides等旱生种在干旱环境中具有高同化速率(A)和较强的抗氧化能力,但是以较高的蒸腾速率(E)和较低的水分利用效率(WUE)为代价的。抵抗策略可能是旱生锦鸡儿植物适应高温、干旱等不利环境的主要机制；C.microphylla具有高光合的潜能,但在干旱、高温等不利环境下以降低气孔导度(gs)和A来增加抵抗力,节水策略可能是其适应干旱环境的主要机制。
3.为了进一步了解锦鸡儿植物的抗氧化系统对干旱环境的适应机制,我们对上述6中不同抗旱性的锦鸡儿植物在一个干旱-复水周期内抗氧化系统及超氧化岐化酶(SOD),过氧化物酶(POD),过氧化氢酶(CAT)的同工酶谱变化进行了研究。结果显示：SOD、POD、“抗坏血酸-谷胱甘肽循环”("ASA-GSH cycle")是旱生锦鸡儿植物抵御干旱,保护细胞正常的生理活动,适应干旱环境的物质基础；更多MnSOD和FeSOD同工酶的表达可能是旱生锦鸡儿植物适应干旱环境的根本原因。CAT在抵御干旱,保护细胞正常生理活动中作用有限。C.roborovskyi和C.tragacanthoides抗氧化能力较强,更适应十旱环境。脯氨酸可能在Carborescens和C.microphylla抵御干旱胁迫中起主要作用。
Caragana Fabr., belonging to the Papilionoideae (Leguminosae) and comprising about100species in the world, are mainly distributed in arid and semi-arid area of Asia and Europe. There are62species,9varieties and12geographic variants of this genus recorded in China. It mainly locates in the cold and drought areas in northwestern China. The species of Caragana play an important role in environmental protection, such as sand dune fixation and water conservation, in arid areas of China. They play important roles in ecological management and they have broad economic values. This paper presented the studies on the primary phylogenetic relationships of Caragana species and on the basis of results,6species represented different water resistance were chosen. The photosynthetic and ecophysiological traits with changes of some anti-oxidative physiological indices under a drying-rehydration cycle were studied. The main results were summarized as follows:
1. Internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA from29taxa of Caragana species and seven close relatives (all belong to Astralinae (Adens) Benth) were used for primary phylogenetic analysis. Length of the entire ITS region ranges from611to614bp in Caragana species sampled. The ITS-1region (229-231bp) was only slightly longer than ITS-2(216-220bp). No length variation was found in5.8S (163bp) region. The aligned sequences nrITS of Caragana are655bp, and170sites are variable, with107phylogenetically informative sites. The ITS sequences data are useful to resolve some relationships at lower taxonomic levels within Caragana. The species of Caragana we studied were distributed into five clades in general accordance with previous morphological and phytogeographical studies. The ITS data reveal that the species of Sect, tragacanthoides were dispersed in MP tree or ME tree. Although the morphology of C. ordosica is similar to C. tibetica, the nrITS results revealed an unexpectedly close relativeship between C. ordosica and C. roborovskyi. The ITS data also indicate C. davazamcii, C. korshinskii. C. intermedia, and C. microphylla are not the same species and Caragana Fabr. is not a monophyletic group with very close connection with Calophaca Fish. exDC.
2. To better understand the physiological characteristics in adapting to arid conditions employed by Caragana plants, the diurnal changes in photosynthetic traits and ecophysiological parameters in C. arborescens Lam.(mesophyte), C. microphylla Lam.(semiarid species), C. roborovskyi Kom., C. stenophylla Pojark., C. acanthophylla Pojark., and C. tragacanthoides Poir.(xerophytic species) were measured in Tenggeli desert. C. microphylla had the highest assimilation rate (A)(9.93±0.05μmolm-2s-1) in the morning but decreased over84%in the midday and resumed only54%in the afternoon. The four xerophytic species exhibited higher A, transpiration rate (E), stomatal conductance (gs), and lower water use efficient (WUE) than C. arborescens. In addition, the xerophytic species had more effective complex antioxidant systems. C. microphylla appears to increase drought and high temperature resistance by reducing stomata conductance and effectively osmotic adjustment, whereas the xerophytic species may use efficient antioxidative systems to adapt to the arid environment.
3. To better understand the possible mechanisms employed by the xerophytic Caragana species against drought stress, the activities of antioxidant enzymes as well as the levels of their osmotically active solutes were studied in six Caragana species grown under a drying-rehydration cycle. Our data demonstrated that the guaiacol peroxidase (POD), superoxide dismutase (SOD) with the "ascorbate-glutathione (AsA-GSH) cycle" may be the effective protector mechanism in xerophytic Caragana species, and that catalase (CAT) may have limited effects under severe stress. Different congener species likewise have differential antioxidant mechanisms to regulate their redox status. It was also determined that drought stress did not induce any new isoenzyme of SOD, POD, and CAT, but that the xerophytic Caragana species considerably have more MnSOD and FeSOD isoenzymes. On the basis of the data obtained, we could infer both C. roborovskyi and C. tragacanthoides plants have more effective complex antioxidant systems against oxidative damage. Free proline may be crucial in the resistance of C.arborescens and C. microphylla to drought stress
引文
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