979 resultados para root : shoot ratio
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Andean montane forests are one of the most diverse ecosystems on Earth, but are also highly vulnerable to climate change. Therefore, the link between plant distribution and ecosystem productivity is a critical point to investigate in these ecosystems. Are the patterns in productivity observed in montane forest due to species turnover along the elevational gradients? Methodological constraints keep this question unanswered. Also, despite their importance, belowground biomass remains poorly quantified and understood. I measured two plant functional traits in seedlings, root:shoot ratio and specific leaf area, to identify different strategies in growth and biomass allocation across elevations. A tradeoff in specific leaf area with elevation was found in only one species, and no generalized directional change was detected with elevations for root:shoot ratio. Lack of information for the ontogeny of the measured plant traits could confounding the analysis.
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Genetic modification of shoot and root morphology has potential to improve water and nutrient 19 uptake of wheat crops in rainfed environments. Near-isogenic lines (NILs) varying for a tillering 20 inhibition (tin) gene and representing multiple genetic backgrounds were investigated in contrasting 21 controlled environments for shoot and root growth. Leaf area, shoot and root biomass were similar 22 until tillering whereupon reduced tillering in tin-containing NILs produced reductions of up to 60% in 23 total leaf area and biomass, and increases in total root length of up to 120% and root biomass to 24 145%. Together, root-to-shoot ratio increased two-fold with the tin gene. The influence of tin on shoot 25 and root growth was greatest in the cv. Banks genetic background, particularly in the biculm-selected 26 NIL, and was typically strongest in cooler environments. A separate de-tillering study confirmed 27 greater root-to-shoot ratios with regular tiller removal in non-tin containing genotypes. In validating 28 these observations in a rainfed field study, the tin allele had a negligible effect on seedling growth but 29 was associated with significantly (P<0.05) reduced tiller number (-37%), leaf area index (-26%) and 30 spike number (-35%) to reduce plant biomass (-19%) at anthesis. Root biomass, root-to-shoot ratio at 31 early stem elongation and root depth at maturity were increased in tin-containing NILs. Soil water use 32 was slowed in tin-containing NILs resulting in greater water availability, greater stomatal 33 conductance, cooler canopy temperatures and maintenance of green leaf area during grain-filling. 34 Together these effects contributed to increases in harvest index and grain yield. In both the controlled 35 and field environments, the tin gene was commonly associated with increased root length and biomass 36 but the significant influence of genetic background and environment suggests careful assessment of 37 tin-containing progeny in selection for genotypic increases in root growth.
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Coffee (Coffea arabica L.) plants were grown in small (3-L), medium (10-L) and large (24-L) pots for 115 or 165 d after transplanting (DAT), which allowed different degrees of root restriction. Effects of altered source : sink ratio were evaluated in order to explore possible stomatal and non-stomatal mechanisms of photosynthetic down-regulation. Increasing root restriction brought about large and general reductions in plant growth associated with a rising root : shoot ratio. Treatments did not affect leaf water potential or leaf nutrient status, with the exception of N content, which dropped significantly with increasing root restriction even though an adequate N supply was available. Photosynthesis was severely reduced when plants were grown in small pots; this was largely associated with non-stomatal factors, such as decreased Rubisco activity. At 165DAT contents of hexose, sucrose, and amino acids decreased in plants grown in smaller pots, while those of starch and hexose-P increased in plants grown in smaller pots. Photosynthetic rates were negatively correlated with the ratio of hexose to free amino acids, but not with hexose content. Activities of acid invertase, sucrose synthase, sucrose-P synthase, fructose-1,6- bisphosphatase, ADP-glucose pyrophosphorylase, starch phosphorylase, glyceraldehyde-3-P dehydrogenase, PPi : fructose-6-P 1-phosphotransferase and NADP : glyceraldehyde-3-P dehydrogenase all decreased with severe root restriction. Glycerate-3-P : Pi and glucose-6-P : fructose-6-P ratios decreased accordingly. Photosynthetic down-regulation was unlikely to have been associated directly with an end-product limitation, but rather with decreases in Rubisco. Such a down-regulation was largely a result of N deficiency caused by growing coffee plants in small pots.
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Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)
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This experiment was conducted in green house conditions to evaluate the DM accumulation in the shoots and in the roots of two cultivars of Lablab purpureus (L.) Sweet. A 2x3 factorial (two cultivars and three evaluation dates) was conducted according to a randomized complete block design with four replications, being the cultivars Highworth and Rongai evaluated at 42, 56, and 70 days after seedling emergence (DASE). The results indicated that the cvs. Highworth and Rongai have the same pattern of DM accumulation in the shoots. In the upper layer of the soil (0-0.20 in) it was found 38.83% and 43.64% of the DM accumulated in the roots down to 2.00 in depth, in the cvs. Highworth and Rongai, respectively. In the deepest layer (1.80-2.00 in) it was found 3.02% and 1.5% of the DM accumulated in the roots of the cvs. Highworth and Rongai, respectively. The root density showed a striking decrease upper layer from the soil (0-0.2 m) down to the depth of 0.60 0.80 in (from 10.83 to 1.75 cm.cm(-3) in the cv. Highworth and from 10.76 to 1.28 cm.cm(-3) in the cv. Rongai). At the bottom layer (1.80-2.00 in) the root density values were 0.98 cm.cm(-3) and 0.59 cm.cm(-3), respectively for the cvs. Highworth and Rongai. The root/shoot ratios were similar in both cvs. and decreased from 42 to 70 DASE showing that the cvs. evaluated had the same dynamics of DM accumulation.
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碳循环研究已经成为目前生态学中的研究热点之一。草地生态系统在全球碳循环中占有重要地位,而且大部分草地处于对全球变化敏感的区域,也是研究生态系统对全球变化响应的关键地区。我国草地面积十分辽阔,占我国陆地面积的40%以上,具有极其重要的生态地位和作用。 在碳蓄积研究中,对地上碳库的估计比较容易,而对地下碳库的估计却由于方法上的局限造成估计的精度不高。因此,研究陆地生态系统中植物群落的地上/地下生物量分配模式,尤其是地下生物量的分配,对于全球碳蓄积的 研究具有重要意义。植物生长与环境因子密切相关,植物的生物量分配与环境因子也必然存在一定的相关关系,了 解植物的生物量分配与环境因子之间的关系,并建立相关模式,不但有利于减少生物量调查的工作量,而且能使碳蓄积量估计的精度大大提高。 (1) 本文通过对锡林郭勒草原上的克氏针茅植物群落进行了为期两年的地上和地下生物量观测,初步得到了克氏针茅群落地上/地下生物量分配规律: 2004年地上生物量表现为逐渐增加的趋势,6、7、8月地上生物量较低,9月显著增加,为8月的4.8倍。2005年地上生物量表现出明显的单峰曲线,8月达到峰值,为150.48 g•m-2。2004年地下生物量6月到7月明显增长,7月、8、9月逐渐减少。2005年地下生物量出现两次增加和两次减小的过程。地下生物量在不同层次的分配数量及所占比例存在差异。0~10cm分配的地下生物量最多,所占比例最大,往下各层基本表现出逐层递减的趋势。其中,0~30cm地下生物量占0~80cm总地下生物量的70%。 地下生物量与地上生物量的比值即为根冠比,根冠比也具有明显的季节变化。2004年从6月到9月根冠比表现为逐渐减小的趋势,最小根冠比值出现在9月,为9.22。2005年为明显的反抛物线形状,最小值出现在8月,为19.93。 (2) 分析克氏针茅群落地上生物量、地下生物量、根冠比三者关系的结果表明:克氏针茅草原群落地上生物量和根冠比的相关关系较为明显,而地下生物量与根冠比的相关关系较差。地上生物量与地下生物量的相关关系较好,呈正相关关系。地上生物量对根冠比变化的贡献较大,而地下生物量对根冠比变化的贡献较小。 (3)在观测生物量的同时,用小气候自动观测系统记录了同时期的降水和气温资料,通过分析克氏针茅群落地上、地下生物量及根冠比与降水、气温的关系,得出以下结论: 2004年和2005年地上生物量均与上月降水量有显著相关关系,而两年的地下生物量与降水量的相关关系较差。两年的根冠比变化与上月降水量也具有较好的相关关系。2004年地下生物量与上月平均气温的相关关系较好。2005年地下生物量与月平均气温的相关关系较差。2004和2005年地上生物量与月平均气温有很好的线性关系,但两年的相关关系正好相反。两年的根冠比与月平均气温的相关关系也表现的恰恰相反。 (4) 最后,分析了温室控制试验羊草种群的生物量分配对水热因子响应的阈值。得出以下结论: 土壤相对含水量范围为6065%、温度为26℃时的地上、地下干重都最大,表明这是羊草地上、地上部分生长最适宜的温度和土壤水分。土壤为相对含水量2530%、温度为20℃时根冠比值最大。 温度、土壤水分以及二者的交互作用对羊草根冠比的影响均达到极显著水平。但根冠比对不同温度和土壤水分的敏感度不同。根冠比在一定的温度范围和土壤相对含水量范围内发生敏感变化。
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针对人工栽培基质中基质配比问题,采用生物试验的方法,对陕西彬县的泥炭、陕西铜川的油页岩、陕西关中土粘化层土壤,进行不同比例的配合后,再对配合基质设制不同处理,进行作物的室内栽培试验,通过生物量、生物性状(根冠比)、及产量分析来判断基质配比的优劣。研究结果表明:随着泥炭比例的增加,小青菜的产量增加,泥炭量达到一定值后,随着泥炭量的增加,小青菜的产量反而降低;泥炭与<1 mm粘土配合(PS)处理中,PS3处理(泥炭∶粘土为3∶1)小青菜地上部分产量最大;在泥炭与7~10 mm粘土配合(PB)处理中,PB2处理(泥炭∶粘土为2∶1)小青菜地上部分产量最大;对小青菜根系生物量来说,在泥炭和粘土比例较小时,小颗粒粘土处理的根量明显高于大颗粒粘土处理,在泥炭和粘土的比例较大时,大颗粒处理高于小颗粒处理。对油页岩处理而言,在泥炭量较少的配合基质中加入8%的油页岩能起到良好的增产作用;而在泥炭含量较多时,对小青菜产量的影响是负面的;石灰处理对小青菜地上部分产量影响都是负面的,但是对小青菜根系生物量的影响基本是正面的。高温处理对小青菜地上部分产量的影响基本上都是负面的,对小青菜根系的生长也基本是起负作用的。根冠比达到某一适宜值...
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人类向大气中排放的大量氮氧化合物和氟氯烃类化合物(CFC’s)引起臭氧分子的分解,导致到达地球表面的紫外辐射增加,特别是UV-B辐射增强。本项目以青杨组杨树为模式植物,从形态和生理方面研究了来自不同UV-B背景下的康定杨与青杨在增强UV-B下的反应及其反应差异,并探讨了干旱、施肥对它们抗UV-B能力的影响。杨树具有分布广、适应性强、在生态环境治理和解决木材短缺方面均占有重要位置,研究成果可为生态系统的恢复与重建提供理论依据和科学指导。主要研究结果有以下: 1. 在温室中经过增强UV-B处理,杨树的外部形态及生理活动受到了一定程度的影响。增强UV-B导致康定杨、青杨的生物量、叶面积及节间长度降低,叶片增厚,SOD活性升高,膜伤害增加,而对叶片数目、R/S、叶绿素A、叶绿素B及整个叶绿素含量没有影响。两种杨树对UV-B胁迫的响应存在差异:在增强UV-B条件下,青杨的植株高度、生物量、叶面积、脯氨酸含量、长期用水效率受到的影响大于康定杨,相比而言,康定杨在比叶面积、叶片厚度、可溶性糖含量、UV-B吸收物质的含量及SOD和GPX活性方面增加的程度大于青杨。这些区别说明,来自于高海拔的康定杨比来自于低海拔的青杨对增强UV-B 具有更强的耐性。我们认为二者在叶片厚度、比叶面积、UV-B吸收物质含量及SOD、GPX活性差异是导致对增强UV-B耐性不同的原因。 2. 干旱与增强UV-B对杨树的生长和生理特性均产生了影响,而且两种胁迫共同作用时干旱表现减弱或加剧了UV-B对杨树某些形态和生理特性的影响。 据试验结果,干旱显著地降低了杨树的株高、叶片数目、叶面积,增加了叶片厚度,促进ABA的积累,提高了CAT活性。对于干旱,两种杨树之间也表现出了一定的差异性。可溶性蛋白质和脯氨酸在青杨叶片中得到显著积累,而在康定杨中没有变化。此外,CAT、长期用水效率在康定杨中受到的影响更加明显。长期用水效率的不同变化趋势说明两种杨树对水分胁迫采用了不同的用水策略,康定杨采用的是节水用水策略,提高用水效率,而青杨采用的是耗水的用水策略。根据干旱对叶面积、脯氨酸、ABA含量、CAT活性及长期用水效率等方面的影响,我们认为来自高海拔地区的康定杨比来自低海拔的青杨有更大的耐旱性,这是对生长环境长期适应的结果。在高海拔地区,因霜冻常带来土壤水分不可利用,降低了根系对水分的吸收,树木容易受到的生理性干旱。另外,高海拔的地区低的气温使植株对严寒有较强的耐性,减少了水分的需要。 生长于增强UV-B下的康定杨和青杨植株表现为高度降低,叶面积缩小,比叶面积增加;叶片栅栏组织、海绵组织均受到增强UV-B的影响,其厚度的增加导致整个叶片变厚。增强UV-B还显著提高了杨树的APX活性、UV-B吸收物质含量,而对叶片数目、ABA、可溶性蛋白质含量及CAT活性没有产生影响。试验中也观察到了两种杨树对增强UV-B响应的差异:与康定杨相比,在增强UV-B下青杨株高、叶面积降低的程度更大一些,SOD活性显著提高。另外UV-B吸收物质受到的影响不同。根据这些差别,高海拔的康定杨(3500 m)比来自低海拔的青杨(1500 m)增强UV-B有较强的耐性。 与水分充足情况下UV-B对植株的影响相比,干旱对杨树抗增强UV-B产生了一定的影响,表现为加剧或减弱UV-B对植物的影响,但这种影响与形态、生理指标有关。当干旱与增强UV-B共同作用时,杨树植株的株高、叶面积进一步降低、叶片进一步增厚。就脯氨酸的积累的而言,在没有水分胁迫时,增强UV-B促使它显著增加,而在干旱处理下这种效果变得不明显。干旱对增强UV-B的影响还与杨树的种类有一定的关系。在康定杨中,干旱减弱了增强UV-B对栅栏组织与海绵组织的影响,且在植株高度、叶面积上表现出累加效应,而在CAT上交互作用显著;但在青杨中干旱则加剧增强UV-B对栅栏组织与海绵组织的影响,在植株高度、叶面积及比叶面积上表现出显著的交互作用。据碳同素分析,在水分充足的条件下,无论是康定杨,还是青杨,增强UV-B均导致其长期用水效率的提高,然而当两种胁迫共同作用时,长期用水效率则表现出差异,在青杨中,长期用水效率得到进一步增高,而康定杨中干旱的效应被增强UV-B所减轻。 3. 田间试验表明,杨树的生长、生理特征都受到养分和增强UV-B的影响。施肥对杨树的影响表现为:提高了叶面积、生物量及SOD的活性,降低了抗坏血酸含量。对于施肥作用,两种杨树的反应也有区别:在康定杨中施肥显著增加了的叶片长度、宽度及光合色素的含量,降低了净光合速率、气孔导度及胞间CO2浓度;在青杨中,则SOD、GPX、APX活性表现增加。从试验看出,施肥对来自于高海拔地区的康定杨(3500 m)的影响较大,对来自低海拔的青杨(1500 m)影响较小,这与它们对原产地的生境适应有一定关系。在康定杨生长的高海拔地区,低温度和湿度不能为地上凋落物或土壤中的根分解提供理想的条件,造成当地土壤的低养分状况,所以当肥料施用以后,效果显著。 经过增强UV-B处理,杨树叶片中UV-B吸收物质含量、GPX的活性得到提高,而脯氨酸、丙二醛、可溶性蛋白质、叶绿素及类胡萝卜素含量没有受到影响。对于增强UV-B两种杨树受到的影响也有所不同:在青杨中增强UV-B导致叶面积缩小,生物量、净光合速率降低,APX的活性及长期用水效率的提高,而对康定杨的这些指标没有产生显著影响,相反抗氧化酶的活性明显高于青杨。这些差异性是由于两种杨树对原产地不同UV-B背景的长期适应结果。康定杨长期生长在较高UV-B环境中,对UV-B有较强的耐性。而青杨适应于较低的UV-B环境,对增强UV-B较为敏感。 试验中施肥也影响了植株对增强UV-B的反应,不过这种影响与杨树的种类及测定指标有一定的相关性。例如,在缺肥的情况下,青杨的长期用水效率和康定杨的叶绿素含量都受到增强UV-B的显著影响,而施肥以后这种影响变得不显著。在缺肥的条件下,GPX、APX在青杨中的活性、GPX在康定杨中的活性对增加UV-B反应不敏感;而施肥以后则变化显著,同样胞间CO2浓度在康定杨也有类似的变化。 For past decades, Ultraviolet radiation, especially UV-B reaching the Earth’s surface increased because of depletion of ozone layer resulted from emission of NxO and CFC’s from human activities. In this experiment, different species of Populus section Tacamahaca Spach from different UV-B background were selected as a model plant to assess the effects of enhanced UV-B radiation. Morphological and physiological traits induced by enhanced UV-B were observed and the different responses between P. kangdingensis and P. cathayana were discussed, furthermore the influences of drought and fertilizer on responses induced by enhanced UV-B were studied. Since poplars play an important role in lumber supply, and are important component of ecosystems due to their fast growth and wide adaptation, the study could provide a strong theoretical evidence and scientific direction for the afforestation, and rehabilitation of ecosystem. The results are as follows: 1. The experiment conducted in a greenhouse indicated that morphological and physiological traits of two poplars were affected by enhanced UV-B radiation. Enhanced UV-B radiation not only reduced biomass, leave area and internode length, but also increased leaf thickness and SOD activity as well as MDA concentration and electrolyte rate. However, no significant changes in leaf numbers, root shoot ratio, and total chlorophyll and chlorophyll component were observed. There were different responses to enhanced UV-B radiation between two species. Compared with P. kangdingensis, cuttings of P. cathayana, exhibited lower height increment and smaller leaf area. In addition, there were significant differences in free proline, soluble protein, and UV-B absorbing compounds, and the activity of SOD and GPX, long-term WUE between them. Differences in plant height, biomass, leaf area, free proline concentration, and long-termed WUE showed that P. cathayana were more affected by enhanced UV-B radiation than P. kangdingensis. In contrast, more increase of specific leaf mass, leaf thickness, and soluble sugar, and UV-B absorbing compounds, and activity of SOD and GPX were observed in P. kangdingensis. According to these results, we suggested that P. kangdingensis from high elevation, which adapted to higher UV-B environments, had more tolerance to enhanced UV-B than P. cathayana from low elevation, which adapted to lower UV-B environment. We believe it was the difference of leaf thickness, specific leaf mass, and UV-B absorbing compounds as well as the activity of SOD and GPX resulted in lower adaptation of P. cathayana to enhanced UV-B radiation. 2. Growth and physiological traits of two poplars were affected by both drought and enhanced UV-B radiation. Moreover, it was observed that when two stresses applied together drought could exacerbate UV-B effects or decrease sensitivity to UV-B. In the experiment, drought significantly decreased plant height, leaf numbers, leaf area, and increased leaf thickness, and ABA, and CAT activity of two poplars. There were significant interspecific differences to drought stress. Exposed to drought, soluble protein and proline concentration were increased in P. cathayana but not in P. kangdingensis. However, more changes in CAT and long-term WUE were observed in kangdingensis. Different change in long-term WUE suggests that two poplars adapted different water-use strategies. P. kangdingensis employ a conservative water-use strategy, whereas P. cathayana employ a prodigal water-use strategy. Based on the differences in leaf area, accumulation of free proline and ABA, CAT activity as well as long-term WUE, we believed that P. kangdingensis from high elevation had a greater tolerance to drought than P. cathayana from low elevation,which is the result of adaptation to local environment. In high elevation area, trees are prone to suffer from physiological drought because of un-movable water caused by frost. Besides lower temperature enable the plants had greater adaptability to frost as a results the requirement of water is reduced Enhanced UV-B radiation decreased shoots height, leaf area, and increased specific leaf mass and thickness of palisade and sponge layer as well as APX activity and UV-B absorbing compounds in both species. Whereas, leaf numbers, ABA content, soluble protein and CAT activity showed no differences to enhanced UV-B radiation. Interspecific differences were also observed. Compared with P. kangdingensis, P. cathayana showed lower shoot height and smaller leaf area, higher SOD activity. Besides, variation in UV-B absorbing compounds was found. These differences suggested that P. kangdingensis from high elevation (3500 m) was more tolerant to enhanced UV-B radiation than P. cathayana from low elevation (1500 m). Compared with morphological and physiological changes induced by enhanced UV-B radiation under well-watered conditions, drought exacerbated or decreased these changes. However, these effects vary with parameters measured. When two stresses applied together, shoot height and leaf area further decreased while leaf thickness further increased. Under well-watered conditions, enhanced UV-B radiation significantly increased proline content, but such effect was not observed under drought conditions. The effect of drought on enhanced UV-B radiation was related to species. For example, drought reduced the effects of enhanced UV-B radiation on palisade parenchyma and sponge mesophyll in P. kangdingensis, and additive effects in shoot height and leaf area and interactive effect CAT activity were observed. In contrast, for P. cathayana drought significantly exacerbated the effects of enhanced UV-B radiation on palisade parenchyma and sponge mesophyll; there were noticeable interaction in shoot height, leaf area and specific leaf mass. As far as long-term WUE is concerned, it was increased by enhanced UV-B radiation under well-watered conditions in both species. While different effect was observed between two species in combination of two stresses. Long-term water use efficiency was further increased in P. cathayana whereas the effect was less significant in P. kangdingensis. 3. The field experiment showed that growth and physiological traits of poplars were affected by nutrition and enhanced UV-B radiation. Fertilization significantly increased leaf area, biomass and SOD activity, reduced Ascorbic acid concentration. There was interspecific difference in response to fertilization. For P. kangdingensis, fertilization significantly increased leaf width, leaf length and photosynthetic pigments content while net photosynthetic rate and stomatal conductance, intercellular CO2 concentration were significantly decreased. However, for P. cathayana, these parameters were unaffected except the increase of SOD, GPX and APX activity. From above, it could concluded that P. kangdingensis from high elevation was more affected by fertilization than P. cathayana, This difference was due to adaptation to local environment., The low temperature and moisture where P. kangdingensis was collected can not provided optimum to decompose roots and litter fall as a result the nutrition in soil was poor. Exposed to enhanced UV-B radiation, for both species UV-B absorbing compounds and GPX activity were significantly increased while proline, MDA, soluble protein, chlorophyll, carotenoids were not affected. Different responses were also observed between the two species. Enhanced UV-B radiation caused significant decreases in leaf area, biomass, net photosynthetic rate and increase in APX activity and long-term WUE in P. cathayana but not in P. kangdingensis. In addition, activity in antioxidant enzymes was much higher in P. kangdingensis than in P. cathayana. In the experiment fertilization affected responses of cuttings to enhanced UV-B radiation, but it concern species and parameters measured. Long-term WUE in P. cathayana and chlorophyll in P. kangdingensis were significantly increased by enhanced UV-B radiation under non-fertilization treatments while the increase was not found under fertilization treatment. In contrast, under no fertilization treatment enhanced UV-B radiation did not affected GPX and APX activity in P. cathayana and GPX in P. kangdingensis while significant increase appeared after application of fertilization. Similar effect of enhanced UV-B radiation on intercellular CO2 concentration in P. kangdingensis was observed.
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本研究通过粗枝云杉不同种群进行的温室半控制试验,采用植物生态学、生理学和生物化学的研究方法,系统地研究了粗枝云杉不同种群抗旱性的生长、形态、生理和生化机理,并结合有关研究进行综合分析,得出主要研究结论如下: 1.粗枝云杉对干旱胁迫的综合反应 粗枝云杉在干旱胁迫下的适应机制为:(1)相对生长速率及植株结构的调整:干旱胁迫下虽然植株相对生长速率显著降低,且有相对较多的生物量向根部分配,但并未发现细根/总根比增加。(2)粗枝云杉对干旱胁迫的光合作用表现为:干旱胁迫显著地降低了控制的理想条件下的气体交换,但干旱胁迫对PSII最大光化学效率(Fv/Fm)没有影响,表明干旱并未影响到光合机构。(3)干旱还影响了很多生理生化过程,包括渗透调解物质(游离脯氨酸)、膜脂过氧化产物、脱落酸(ABA)含量的增加,以及保护酶活性的升高。这些结果证明植物遭受干旱胁迫后发生了一系列的形态、生理和生化响应,这些变化能提高干旱时期植物的存活和生长能力。 2.粗枝云杉不同种群对干旱胁迫反应的种群差异 粗枝云杉三个种群-干旱种群(四川丹巴和甘肃迭部)和湿润种群(四川黑水)对干旱适应不同,这种不同应归因于它们采用的用水策略不同:在水分良好和干旱胁迫条件下,受试种群在相对生长速率和水分利用效率(WUE)方面都表现出显著的种群间差异。与湿润种群相比,干旱种群在两种水分条件下有更高的WUE。粗枝云杉不同种群的碳同位素组分(δ13C)只在干旱胁迫下有显著差异,并且这种差异在水分良好时比干旱胁迫条件下小,说明生理响应和干旱适应性之间的关系受植物内部抗旱机制和外部环境条件(如水分可利用性)或两者互作效应的影响。这些结果说明干旱种群和湿润种群所采用的用水策略不同。干旱种群有更强的抗旱能力,采用的是节水型的用水策略,而湿润种群抗旱能力较弱,采用的是耗水型的用水策略。 3. 遮荫对粗枝云杉不同种群抗旱性影响 干旱胁迫显著降低了全光条件下叶相对含水量(RWC)、相对生长速率、气体交换参数、PSII的有效量子产量(Y),提高了非光化学猝灭效率(qN)、水分利用效率、脯氨酸(PRO)积累、脱落酸(ABA)含量及保护酶活性。然而这种变化在遮荫条件下不明显。我们得出结论适度遮荫降低了干旱对植物的胁迫作用。另一方面,在干旱条件下,与湿润种群相比,干旱种群抗旱性更强,表现在干旱种群净光合速率与单位重量上叶氮含量(Nmass)降低较少。另外,干旱种群表现出更为敏感的气孔导度,更高的热耗散能力(qN)能力、用水效率、ABA积累、保护酶活性,以及更低的总用水量、相对生长速率。这一结果表明这两种群采用不同的生理策略对干旱和遮荫做出反应。许多生长和生理反应差异与这两个种群原产地气候条件相适应。 4. 外源脱落酸(ABA)喷施对粗枝云杉不同种群抗旱性影响 外源ABA喷施在干旱和水分良好条件下均不同程度地提高了根/茎比,表明根和茎对ABA敏感程度不同。实验结果还表明,外源ABA喷施对这两个种群在干旱胁迫期间影响不同。干旱胁迫期间,伴随着ABA喷施,湿润种群净光合速率(A)显著降低,而干旱种群净光合速率变化不明显。另一方面,外源ABA喷施显著提高了干旱条件下干旱种群的单位叶面积重(LMA)、根/茎比、细根/总根(Ft)比、水分利用效率(WUE)、ABA含量, 以及保护酶活性。然而,外源ABA喷施对湿润种群的上述测定指标没有显著影响。这一结果表明干旱种群对外源ABA喷施更为敏感, 反应在更大的气孔导度降低,更高的生物量可塑性,及更高的水分利用效率、ABA含量和保护酶活性。综上所述,我们得出结论,粗枝云杉对外源ABA敏感性因种群的不同而不同。该研究结果可为两个明显不同种群在适应分化方面提供强有力的证据。 Arid or semi-arid land covers more than half of China's land territory. In arid systems, severe shortages of soil water often coincide with periods of high temperatures and high solar radiation, producing multiple stresses on plant performance. Protection from high radiation loads in shaded microenvironments during drought may compensate for a loss of productivity due to reduced irradiance when water is available. Additionally, ABA, a well-known stress-inducible plant hormone, has long been studied as a potential mediator for induction of drought tolerance in plants. Picea asperata Mast., which is one of the most important tree species used for the production of pulp wood and timber, is a prime reforestation species in western China. In this experiment, different population of P. asperata were used as experiment material to study the adaptability to drought stress and population differences in adaptabiliy, and the effects of shade and exogenous abscisic acid (ABA) application on the drought tolerance. Our results cold provide a strong theoretical evidence and scientific direction for the afforestation, and rehabilitation of ecosystem in the arid and semi-arid area, and provide a strong evidence for adaptive differentiation of different populations, and so may be used as criteria for species selection and tree improvement. The results are as follows: 1. A large set of parallel response to drought stress Drought stress caused pronounced inhibition of the growth and increased relatively dry matter allocation into the root; drought stress also caused pronounced inhibition of photosynthesis, while drought showed no effects on the maximal quantum yield of PSII photochemistry (Fv/Fm) in dark-adapted leaves, indicating that drought had no effects on the primary photochemistry of PSII. However, in light-adapted leaves, drought reduced the quantum yield of PSII electron transport (Y) and increased the non-photochemical quenching (qN). Drought also affected many physiological and biochemical processes, including increases in superoxide dismutase (SOD), ascorbate peroxidase (APX) activities, malondialdehyde and ABA content. These results demonstrate that there are a large set of parallel changes in the morphological, physiological and biochemical responses when plants are exposed to drought stress; these changes may enhance the capability of plants to survive and grow during drought periods. 2. Difference in adaptation to drought stress between contrasting populations of Picea asperata There were significant population differences in growth, dry matter allocation and water use efficiency. Compared with the wet climate population (Heishui), the dry climate population (Dan ba and Jiebu) showed higher LMA, fine root/total root ratio and water use efficiency under drought-stressed treatments. The results suggested that there were different water-use strategies between the dry population and the wet population. The dry climate population with higher drought tolerance may employ a conservative water-use strategy, whereas the wet climate population with lower drought tolerance may employ a prodigal water-use strategy. These variations in drought responses may be used as criteria for species selection and tree improvement. 3. The effects of shade on the drought tolerance For both populations tested, drought resulted in lower needle relative water content (RWC), relative growth rate (RGR), gas exchange parameters and effective PSII quantum yield (Y), and higher non-photochemical quenching (qN), water use efficiency (WUE), proline (PRO) and abscisic acid (ABA) accumulation, superoxide dismutase (SOD), ascorbate peroxidase (APX) activities as well as malondialdehyde (MDA) levels and electrolyte leakage in sun plants, whereas these changes were not significant in shade plants. Our study results implied that shade, applied together with drought, ameliorated the detrimental effects of drought. On the other hand, compared with the wet climate population, the dry climate population was more tolerant to drought in the sun treatment, as indicated by less decreases in A and mass-based leaf nitrogen content (Nmass), more responsive stomata, greater capacity for non-radiative dissipation of excitation energy as heat (analysed by qN), and higher WUE,higher level of antioxidant enzyme activities,higher ABA accumulation as well as lower MDA content and electrolyte leakage. Many of the differences in growth and physiological responses reported here are consistent with the climatic differences between the locations of the populations of P. asperata. 4. The effects of exogenous abscisic acid (ABA) application on the drought tolerance For both populations tested, exogenous ABA application increased root/shoot ratio (Rs) under well-watered and drought-stressed conditions, indicating that there was differential sensitivity to ABA in the roots and shoots. However, it appeared that ABA application affected the two P. asperata populations very differently during drought. CO2 assimilation rate (A) was significantly decreased in the wet climate population, but only to a minor extent in the dry climate population following ABA application during soil drying. On the other hand, ABA application significantly decreased stomatal conductance (gs), transpiration rate (E) and malondialdehyde (MDA) content, and significantly increased leaf mass per area (LMA), Rs, fine root/total root ratio (Ft), water use efficiency (WUE), ABA contents, superoxide dismutase (SOD), ascorbate peroxidase (APX) and catalase (CAT) activities under drought condition in the dry climate population, whereas ABA application did not significantly affect these parameters in the wet population plants. The results clearly demonstrated that the dry climate population was more responsive to ABA application than the wet climate population, as indicated by the strong stomata closure and by greater plasticity of LMA and biomass allocation, as well as by higher WUE, ABA content and anti-oxidative capacity to defense against oxidative stress, possibly predominantly by APX. We concluded that sensitivity to exogenous ABA application is population dependent in P. asperata. Our results provide strong evidence for adaptive differentiation between populations of P. asperata.
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沙棘广泛分布于亚欧大陆的温带地区和亚洲亚热带的高海拔地区。沙棘能适应多种生态环境,能耐受多种逆境(如干旱、低温、高温和盐害等)。在中国,沙棘常常被用作植被恢复中的先锋树种而大量栽培。本文以中国沙棘为试验材料,探索沙棘适应干旱机制,以及沙棘对干旱胁迫的适应机制是否存在种群间的差异,同时试图通过分析干旱胁迫下沙棘叶片蛋白质表达变化探索沙棘适应干旱胁迫的分子机理。 对三个分别来自低海拔湿润地区、低海拔干旱地区和高海拔湿润地区的中国沙棘种群进行干旱胁迫处理。干旱胁迫能提高根冠比,比叶面积,降低平均叶面积和总生物量,提高沙棘的抗氧化性酶活性、脯氨酸含量、脱落酸(ABA)含量、降低光合作用,提高长期用水效率。实验中的这两个低海拔种群比高海拔种群抵抗干旱的能力更强,不同的种群采用了不同的策略来耐受干旱胁迫和过氧化胁迫。 在2004 年度的实验中,干旱胁迫处理下,高海拔湿润种群(道孚种群)严重失水,生长也受到更大的抑制,非气孔因素在抑制光合作用方面占支配地位,抗坏血酸含量下降,ABA和脯氨酸含量增加幅度比九寨沟种群的要高,这可能是因为道孚种群严重失水而引起的;而低海拔湿润种群(九寨沟种群)的体内水分状况几乎不受干旱的影响,生长情况也较道孚种群要好。 在2005 年度的试验中,和高海拔湿润地区种群(道孚)相比较,低海拔干旱地区种群(定西)在叶片相对水含量、根冠比、抗氧化酶活性(过氧化氢酶、抗坏血酸过氧化物酶和谷胱甘肽过氧化物酶)、保护性物质(脯氨酸,脱落酸)含量等方面都要高,光能热耗散能力也更强,而且气体交换参数(气孔扩散阻力和胞间CO2浓度等)对干旱也更不敏感。 分析了干旱胁迫下沙棘叶片蛋白质表达的变化。共发现319 个蛋白质,有4 个蛋白在干旱胁迫下消失(Putative ABCtransporter ATP-binding protein 、Hypothetical proteinXP-515578,热激蛋白Hslu219 和一个没得到鉴定的蛋白),4 个只在干旱胁迫下出现(没命名的蛋白质产物,对甲基苯-丙酮酸双加氧酶,NTrX 和一个没得到鉴定的蛋白),46 个蛋白质的表达丰度变化显著,包括32 个干旱负调蛋白,14 个干旱正调蛋白(3 个Rubisco 的大亚基、J-type–co-chaperone Hsc20、putative protein DSM3645-2335、putative acyl-COA 脱氢酶、nesprin-2 和两个没有得到鉴定的蛋白质)。这些蛋白质参与了氮代谢调控、抗氧化行物质的合成、脂肪酸β-氧化、核骨架构造、[Fe-S]基团组装、物质跨膜运输、细胞分裂或作为分子伴侣和蛋白质酶起作用。putative ABC transporter ATP-binging protein、NtrX、nesprin-2 和Hslu 是本试验新发现的高等植物蛋白,我们主要从它们的保守结构域或在其他生物中的同源物来猜测它们的功能。实验结果为我们研究植物抗干旱机制提供了新线索和新视野。 Seabuckthorn (Hippophae rhamnoides L.) is widly distributed throughtout the temperatureresiogn of Europe and Asia and sub-tropical plateau zone of Asia. H. rhamnoides can adapatseveral different environments, and can tolerant several envioronmental stresses (e.g, lowtemperature, high temperature, drought and salty). It has been widely used in forest restoration asthe pioneer species in China. In present study, we applied H.rhamnoides subsp. Sinensis asexperimental materials to study its drought-tolerant mechanism, and expected to findpopulational difference in drought-tolerant mechanism that may exist among populations, and tryto get some insight in drought-tolerant mechanism of it at morecular level through analyzing thechange of leaf protein expression. Three populations from high altitude wet zone, low altitude wet zone and low altitude arid znoe,respectively, were applied in our experiment, and were subjected to drought. Drought increasedthe root/shoot ratio(RS), special leaf area, long-term water use efficinency, activity of antioxidantenzymes, proline content and abscisic acid (ABA) content, declined the net photosynthesis rate(A), average leaf area (ALA), total biomass (TB). Both two low altitude populations were moredrought-tolerant than the high altitude population, and different population applied differentstratedgies to tolerant oxidant stress and drought stress. The results of the exprement in 2004 showed that Daofu population was more drought-sensitivethan Jiuzhai population. Under drought conditions, leaf relative water content (RWC) greatlydecreased in Daofu population, but not in Jiuzhai population. The large loss of water in Daofupopulation resulted in a limitation on A mainly caused by non-stomatal factors, severer suppression in growth rate and a significant reduction in ascorbic acid (AsA) content, comparedwith Jiuzhai population. The greater increase in content of ABA and proline in Daofu populationmay be also induced by large loss in water, so that enable plants to cope with sever drought. In the exprement of 2005, drought significantly increased RS, activities of catalase (CAT),peroxidase (POD), glutathione peroxidase (GPX) and ascorbate peroxidase (APX), and alsosignificantly increased ABA and proline contents. On the other hand, compared with Daofupopulation, drought induced larger RS and activities of CAT, GPX and APX, and higher ABAcontent in Dingxi population, whereas gas exchange traits, e.g., stomatal limitation value (LS) andintercellular CO2 concentration (Ci), were less responsive to drought in Dingxi population thanthose in Daofu population. All these factors enable Dingxi population to tolerant drought betterthan Daofu population. The leaf protein profile of seabuchthorn subjected to drought was analyzed. Altogether 319proteins were detected in well-watered sample, four proteins disappeard by drought (putativeABCtransporter ATP-binding protein, hypothetical protein XP-515578, Hslu219and aunidentified protein), four only appeared under drought (a probable nitrogen regulation protein(NtrX), a 4-hydroxyphenylpyruvate dioxygenase , an unnamed protein product and an identified protein), 32 drought down-regulated proteins, and 14 drought up-regulated proteins (nine wereidentified: three large subunits of Rubisco, a hypothetical protein DSM3645-23351, a putativeacyl-COA dehydrogenase, a nesprin-2, a J-type-co-chaperone HSC20 and two unmatchedproteins). These proteins may involve in β-oxidation, cross-membrane transport, cell division,cytoskeleton stabilization, iron-sulfur cluster assembly, nitrogen metabolism regulation andantioxidant substance biosynthesis or function as molecular chaperone or protease. Four proteins(a putative ABC transporter ATP-binging protein, NtrX, nesprin-2, Hslu) were new found in highplants, and their functions were estimated from their conserved domain or their homologues inother organism. Our results provided new clue and new insight for us to study thedrought-tolerant mechanism in plants.
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目前,随着全球气候变化的加剧,水分短缺更加明显。在干旱与半干旱地区,水分胁迫是影响植物存活和生长的主要限制因子。同时,随着大气平流层中臭氧浓度的减少,过量的紫外辐射(UV-B)到达地球表面,一些地区的植物不可避免地受到增强UV-B 和水分胁迫的共同作用。文献表明在UV-B 增强的情况下,干旱表现为减弱或增强UV-B 对植物的影响,这与种、品种有一定的相关性。另外,脱落酸(ABA)是近年来研究报道最多的信息调控物质,与植物抗旱性途径有较大的关系,但其对植株抗UV-B 的影响还有待于研究。本论文以滇杨(Populus yunnanensis)为模式植物,从形态和生理方面研究了增强UV-B、干旱和脱落酸对它的影响,并探讨了UV-B 与干旱的互作效应以及喷施脱落酸对植株抗旱性和抗UV-B 能力的影响。主要研究结果如下:1. 增强的UV-B 和干旱胁迫都影响了滇杨的形态生长和生理生化反应。它们都导致了滇杨植株的株高、基茎、整株叶面积、平均叶面积、总生物量和净光合速率的显著降低,使得叶片增厚,过氧化物酶(GPX)活性升高,脯氨酸和花色素苷含量增加,膜脂过氧化程度增大。不同的是干旱显著降低了植株叶片数目,增大了根/冠比(Rs)、细根/总根比(Ft)、提高了内源ABA 含量、碳同位素(δ13C)以及紫外吸收物质含量和超氧化物歧化酶(SOD)的活性,而UV-B 对它们没有影响。干旱与UV-B 的复合作用加剧了任一单独胁迫对植株的抑制,表现为更小的株高、基茎、整株叶面积、平均叶面积、总生物量,更低的光合作用和更高的MDA 含量。而且UV-B 辐射降低了干旱胁迫下生物量分配的可塑性,表现为降低了干旱情况下的Rs 和Ft,ABA 的含量也显著下降,复合胁迫下脯氨酸含量和过氧化氢酶(CAT)的活性比任一单独胁迫时都要低。这些实验结果表明,增强的UV-B 与干旱的复合胁迫加剧了对植株的抑制作用。II2. 干旱情况下同时施加外源ABA 提高了植株的根/冠比、细根/总根比和单位面积叶重,即提高了干旱胁迫下植株对生物量分配的可塑性。而且外源ABA 使干旱胁迫下的长期用水效率、ABA 含量、脯氨酸含量、GPX 活性进一步增加,并有效调节了活性氧代谢的平衡,抑制了受旱植株MDA 的增加。结果表明,外源ABA 的喷施提高了滇杨植株的抗旱性。3. 在增强的UV-B 情况下,外源ABA 加剧了UV-B 对滇杨形态生长的抑制效果,表现为进一步降低了滇杨植株的整株叶面积、平均叶面积、单位面积叶重和总生物量,而且ABA 还降低了UV-B 胁迫下的净光合速率和脯氨酸的含量,增大了MDA 的含量。通过以上的数据我们可以看出,外源ABA 虽然提高了滇杨植株的抗旱性,但却加剧了UV-B 胁迫对植株的抑制作用。Currently, drought is one of the most serious environmental stresses. In arid and semi-aridregions, drought is a major constraint imposed on tree survival and growth. The decrease ofozone layer leads to a significant increase in ultraviolet-B (UV-B, 280-320 nm) radiationreaching the earth surface. In some places, plants suffer both UV-B and water stresssimultaneously. Their combination will increase or decrease the sensitivity of plants to UV-Bstress which lies on the species. On the other hand, abscisic acid (ABA), as a plant homoneand growth regulator, is better for plants resistant to drought stress, but it is uncleared aboutthe relationship between exogenous ABA and supplemental UV-B. In the present study, weemployed Populus yunnanensis Dode as a model species to characterize the growth andecophysiological responses of woody plants to supplemental UV-B, drought and exogenous ABA. The results are as follows:1. Both supplemental UV-B and drought affected the morphological, physiological andbiochemical responses of P. yunnanensis. They decreased the plant height, basal diameter,total leaf area, average leaf area, biomass and photosynthesis, and increased specific leaf mass,the activity of guaiacol peroxidase (GPX), the content of proline, anthocyanins andmalondialdehyde (MDA). However, drought decreased the leaf number and increasedroot/shoot ratio, fine root/total ratio, the activity of superoxide dimutase (SOD) and thecontents of ABA, carbon isotope composition (δ13C), UV-absorbing compounds. Whilesupplemental UV-B had no effects on them. The combination of drought and UV-Baugmented the growth inhibtion acting as further lower plant height and smaller basaldiameter, leaf area, biomass and higher MDA content. And compared with drought stress,root/shoot ratio and fine root/total root ratio decreased under the combination stresses. The photosynthesis, proline content and Catalase (CAT) activity became lower under combinationstresses than that of either stress lonely. According to these results, we suggested that,compared with the effect of stress lonely, the combination of supplemental UV-B and droughtdid not mitigate the harmful effect, but augmented it.2. Under drought conditions, exogenous ABA increased root/shoot ratio, fine root/total rootratio and the specific leaf mass. That was to say exogenous ABA increased plant plasticityunder drought conditions. Also ABA content, proline content, activity of GPX and δ13C wereenhanced further. In addition the enhancement of MDA was restrained. So the resultssuggested that exogenous ABA increased the seedling capacity of resistance to drought.3. Under supplemental UV-B conditions, exogenous ABA augmented the growth restrain ofUV-B to seedlings, which acted as further decreased leaf area, specific leaf mass and biomass.Compared with UV-B stress alone, proline content and photosynthesis were decreased andMDA content was increased under the combination of UV-B and ABA. These resultssuggested that although exogenous ABA increased the seedling capacity of resistance todrought, it augmented the growth restrain of supplemental UV-B to P. yunnanensis.
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杨树具有分布广、适应性强的特征,在生态环境治理和解决木材短缺方面均占有重要位置。青杨(Populus cathayana Rehd.)是青杨派树种的重要成员之一,也是生长较迅速、易繁殖的重要杨树资源。本研究选取了来自不同气候地区的青杨两种群为材料,采用植物生态学、生理学和生物化学的研究方法,系统地研究了青杨对干旱与遮荫、干旱与外源脱落酸(ABA)喷施的生长、形态、生理和生化响应及种群间差异,研究成果可为我国干旱半干旱地区的造林以及生态恢复提供理论依据和科学指导。主要研究结论如下:1.青杨在干旱胁迫下的适应机制为:生长性状及生物量的分配变化:干旱胁迫下虽然植株生长受抑,株高、基茎及各部分生物量都显著减小,但有相对较多的生物量向根部分配,根/冠比以及细/粗根比增加。青杨对干旱胁迫的光合作用表现为:干旱胁迫降低了青杨的净光合速率、蒸腾速率、气孔导度以及光合氮利用效率,提高了瞬时用水效率。干旱还引起了活性氧的产生,使得膜脂过氧化产物丙二醛(MDA)增加,同时也增强了植物抗氧化酶系统(如超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)活性的增加)及非酶系统的能力(如抗坏血酸(AsA)含量的增加)。干旱降低了植物叶片的相对含水量,而促进了渗透调节物质(游离脯氨酸及可溶性糖)的积累,增加了植物的渗调能力。干旱下青杨两种群的内源ABA含量显著增加,碳同位素组分(δ13C)也显著提高。这些结果证明植物遭受干旱胁迫后发生一系列的形态、生理和生化响应,这些变化能提高植物在干旱下的存活和生长能力。2.青杨两种群对干旱胁迫反应的种群差异:与来自湿润地区的汉源种群相比,来自干旱地区的乐都种群在干旱条件下生物量向根系分配的可塑性更强,同时具有更强的抗氧化系统能力,所受到活性氧的伤害也更少,并且累积更多的脯胺酸和ABA,具有更高的δ13C。这些都说明了乐都种群对干旱的适应性比汉源种群更强。两种群对干旱的响应差异应归于它们的用水策略的不同:汉源种群来自湿润地区,采用了耗水型的用水策略,抗旱能力较弱;而乐都种群,来自干旱地区,通常采用节水型的用水策略,有更强的抗旱能力。3.遮荫对青杨两种群抗旱性的影响:遮荫对青杨抗旱性的影响决定于遮荫程度的不同,我们的结果表明中度的遮荫可以有效的提高干旱下植物的生长,对干旱胁迫有明显的缓解作用,具体体现在中度遮荫下受旱植物的叶片相对含水量得到提高,使得植物体内水分状况得到了改善;光合速率并未降低,植物光合氮利用效率增加,说明中度的遮荫并未明显限制植物的碳获得;抗氧化酶活性与膜脂过氧化产物MDA含量的同时降低,说明中度遮荫下所受到的活性氧伤害减少;中度遮荫下的ABA及δ13C的变化也不如在全光下变化明显,这也说明中度遮荫缓解了干旱胁迫。但是重度的遮荫却对干旱胁迫有明显的加剧作用,主要表现在重度遮荫降低了植物的光合速率,严重抑制了植物的生长;同时重度遮荫下脯胺酸含量和抗氧化酶活性的急剧下降,导致了植物渗调能力的下降及膜脂过氧化产物MDA的显著升高;重度遮荫还显著降低了内源ABA的累积和δ13C,降低了植物的抗旱能力。此外,青杨两种群在对干旱和遮荫的响应中,也表现出种群差异。汉源种群,来自湿润且年日照辐射较少的地区,表现出相对更强的耐荫性和需水性。而乐都种群,来自干旱且年日照辐射丰富的地区,表现出相对更强的耐旱性和需光性。这说明了植物对环境胁迫的耐受性是其长期适应原生境的结果,并且来自不同气候地区的两种群在面临环境胁迫时会采取不同的生存策略。4. 外源ABA喷施对青杨两种群抗旱性的影响:外源ABA的喷施可以提高两种群的抗旱性,具体表现为外源ABA喷施促进了青杨根系的生长,显著提高了干旱下植物的根/冠比和细/粗根比,减少了比叶面积;在生理生化方面,外源ABA降低了干旱下植物叶片的气孔导度,降低了蒸腾速率和净光合速率,但提高了瞬时用水效率,提高了叶片的相对含水量,增加了干旱下植物的保水能力。外源ABA进一步增加了干旱下植物内源ABA的积累,促进了植物渗调物质如脯胺酸和可溶性糖的积累,增加了抗氧化酶系统(如SOD、APX、CAT)的活性和非酶系统AsA的含量,降低了活性氧(如超氧阴离子(O2和过氧化氢(H2O2))对植株的伤害。此外,外源ABA还进一步提高了干旱下植物的δ13C,提高了植物的长期用水效率,由此提高了植物的抗旱能力。另一方面,两种群对外源ABA和干旱的响应也有所差别。来自湿润地区的汉源种群,对干旱较为敏感,所受干旱的影响也较大,而外源ABA的喷施对汉源种群抗旱性的提高作用也更为突出。乐都种群,由于其长期适应干旱地区的生长,本身已具有较强的抗旱能力,因此外源ABA喷施对其抗旱性的提高不如对汉源种群的效果明显。由此我们可以得出对于一些抗性弱或干旱敏感的物种或者种群,可以采用外施ABA的方法来提高其抗性。Poplars play an important role in lumber supply, and are important component ofecosystems due to their wide distribution and well adaptation. Populus cathayana Rehd.,which belongs to Populus Sect. Tacamahaca Spach, is one of the most important resources ofpoplars for its fast growth and reproductive. In this study, different populations of P.cathayana were used as experiment material to investigate the adaptability to drought stressand population differences in adaptability, and the effects of shade and exogenous abscisicacid (ABA) application on the drought tolerance. Our results could provide a strongtheoretical evidence and scientific direction for the afforestation, and rehabilitation ofecosystem in the arid and semi-arid area, and provide a strong evidence for adaptivedifferentiation of different populations, and so may be used as criteria for species selectionand tree improvement. The results are as follows:1. A large set of parallel response to drought stress: Drought stress caused pronouncedinhibition of the growth and increased relatively dry matter allocation into the root. For thetwo populations, the shoot height, basal diameter and total biomass were decreased but theroot/shoot ratio and fine root/coarse root ratio were increased under drought conditions;Drought stress caused pronounced inhibition of photosynthesis, decreased the stomatalconductance, transpiration rate, and photosynthetic nitrogen-use efficiency (PNUE) butincreased the instantaneous water use efficiency. Drought significantly improved the levels ofreactive oxygen species and malondialdehyde (MDA) and to induce the entire set ofantioxidative systems including the increase of activities of superoxide dismutase (SOD),ascorbate peroxidase (APX), catalase (CAT) and ascorbate (AsA) content. Drought decreased the leaf relative water content (RWC) but improved the capability of osmotic adjustmentindicated by the higher proline accumulation. Drought also increased the ABA content andcarbon isotope composition (δ13C), which indicating the long period water use efficiency wasimproved under drought. These results demonstrate that there are a large set of parallelchanges in the morphological, physiological and biochemical responses when plants areexposed to drought stress; these changes may enhance the capability of plants to survive andgrow during drought periods.2. Difference in adaptation to drought stress between contrasting populations of P.cathayana: Compared with the Hanyuan population (wet climate), the Ledu population (dryclimate) showed higher root/shoot ratio and water use efficiency, exhibited higherantioxidative systems capability thus resulting in less oxidative damage, accumulated moreABA and free proline content under drought conditions. The results suggested that there weredifferent water-use strategies between the two populations. The Ledu population, whichcomes from dry climate region, with higher drought tolerance, may employ a conservativewater-use strategy, whereas the Hanyuan population, which comes from wet climate, withlower drought tolerance, may employ a prodigal water-use strategy. These variations indrought responses may be used as criteria for species selection and tree improvement.3. The effects of shade on the drought tolerance: The reduction in the availability of lightand water affected the morphological and physiological responses of the two P. cathayanapopulations. In addition, the light environment modified the growth responses of P.cathayana seedlings to varying water environments in different ways depending upon theintensity of the light levels considered. There is an apparent alleviation to drought effects bymoderate shade in P. cathayana seedlings, as indicated by the higher leaf RWC, and unchanged net photosynthesis and PNUE, as well as by the lower antioxditative enzymeactivity, MDA, ABA and δ13C levels, which implied moderate shade did not significantlylimited the carbon acquisition or inhibited the plant growth, but ameliorated the detrimentaleffects of drought. On the other hand, an apparent aggravation to drought effects by severeshade was also observed, as indicated by the pronounced decrease of plant growth and net photosynthesis, the lower total biomass, ABA level, δ13C, free proline content andantioxditative enzyme activity and higher MDA accumulation. By contrast, the twopopulations showed different responses to shade and drought. The Hanyuan population,which comes from a riparian basin having a relatively wet climate and less annual solarradiation, is more sensitive to drought but more tolerant to shade. The Ledu population, whichcomes from a mountainous plateau with less rainfall and with more annual solar radiation, ismore tolerant to drought but more sensitive to shade. The results demonstrated that theendurance of plants to stress is a result of long-term evolution and adaptation to theenvironment, as suggested by the different strategies employed by the P. cathayanapopulations originating from contrasting habitats when they were exposed to drought andshade.4. The effects of exogenous ABA application on the drought tolerance: For bothpopulations under drought conditions tested, exogenous ABA application significantlyimproved the root/shoot ratio, fine root/coarse root ratio, and decreased the specifical leaf area.On the physiological and biochemical traits, exogenous ABA application significantlydecreased stomatal conductance, transpiration rate and net photosythesis but increased theinstance water use efficiency and leaf RWC. On the other hand, exogenous ABA applicationsignificantly increased endogenous ABA, proline, solube sugar and AsA content, as well asSOD, APX and CAT activities, thus reduced the damage of reactive oxygen species. Moreover,the long period water use efficiency as indicated by δ13C was also improved by exogenousABA application. In additionally, there was different responsive between the two populationsto drought and exogenous ABA application. The Hanyuan population, which comes from wetclimate region, is more sensitive to drought, and the effect of exogenous ABA is moreobviously than that in the Ledu population, which comes from dry climate region and is moredrought-responsive. Therefore, we can use exogenous ABA application to improve theresistance of plants, especially for the drought- sensitive species or populations.
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克隆植物被认为比非克隆植物更宜于利用异质性环境。在复杂的空间异质环境中,克隆植物可能形成了各种有效利用环境异质性的适应对策。对于克隆植物适应机制的研究,前人已做了大量的工作,特别是从形态和生物量分配等方面对简单异质生境下克隆植物的克隆整合和克隆分工进行了详细的研究。本研究以分布广泛的克隆植物东方草莓(Fragaria orientalis)作为研究对象,应用野外调查和实验生态学方法,采用多对比度单资源模型和不同向双资源模型,从形态和生理生态的角度,研究复杂异质生境下克隆植物的整合和分工及其耗益问题,分析不同类型的生境对克隆植物整合和分工的修饰作用,进而探讨克隆植物对异质生境的适应策略。克隆构型和分株种群特征是植物克隆生长及其生态适应对策研究的基本内容。本文通过野外调查,研究在不同光照条件下东方草莓克隆构型、分株种群特征以及点分布格局。结果表明:东方草莓的克隆构型随光照发生相应的变化,低光照下其匍匐茎节间长和分枝角度均增大而分枝强度减小;随光照减弱,东方草莓分株种群的生物量、根冠比和分株种群密度显著降低;不同光照下东方草莓分株均以随机分布为主但不同尺度下有所差异,其分布格局强度依次为旷地<林缘<林下。结合克隆植物对资源的利用对策,探讨了克隆构型和分株种群特征以及分布格局随环境条件变化的生态适应意义。不同生境斑块条件下克隆植物可能采取不同的适应对策。采用盆栽实验,研究不同水分对比度下克隆整合及其生理生态特征,并对单向和交互资源中东方草莓的克隆整合做了对比研究。结果显示:高的水分对比度能够促进东方草莓的克隆整合,并能刺激相连分株增加光合作用,东方草莓体内的氧化—抗氧化系统也II随对比度做出相应的反应。耗-益分析表明胁迫分株的受益是以供给分株的损耗为代价的,但从克隆片段总体来说是受益的。单向资源中东方草莓生长的绝对值高于交互资源,但耗-益分析表明生长于交互资源下东方草莓的克隆整合获益大于生长于单向资源下东方草莓的克隆整合获益。长期生长于特定生境的克隆植物,在进化过程中其克隆整合和克隆分工在对资源异质性的适应策略方面可能有所侧重。采用盆栽实验对来自不同海拔梯度的东方草莓的克隆整合和克隆分工对异质资源的适应对策进行了研究。实验结果表明,来自高海拔的东方草莓可塑性较差。来自两个海拔的东方草莓对切断匍匐茎的表现有所差异,总体上切断匍匐茎对来自高海拔的东方草莓影响更大些。另外,来自高海拔的东方草莓表现出更高的克隆分工。IIIClonal plants are known to be more suitable for the habitats of heterogeneousresources than nonclonal plants, perhaps due to their well developed adaptivestrategies to environmental heterogeneity. Many studies have been done on theadaptive mechanisms of clonal plants, especially on the clonal integration anddivision of labor with morphology and biomass allocation under simpleheterogeneous habitats. Based on field surveys, laboratory experiments, multi-contrastunidirectional resource model and reciprocal resource model, Fragaria orientalis, aRosaceae stoloniferous herb that widely distributes in China, was used to study thisplant’s morphological and physiological responses to complicated heterogeneoushabitats in terms of its clonal integration, division of labor and cost-benefit, as well astheir modifications by different habitats, so as to better understand the adaptivestrategies of clonal plants under heterogeneous environments.Clonal architecture and ramet population characteristics are of the major concernin the studies on growth and adaptive strategies of clonal plants. Clonal architecture,ramet population characteristics and spatial point pattern of F. orientalis underdifferent light intensity were studied with field observations. The results showed that,clonal architecture changed with light availability: Internode-lengths and branchangels of stolons were larger while branch intensities were smaller under lower lightintensity than those under higher light intensity; Biomass of ramet population,root-shoot ratio and density of ramet population decreased significantly with reduce oflight intensity; Under all light intensities, spatial pattern of ramets was mainlyrandomly distributed but it changed with different scales, with pattern intensity as:open space < forest edge < understory. Adaptation significance of the clonal architecture, the ramet population characteristics and the spatial pattern changing withdifferent environments was discussed according to these results.Clonal plants may take different adaptive strategies under different patches. Withpot culture, clonal integration and physiological parameters of F. orientalis underdifferent water contrasts were studied, and clonal integration under unilateralresources and reciprocal resources were also compared. The results suggested that,high water contrast improve the clonal integration of F. orientalis and increase thephotosynthesis of connected ramets. Oxidative and antioxidative system of F.orientalis also responded with changing water contrasts. According to cost-benefitanalysis, the drought-stressed ramets obtained benefits from the connectedwell-watered ramets, and as a whole, the clonal fragment could also get benefits.Growth of F. orientalis in homogeneous resources was better than that inheterogeneous resources, but the whole plant got more benefit through clonalintegration in heterogeneous resources than in homogeneous resources.Pot culture experiments were also used to study the adaptive strategies inutilizing heterogeneous resources by the plant populations from different altitudes.The results showed that, F. orientalis from alpine zones were shorter and lessexpanded with poorer clonal plasticity than those from middle mountains. F.orientalis from two different altitudes showed different responses to stolon severing,and as a whole, stolon severing had more influence on F. orientalis from alpine zones.In addition, F. orientalis from alpine zones exhibited higher division of labor, whichsuggested that clonal plants from different habitats develop their own adaptivemechanisms in their clonal integration and division of labor in response toenvironmental heterogeneity.
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干旱环境常常由于多变的降水事件和贫瘠土壤的综合作用,表现出较低的生产力和较低的植被覆盖度。全球性的气候变暖和人类干扰必将使得干旱地区缺水现状越来越严竣。贫瘠土壤环境中已经很低的有效养分含量也将会随着干旱的扩大而越来越低。干旱与半干旱系统中不断加剧的水分与养分的缺失将严重限制植物的生长和植被的更新,必然会使得已经恶化的环境恶化速率的加快、恶化范围的加大。如何抑制这种趋势,逐步改善已经恶化的环境是现在和将来干旱系统管理者面临的主要关键问题。了解干旱系统本土植物对未来气候变化的适应机制,不仅是植物生态学研究的重要内容,也对人为调节干旱环境,改善干旱系统植被条件,提高植被覆盖度具有重要的实践意义。 本研究以干旱河谷优势灌木白刺花(Sophora davidii)为研究对象,通过两年大棚水分和施N控制实验和一个生长季野外施N半控制实验,从植物生长-生理-资源利用以及植物生长土壤环境特征入手,系统的研究了白刺花幼苗生长特性对干旱胁迫和施N的响应与适应机制,并试图探讨施N是否可调节干旱系统土壤环境,人工促进干旱条件下幼苗定居,最终贡献于促进植被更新实践。初步研究结论如下: 1)白刺花幼苗生长、生物量积累与分配以及水分利用效率对干旱胁迫和施N处理的适应白刺花幼苗株高、基径、叶片数目、叶面积、根长、生物量生产、相对含水量和水分利用效率随着干旱胁迫程度的增加而明显降低,但地下部分生物量比例和R/S随着干旱胁迫程度的增加而增加。轻度施N处理下幼苗株高、基径、叶片数目、叶片面积和生物量生产有所增加。但重度施N处理下这些生长指标表现出微弱甚至降低的趋势。严重干旱胁迫条件下,幼苗叶面积率、R/S、相对含水量和水分利用效率也以轻度施N处理为最高。 2)白刺花幼苗叶片光合生理特征对干旱胁迫和施N处理的适应叶片光合色素含量和叶片光合效率随着干旱胁迫程度的增加而显著降低,并且PS2系统在干旱胁迫条件下表现出一定程度的光损害。但是比叶面积随着干旱胁迫程度的增加而增加。在相对较好水分条件下幼苗净光合速率的降低可能是因为气孔限制作用,而严重干旱胁迫条件下非气孔限制可能是导致幼苗叶片光合速率下降的主要原因。叶片叶绿素含量、潜在光合能力、羧化效率、光合效率以及RUBP再生能力等在施N处理下得到提高,并因而改善干旱胁迫条件下光合能力和效率。虽然各荧光参数对施N处理并无显著的反应,但是干旱胁迫条件下qN和Fv/Fm在轻度施N处理下维持相对较高的水平,而两年连续处理后在严重干旱胁迫条件下幼苗叶片光合效率受到重度施N处理的抑制,并且Fv/Fm和qN也在重度施N处理下降低。 3)白刺花幼苗C、N和P积累以及N、P利用效率对干旱胁迫和施N处理的适应白刺花幼苗C、N和P的积累,P利用效率以及N和P吸收效率随干旱胁迫程度的增加而显著降低,C、N和P的分配格局也随之改变。在相同水分处理下,C、N和P的积累量、P利用效率以及N和P吸收效率在轻度施N处理下表现为较高的水平。然而,C、N和P的积累量和P利用效率在重度施N处理下不仅没有表现出显著的正效应,而且有降低的趋势。另外,在相同水分条件下白刺花幼苗N利用效率随着施N强度的增加而降低。 4)白刺花幼苗生长土壤化学与微生物特性对干旱胁迫和施N的适应白刺花幼苗生长土壤有机C、有效N和P含量也随干旱胁迫程度的增加而明显降低。干旱胁迫条件下土壤C/N、C/P、转化酶、脲酶和碱性磷酸酶活性的降低可能表明较低的N和P矿化速率。尽管微生物生物量C、N和P对一个生长季干旱胁迫处理无显著反应,但微生物生物量C和N在两年连续干旱胁迫后显著降低。土壤有机C和有效P含量在轻度施N处理下大于重度施N处理,但是有效N含量随着施N强度的增加而增加。微生物生物量C和N、碱性磷酸酶和转化酶活性也在轻度施N处理下有所增加。但是碱性磷酸酶活性在重度施N处理下降低。 5)野外条件下白刺花幼苗生长特征及生长土壤生化特性对施N的适应植物生长、生物生产量、C的固定、N、P等资源的吸收和积累、其它受限资源的利用效率(如P)在轻度施N处理下均有所增加,但N利用效率有所降低。幼苗生物生产量及C、N和P等资源的分配格局在轻度施N处理下也没有明显的改变。白刺花幼苗叶片数目、生物生产量和C、N、P的积累量在重度施N处理下虽然也相对于对照有所增加,但幼苗根系长度显著降低。生物量及资源(生物量、C、N、P)在重度施N处理下较多地分配给地上部分(主要是叶片)。另外,土壤有机C、全N和有效N含量随外源施N的增加而显著增加,土壤pH随之降低,但土壤全P含量并无显著反应。其中有机C含量和有效P含量以轻度施N处理最高。微生物生物量C、N和P在轻度施N处理下也显著增加,而微生物生物量C在重度施N处理下显著降低。同时,转化酶、脲酶、碱性磷酸酶和中性磷酸酶活性在施N处理下也明显的提高,但酸性磷酸酶和过氧化氢酶活性显著降低,其中碱性磷酸酶和中性磷酸酶活性以轻度施N处理最高。 综合分析表明,干旱河谷水分和N严重限制了白刺花幼苗的生长。施N不能完全改变干旱胁迫对白刺花幼苗的抑制的作用,但是由于施N增加土壤N有效性,改善土壤一系列生物与化学过程,幼苗的生长特性也对施N表现出强烈的反应,表现为植物结构与资源分配格局的改善,植物叶片光合能力与效率的提高,植物生长以及利用其他受限资源(如水分和P)的效率的增加,致使植物自身生长及其生长环境在干旱环境下得到改善。但是过度施N不仅不能起到改善干旱胁迫下植物生长环境、促进植物生长的作用,反而在土壤过程以及植物生长过程中加重干旱胁迫对植物的伤害。因此,建议在采用白刺花作为先锋种改善干旱河谷系统环境的实践中,可适当施加N以改善土壤环境,调节植物利用与分配资源的效率,促进植物定居,得到人工促进种群更新的目的。但在实践过程中也要避免过度施N。 Arid regions of the world are generally noted for their low primary productivity which is due to a combination of low, unpredictable water supply and low soil nutrient concentrations. The most serious effects of global climate change and human disturbances may well be those which related to increasing drought since drought stress has already been the principal constraint in plant growth. The decline in total rainfall and/or soil water availability expected for the next decades may turn out to be even more drastic under future warmer conditions. Nevertheless, water deficit is not the only limiting factor in arid and semiarid environments. Soils often suffer from nutrient (especially N and P) deficiencies in these ecosystems, which can also be worsened by climate change. How to improve the poor soil quality and enhance the vegetation coverage is always the problem facing ecosystem managers. The adaptive mechanisms of native plant to future climate change is always the focus in plant ecology, it also plays important roles in improving vegetation coverage by manual controlled programmes. Sophora davidii is a native perennial shrub of arid valleys, which is often predominant on eroded slopes and plays a vital role in retaining ecological stability in this region. It has been found that S. davidii was better adapted to dry environment than other shrubs, prompting its use for re-vegetation of arid lands. A two-years greenhouse experiment and a field experiment were conducted in order to understand the adaptation responses of Sophora davidii seedlings to different water and N conditions, and further explore if additional N supply as a modified role could enhance the adaptation ability of S. davidii seedlings to dry and infertile environment. Two-month old seedlings were subjected to a completely randome design with three water (80%, 40% and 20% water field capacity (FC)) and three N supply (N0: 0, Nl: 92 and Nh: 184 mg N kg-1 soil) regimes. Field experiment was arranged only by three N supplies in the dry valley. 1) The growth, biomass partitioning and water-use efficiency of Sophora davidii seedlings in respond to drought stress and N supply Seedlings height, basal diameter, leaf number, leaf area, root length, biomass production, relative water content (RWC) and WUE were decreased with increase of drought stress. An increase in below-ground biomass was observed indicating a higher root/shoot ratio (R/S) under drought stress conditions. Low N supply increased seedlings height, basal diameter, leaf number, leaf area, and biomass production, but decreased root length. In contrast, these growth characteristics showed little or negative effect to high N supply treatment. Leaf percentages increased with increase of N supply, but fine root percentages decreased. In addition, Low N supply rather than the other two N treatments increased leaf area ratio (LAR), leaf/fine root mass ratio (L/FR), R/S and RWC under severe drought stress (20%FC), even though these parameters could increase with the high N supply treatment under well-watered condition (80%FC). Moreover, Low N supply also increased WUE under three water conditions, but high N supply had little effect on WUE under drought stress conditions (40%FC and 20%FC). 2) Leaf gas exchange and fluorescence parameters of Sophora davidii seedlings in respond to drought stress and N supply Leaf area (LA), photosynthetic pigment contents, and photosynthetic efficiency were decreased with increase of drought stress, but specific leaf area (SLA) increased. Photodamage in photosystem 2 (PS2) was also observed under drought stress condition. The decreased net photosynthetic rate (PN) under relative well-watered water conditions might result from stomatal limitations, but the decreased PN under other hand, photosynthetic capacity by increasing LA, photosynthetic chlorophyll contents, Pnmax, CE, Jmax were increased with increase N supply, and photosynthetic efficiency was improved with N supply treatment under water deficit. Although N supply did a little in alleviating photodamages to PS2 caused by drought stress, low N supply enhanced qN and kept relative high Fv/Fm under drought stress condition. However, high N supply inhibited leaf photosynthetic efficiency, and declined Fv/Fm and qN under severe drought stress condition after two year continues drought stress and N supply. 3) Carbon accumulation, nitrogen and phosphorus use efficiency of Sophora davidii seedlings in respond to drought stress and N supply C, N and P accumulation, NUE , N and P uptake efficiency (NUtE and NUtE ) P N P were decreased with increase of drought stress regardless of N supply. On the other hand, the S. davidii seedlings exhibited strong responses to N supply, but the responses were inconsistent with the various N supply levels. Low N supply rather than the other two N treatments increased C, N and P accumulation, improved NUEP, NUtE and NUtE under corresponding water condition. In contrast, high N supply N P did few even depressed effects on C, N and P accumulation, and NUEP, although NUtEN and NUtEP could increase with high N supply under corresponding water conditions. Even so, a decrease of NUEN was observed with increase of N supply under corresponding water conditions. 4) Soil microbial and chemical characters in respond to drought stress and N supply The content of soil organic C, available N and P were decreased with increase of drought stress. Decreases in C/N and C/P, and invertase, urea and alkaline phosphatase activity were also observed under drought stress conditions, indicating a lower N and P mineralization rate. Although microbial biomass C, N and P showed slight responses to drought stress after one growth period treatment, microbial biomass C and N were also decreased with increase of drought stress after two year continuous treatment. The content of soil organic C and available P showed the stronger positive responses to low N supply than which to high N supply, although than the other two N treatments increased microbial biomass N and invertase activity under severe drought stress condition, even though invertase activity could increase with high N supply treatment under relative well-water conditions. Moreover, low N supply treatment also increased C/P and alkaline phosphatase activity which might result from higher P mineralization, but high N supply did negative effects on alkaline phosphatase activity. 5) The growth characteristics of Sophora davidii seedlings and soil microbial and chemical characters in respond to N supply under field condition Low N supply facilitated seedlings growth by increasing leaf number, basal diameter, root length, biomass production, C, N and P accumulation and absorption, and enhancing the use efficiency of other limited resources as P. Compared to control, however, low N supply did little effect on altering biomass, C, N and P portioning in seedlings components. On the contrary, high N supply treatment also increased leaf number, biomass and C, N and P accumulation relative to control, but significantly decreased root length, and altered more biomass and resources to above-ground, which strongly reduced the ability of absorbing water under drought condition, and thus which might deep the drought stress. In addition, N supply increased soil C, N and available N content, but declined pH and showed little effects on P content. Low N supply showed higher values of soil C and available P content. Low N supply also increased microbial biomass C, N and P, although high N supply decreased microbial biomass C. N supply significantly enhanced soil invertase, urea, alkaline and neutral phosphratase activity, while declined acid phosphratase and catalase activity. Low N supply exhibited higher alkaline and neutral phosphratase activity compared to the others. The results from this study indicated that both drought and N limited the growth of S. davidii seedlings and their biomass production. Regardless of N supply levels, drought stress dramatically reduced the seedlings growth and biomass production. Although plant growth parameters, including basal diameter, height, leaf number, and biomass and their components were observed to be positive responses to low N supply, N supply alone can not alter the diminishing tendency which is caused by drought. available N content increased with increase N supply. In addition, low N supply rather These findings imply that drought played a primary limitation role and N was only the secondary. Even so, appropriate N supply was seemed to enhance the ability that S. davidii seedlings adapted to the xeric and infertile environment by improving soil processes, stimulating plant growth, increasing recourses accumulation, enhancing use efficiency of other limited resources, and balancing biomass and resources partitioning. Appropriate N supply, therefore, would be recommended to improve S. davidii seedling establishment in this region, but excess N supply should be avoided.
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利用田间小区试验研究了不同土壤水分条件下N肥对根系生长(根长、根重和根冠比)的影响。结果表明,玉米拔节期和开花期无论水分条件如何,施N肥可增加其总根长、表层根长和根重并使根冠比下降;灌浆期施N肥可增加总根长和表层根长,但正常水分条件下N肥使根重和根冠比下降,而干旱条件下N肥对根重和根冠比则无影响。不同处理根冠比大小与N吸收和分配有很大关系。