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岷江上游干旱河谷区水土流失强烈,地质灾害频繁,生态环境十分脆弱,而土壤条件恶劣(水分不足和养分缺乏)是阻碍该区植被恢复的关键因子,因此研究水分和乡土灌木生长对土壤的影响对该区的生态恢复具有指导意义。本文通过定点模拟实验,选取三种优势豆科灌木为研究对象,分别是白刺花(Sophora davidii)、小马鞍羊蹄甲(Bauhinia faberi var. microphylla)和小雀花(Campylotropics polyantha),设置5 个水分梯度,分别为100%、80%、60%、40%和20%田间持水量(FC),对栽种植物与不种植物下土壤理化性质和酶活性进行测定分析,系统比较和研究了不同水分条件和不同乡土灌木生长对干旱河谷区土壤结构、养分循环、酶活性以及微生物量的影响。主要结果如下:1. 无论生长植物与否,土壤的毛管持水量和毛管孔隙度都随着水分含量的减少而降低,最大持水量、总孔隙度和容重变化不大,相应地,土壤中的非毛管孔隙随含水量的减少而升高。各水分条件下,种植植物的毛管持水量和毛管孔隙度低于无植物生长的土壤,非毛管孔隙度相应地高于无植物土壤。土壤含水量在100%-40% FC 时,三种豆科灌木的毛管持水量和毛管孔隙度存在差异,而20% FC 条件下,三种豆科灌木土壤的物理性质基本相同。2. 水分胁迫影响土壤中养分的矿化和积累,主要表现在降低了水溶性碳和铵态氮的含量,中等程度胁迫时(60% FC)促进了有机碳和硝态氮的富集,对速效钾和有效磷没有明显作用。种植豆科灌木后各水分梯度上都增加了有机碳、铵态氮、速效钾和有效磷的积累。增加程度上三种豆科灌木间有一定差异,对于土壤有机碳总量,种植白刺花和小马鞍羊蹄甲明显高于小雀花,同样的情况还出现在铵态氮和速效钾上,但是对于有效磷,种植小雀花后的增加程度则明显高于白刺花和小马鞍羊蹄甲。种植豆科灌木不仅增加了土壤养分的相对含量,也改变了其在水分梯度上的变化趋势及其变化幅度,这种作用主要体现在碳元素和氮元素上。3. 无植物生长时脲酶活性随水分含量的减少而升高,水分胁迫对磷酸酶和过氧化氢酶的作用不显著,蔗糖酶也保持在相对较高的水平。种植植物后,蔗糖酶、磷酸酶活性与无植物时相比有较大幅度的提高,种植白刺花的脲酶活性也升高,其升高的程度在不同水分含量时不同。种植植物还降低了酶活性在水分梯度上的变幅,使之在水分梯度间的差异显著性降低。脲酶活性在指示土壤性质改变方面是较敏感的指标,其它三种酶在不同植物间的差异不明显。4. 在无植物生长时,中等程度的水分胁迫(60% FC)提高了土壤微生物量碳含量,过高或过低的土壤水分均不利于微生物碳的积累。种植小马鞍羊蹄甲后微生物量碳在水分梯度上的变化趋势与无植物生长时一致,而种植白刺花和小雀花后微生物量碳随着水分含量的减少而降低。不同种类植物的微生物量碳在水分梯度上的变化特征也不同,100% FC 条件下三种植物间没有差异,80%和60% FC 条件下小马鞍羊蹄甲显著高于白刺花和小雀花,40%和20% FC 条件下白刺花和小马鞍羊蹄甲也显著高于小雀花,说明不同种类植物随着干旱胁迫程度的加深微生物量碳的降低幅度不同,在极度干旱时,白刺花和小马鞍羊蹄甲土壤依然保持了较高的微生物活性,而小雀花土壤微生物量则明显下降。The dry valley of the upper reaches of the Minjiang River is seriously degradedmountain ecosystem. It was endangered by extremely soil lost and frequentlygeological disaster. Previous studies showed that short of water and nutrients in soilwas the principal limiting factors of vegetation restoration in this area. The typical soiland three dominant leguminous shrubs Sophora davidii, Bauhinia faberi var.microphylla and Campylotropics polyantha in upper reaches of arid Minjiang Rivervalley were considered as experimental material. Two-month old seedlings of eachspecies were exposed to five water supplies (100%, 80%, 60%, 40% and 20% waterfield capacity (FC)) in a temperature and light-controlled greenhouse. Afterthree-month water treatment, soil physiochemical variables and soil microbialactivities were determined by conventional methods. The main results showed that:1. Soil capillary capacity and capillary porosity decreased along water supplyregimes in all treatments, while saturated water capacity, total porosity and bulkdensity kept in a relatively stable level, as a result, the non-capillary porosity andcapacity increased with decrease of water supply. Compared to non-planted soil, theplant-soil systems had a higher non-capillary porosity and capacity, suggestingappropriate oxygen was present in soil to maintain the living of microorganism. Soilof three type shrub species shared the same capillary capacity and capillary porosityunder 20% FC.2. Water soluble carbon and NH4+-N decreased in response to water stress, whiletotal organic carbon and NO3--N promoted by moderate water stress and inhibited by 100% and 20% FC. Total organic carbon, NH4+-N, rapidly available K and availableP increased after the planting of leguminous shrubs in five water supply regimescompared to non-planted soil. For TOC, NH4+-N and rapidly available K, thepromotion effect was higher in S. davidii and B. faberi var. microphylla than C.polyantha planted soil, while available P displayed the opposite side. The planting ofshrubs also reduced the variance of observed traits along water supply gradients.3. Drought stress increased urease activity in non-planted soil, while insignificantdifferences were observed in phosphatase and catalase activity among five watersupply regimes. The planting of leguminous shrubs facilitated the β-glucosidase andphosphatase activity compared to the non-planted soil. It also reduced the variance ofenzyme activity along water supply gradients. Urease was more sensitive to waterstress than other three enzymes.4. Soil water content significantly affected microbial biomass carbon andCmic:Corg. S. davidii and B. faberi var. microphylla showed more drought toleranceability than C. polyantha, attributing not only to their relatively smaller variance ofmicrobial biomass carbon along soil water supply gradients, but also to the highlevel of microbial activity under severe water stress. S. davidii and B. faberi var.microphylla benefited reproduction of soil microorganism at 60%-80% FC, whilesevere drought limited it due to the competition of water and nutrients between plantand soil microorganism.
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随着全球气候变暖和温室效应加剧,干旱和荒漠化成为威胁人类生存和发展的主要 灾害,许多被子植物对干旱胁迫的生理、生态和生化响应已逐步得以报道,但很少有开 展干旱胁迫对雌雄异株植物的影响方面的研究。由于这类植物在长期进化过程中已经在 生长、性比、生殖格局、空间分布、资源配置和生物量分配等方面形成了明显的性别差 异,因此,干旱胁迫必将对其雌雄植株产生不同的生理生态影响。本研究以青杨为模式 植物,采用植物生态、生理及生物化学等研究方法,系统研究青杨雌雄植株在常温、增 温以及喷施外源脱落酸的条件下对干旱胁迫的响应,揭示其在生长形态、生物量分配、 光合作用、用水效率和生理生化等方面的性别间差异。主要研究结果如下: 1. 青杨雌雄植株对干旱胁迫的综合响应。 与较好水分条件相比,干旱胁迫显著降低了青杨雌雄植株的光合作用和生长发育, 影响了许多生理生化过程,并导致雌雄植株在生长发育、气体交换、用水效率、膜脂抗 氧化和抗氧化系统酶活性方面表现出显著的性别间差异。在较好水分条件下,雌雄植株 之间在株高、基径、生物量、净光合速率、蒸腾速率、用水效率以及丙二醛、脱落酸和 游离脯氨酸等生化物质含量方面均无显著差异。但在干旱胁迫下,雄株在生长发育、气 体交换、水分利用效率、膜脂过氧化保护和抗氧化系统酶活性方面均显著高于雌株,表 现出比雌株更高的株高、基径、叶面积、总叶片数、总生物量、总色素含量、类胡萝卜 素含量、净光合速率、蒸腾速率、羧化效率、光系统II最大光化学效率、内在水分利用 效率、碳同位素组分、过氧化氢酶和过氧化物酶活性等,而在CO2补偿点、比叶面积、 叶绿素a/b、丙二醛、脱落酸和超氧化物歧化酶活性等指标上显著低于雌株。与雌株相比, 雄株表现出更高的干旱胁迫适应能力,而雌株的生长发育和生理生化过程更易遭受干旱 胁迫的影响。 2. 干旱胁迫下的青杨雌雄植株对增温处理的综合响应 与环境温度相比,增温在干旱胁迫前后均显著促进了雌雄植株的生长发育、气体交 换,降低水分利用效率,影响生化物质含量,并促使青杨雌雄植株之间在干旱胁迫下表 现出显著的差异。在较好水分条件下,增温导致雌株的株高、基径、叶面积、总叶片数、 总生物量和超氧化物歧化酶活性显著高于雄株,而用水效率、丙二醛、脱落酸和游离脯 氨酸、抗坏血酸过氧化物酶和过氧化物酶活性低于雄株。在干旱胁迫下,增温将导致雄 株的株高、基径、叶面积、总生物量、净光合速率、蒸腾速率、气孔导度、总色素含量、 相对含水量、过氧化氢酶和抗坏血酸过氧化物酶活性等显著高于雌株,而光系统II 最大 光化学效率、内在水分利用效率、碳同位素组分、丙二醛、脱落酸、游离脯氨酸和超氧 化物歧化酶活性显著低于雌株。与雄株相比,水分较好条件下的增温有利于促进雌株的 生长发育,并在生理生态特征上优于雄株。而干旱胁迫下的增温则加剧了水分胁迫强度, 致使雌株的生长发育遭受比雄株更多的负面影响。 3. 干旱胁迫下的青杨雌雄植株对喷施外源脱落酸处理的综合响应 与对照相比,在干旱胁迫下喷施外源脱落酸可显著增加青杨雌雄植株的生长发育、 气体交换、降低水分利用效率,影响了生化物质含量,并导致青杨雌雄植株之间在干旱 胁迫下表现出显著的生理生态差异。在干旱胁迫下,喷施外源脱落酸致使雌株的株高、 叶面积、叶干重、细根干重、总生物量、净光合速率、蒸腾速率、气孔导度、光系统II 最大光化学效率、非光化学淬灭系数、相对含水量、总光合色素、类胡萝卜素、脱落酸、 超氧化物歧化酶和过氧化物酶活性的增加量显著高于雄株,而根重比、根冠比、细根/ 总根、比叶面积、内在水分利用效率、碳同位素组分、丙二醛、脯氨酸、过氧化氢酶和 抗坏血酸过氧化物酶活性等指标的减少量上显著低于雄株。与对照相比,干旱胁迫下的 喷施外源脱落酸则一定程度能减缓植株遭受胁迫的压力,促进植株生长和气体交换,减 少了植株体内的过剩自由基数量,并促使雌株的生长发育和光合能力显著提高,增强其 抗干旱胁迫能力。 With development of global warming and greenhouse effect, drought and desertification have been became main natural disasteres in resent years. Studies on ecophysiological responses of most angiosperm species to environmental stress have been reported, but little is known about dioecious plant responses to drought stress. Since significant differences on growth, survival, reproductive patterns, spatial distribution, as well as resource allocation between males and females of dioecious plant have been formed during evolutionary process, sexual different ecophysiological responses should be caused by drought stress. In this experiment, Populus cathayana Rehd. was used as model plant to study the sex-related responses to drought by using the ecological, physiological and biochemical methods under normal atmospheric temperature, elevated temperatures and exogenous abscisic acid (ABA) application treatment respectively, and to expose the sexual differences in growth, biomass allocation, photosynthesis, water use efficiency and some biochemical material contents in the males and females of dioecious plant. The results are follows: 1. A large set of parallel responses of males and females of P. cathayana to drought stress Compared with well-watered treatment, drought significantly decreased growth and photosynthesis of P. cathayana individuals, affected some physiological and biochemical processes, and induced males and females to exhibit obvious sexual differences in growth, gas exchange, water use efficiency, lipid peroxidation protection and antioxidant defenses enzyme system. Under well-watered treatment, there were no significant sexual differences in height growth (HG), basal diameter (BD), dry matter accumulation (DMA), net photosynthesis rate (A), transpiration (E), water use efficiency (WUE), and malondialdehyde (MDA), abscisic acid (ABA) and praline (Pro). However, under drought stress, males were found to exhibit higher HG, BD, leaf area (LA), total leaf number (TLA), DMA, total chlorophyll contents (TC), carotenoids content (Caro), A, E, carboxylation efficiency (CE), the maximum efficiency of PSII (Fv/Fm), intrinsic water use efficiency (WUE ), carbon isotope composition (δ13C), catalase (CAT), peroxidase (POD) and lower CO2 compensation point (Γ), specific leaf area (SLA), chlorophyll a/b ratio (Chla/Chlb), MDA, ABA and superoxide dismutase (SOD) than females. The results suggest that males possess greater drought resistance than do females and females suffer more negative effect on growth and development, physiological and biochemical processes than males under drought stress. 2. A large set of parallel responses of drought-stressed males and females of P. cathayana to elevated temperatures Compared with environmental temperature, elevated temperature treatment significant increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents of P. cathayana individuals, and induced males and females to exhibit obvious differences under drought stress. Under good water condition, elevated temperature treatment caused females to show significant higher HG, BD, LA, TLN, DMA, SOD activity, and great lower WUE, MDA, ABA, Pro, ascorbate peroxidase (APX) and POD than do males. On contrary, under drought condition, elevated temperature treatment induced males to exhibit higher HG, BD, LA, DMA, A, E, stomatal conductance (gs), relative water content (RWC), CAT, APX activity but lower Fv/Fm, WUE, δ13C, MDA, ABA, Pro, SOD activity than do females. The results suggest that females will benefit from elevating temperature under good water condition by possessing better ecophysiological processes than that of males, but will suffer from greater negative effects than do males when grown under drought stress with elevated temperature treatment. 3. A large set of parallel responses of drought-stressed males and females of P. cathayana to exogenous ABA application Compared with controls, exogenous ABA application under drought greatly increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents in P. cathayana individuals, and induced males and females to exhibit obvious sexual differences under drought. Under drought stress, exogenous ABA application induced females to exhibit more increases in HG, LA, leaf weight (LW), fine root weight (FRW), DMA, A, E, g, Fv/Fm, non-photochemical quenching coefficient (qN), RWC, TC, Caro, ABA, SOD, POD s activity than males, but to show lower decreases in root/weight ratio (RWR), root mass/foliage area ratio (RF), fine root/total root ratio (FT), SLA, WUE, δ13C, MDA, Pro, CAT, APX than males. The results suggest that exogenous ABA application under drought stress will eliminate negative damages caused by drought stress at a certain extent,promote the growth and gas exchange of plant and decrease the number of superfluous 1O2 in plant cells of males and females of P. cathayana. Furthermore, exogenous ABA application promoted more drought resistance in females than in males by increasing more growth and photosynthetic capacity in females under drought stress.
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臭氧层损耗导致的地球表面UV-B辐射增强以及温室气体增多引起的气候变暖是当今两大全球环境问题。UV-B辐射增强和气候变暖对陆地植物和生态系统产生深远影响,并已成为全球变化研究的重要议题。作为世界第三极的青藏高原,UV-B 辐射增强以及气候变暖现象尤为突出。本试验所在林区是青藏高原东缘的主要林区,具有大面积的亚高山人工针叶成熟林,在全球变化背景下该森林的天然更新潜力如何是急待回答的重要问题。基于此,本研究围绕森林树种的种子和幼苗这一更新的重要阶段,开展了气候变暖、UV-B辐射增强和联合胁迫对云杉种子萌发及幼苗定居影响的研究,旨在全球变化背景下,探讨全球变暖、UV-B 辐射增强和联合胁迫是否对西南地区大面积人工亚高山针叶林更新的种子萌发和幼苗定居阶段产生影响。 本文以青藏高原东缘亚高山针叶林主要树种云杉为研究对象,研究云杉种子萌发及幼苗的生长和生理对UV-B辐射增强与气候变暖的响应。采用UV-B荧光灯(UV-lamp)来模拟增强的UV-B 辐射,此外,采用开顶式有机玻璃罩(OTCs)来模拟气候变暖。本试验包括四个处理:(1)大气UV-B 辐射+大气温度(C);(2)大气UV-B 辐射+模拟气候变暖(W);(3)增强的UV-B辐射+大气温度(U);(4)增强的UV-B辐射+模拟气候变暖(U+W)。 根据本试验结果,UV-B辐射增强对云杉种子萌发没有显著影响,它对萌发云杉幼苗的影响主要体现在幼叶展开以后。根据两年的试验结果,增强的UV-B辐射降低了云杉幼苗抗氧化酶活性,降低了抗氧化物质的含量,此外,造成了膜质的过氧化,表现为MDA在针叶中的积累。增强的UV-B照射处理萌发云杉幼苗两年后,幼苗的生长受到显著抑制。我们的结果显示,OTCs分别提高了空气(10 cm)和土壤(5 cm)温度1.74℃和0.94 ℃。增温显著地促进了云杉种子提前萌发,提高了萌发速率和萌发比率,而且,明显地促进了幼苗的生长,表现为株高和生物量累积的显著增长。此外增温还有利于云杉幼苗根的伸长生长以及生物量的累积,这可以使云杉幼苗更好地利用土壤中的水分和营养元素。 根据本试验结果,温度升高显著地促进了增强UV-B辐射下云杉萌发幼苗的生长,这说明,温度升高缓解了UV-B辐射增强对云杉萌发幼苗的负面影响。这种缓解作用可能是温度升高对UV-B辐射增强处理下幼苗的抗氧化系统活性改善的结果。温度升高还缓解了高UV-B辐射对云杉幼苗根生长的抑制作用,这也可能是增温缓解伤害的原因之一。此外,根据我们的试验结果,增温与UV-B辐射增强联合作用(U+W)下云杉萌发幼苗的生长状况好于大气温度与大气UV-B辐射联合(C)处理,表现为株高、地径、根长和生物量积累均高于C处理,因此可以推断,UV-B辐射增强与气候变暖同时存在对萌发幼苗在两年之内的生长没有产生抑制作用,也就是说,气候变暖的缓解作用完全弥补了UV-B辐射增强的有害作用。 同样,增强的UV-B辐射显著影响了云杉幼苗的光合作用,表现为净光合速率(Pn)和表观量子效率(Φ)的提高,此外,根据我们的试验结果,它还造成了PSII的光抑制。增强的UV-B辐射显著抑制了云杉幼苗对营养元素的吸收,表现为大量营养元素、碳、钙、镁和锌含量的降低,但是,它却显著促进了铁在植株体内的积累。增温显著地提高了净光合速率,但是,它对光系统II(PSII)的光化学效率影响不大。温度升高缓解了UV-B增强对云杉幼苗光合作用的伤害,表现为净光合速率、表观量子效率以及PSII光化学效率的提高。此外,温度升高还缓解了UV-B辐射增强对离子吸收的抑制作用。 Enhanced UV-B radiation due to the reduction of O3 layer and global warming induced by increased greenhouse gases in the air have become the two pressing aspects of global climate changes. Moreover, enhanced UV-B radiation and warming have profound and long-term impacts on terrestrial plants and ecosystems, and the studies focusing on the two factors have attracted many attentions. Qinghai-Tibetan Plateau is the third in elevation in the world, and enhanced UV-B radiation and climate warming are especially prominent in this region. Our research located in the main forest belt in the eastern Qinghai-Tibetan Plateau where large areas of subalpine coniferous forests distributed. Based on that, we carried out a research to study the effects of enhanced UV-B radiation and climate warming on seed germination and seedlings growth of seedlings which are the important basic stage in forest regeneration. This research was arranged by a complete factorial design and included two factors (UV-B radiation and temperature) with two levels. The UV-lamps were used to manipulate the supplemental UV-B radiation and open-top chambers (OTCs) were adopted to increase temperature. The four treatments were: (1) C, ambient UV-B without warming; (2) U, enhanced UV-B without warming; (3) W, ambient UV-B with OTCs warming; (4) U+W, enhanced UV-B with OTCs warming. The main results were exhibited as follows: 1. Based on our results in this research, OTCs increased temperature on average 1.74℃ in air (10 cm above ground) and 0.92 ℃ in soil (5 cm beneath ground). Furthermore, OTCs also slightly reduced soil moisture and relative air humidity, however, the differences was not statistically significant. 2. Our results showed that enhanced UV-B had no significant effects on the seeds germination of P. asperata. Enhanced UV-B affected sprouts of P. asperata until the needles unfolded. During two years, enhanced UV-B inhibited the efficiency of the antioxidant defense systems, and as a result, it induced oxidant stress and the accumulation of MDA in needles. After two years of exposure to enhanced UV-B, the growth of P. asperata sprouts was markedly restrained compared with those under ambient UV-B radiation and temperature (C). Warming significantly stimulated the germination speed and increased the germination rate of P. asperata seeds. In the next place, it prominently facilitated the growth of P. asperata sprouts, represented as improvements in stem elongation and biomass accumulation. Furthermore, warming also increased root growth of P. asperata sprouts, which could made sprouts more efficient to use water and nutrient elements in soil. In this research, warming alleviated the deleterious effects of enhanced UV-B on P. asperata sprouts. It markedly stimulated the growth of P. asperata sprouts exposed to enhanced UV-B. The ease effects of warming on the abilities of the antioxidant defense systems might account for its amending effects on growth. After two years of exposure to enhanced UV-B radiation and warming, the growth of P. asperata sprouts was better than those under ambient UV-B radiation without warming (C), which could be seen from the higher plant height, basal diameter, root length and total biomass accumulation compared with C. 3. Enhanced UV-B radiation significantly influenced the photosynthesis processes of two-year old P. asperata seedlings. Our results showed that enhanced UV-B reduced the net photosynthetic rate (Pn) and the apparent quantum efficiency (Φ), and induced photoinhibition of photosynthetic system II (PSII). Enhanced UV-B significantly decreased the concentration of nitrogen (N), phosphorous (P), potassium (K), calcium (Ca), magnesium (Mg) and zinc (Zn), however, it increased the accumulation of iron (Fe) in the whole plant of P. asperata seedlings. Warming significantly stimulated Pn of P. asperata seedlings but it had no prominent impacts on the photochemical efficiency of PSII. In our research, warming also alleviated the harmful effects of enhanced UV-B on photosynthesis and absorption of ions of P. asperata seedlings. It increased Pn, Φ and the photochemical efficiency of PSII in seedlings exposed to enhanced UV-B. Moreover, warming also increased the absorption of ions of the seedlings exposed to enhanced UV-B radiation.
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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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近年来,随着对作物重茬(连年种植)障碍原因的深入研究,植物的化感作用越来越受到国内外众多学者的重视。而作为重要调料和药用植物的生姜,其连作障碍也备受关注,系统地研究生姜化感作用将有助于理解和最终解决生姜连作障碍问题。本文通过研究生姜不同部位、不同浓度的水浸液对与其间作的两个物种(大豆和四季葱)种子的萌发及幼苗生长的影响,从而证明生姜化感作用的存在;并通过温室盆栽实验研究了生姜的自毒作用(即研究生姜不同部位、不同浓度的水浸液对其幼苗的形态、生理生化、光合作用、土壤酶、土壤微生物多样性及土壤养分的影响),从而揭示生姜退化和衰老的机制,并为生姜筛选出合适的间作物种提供科学依据,对生姜连作障碍提出科学的解决方法。主要研究结果如下: 1. 与对照相比,生姜所有部位(根茎、茎、叶)、所有浓度(10、20、40、 80 g l-1)的水浸液均抑制了大豆种子和葱籽的萌发率、幼苗生长、水分吸收和脂肪酶活性,并且其抑制程度随着水浸液浓度的增加而增强,其生姜各部位水浸液抑制效应的强弱顺序为茎>叶>根茎。这一结果表明生姜根茎、茎、叶含有能够抑制大豆种子和葱籽种子萌发和幼苗生长的水溶性化感物质。根茎是生姜的主要收获部位,而生姜的残株(主要是茎和叶)应该从大田中处理掉以减轻其抑制效应。生姜水浸液中主要化感成分包括:根茎水浸液中主要是丁香酸和伞花内脂;茎水浸液中主要是阿魏酸,且其含量最高为73.4 ug/g;叶水浸液中除了阿魏酸,其他六种物质均检测出来,但含量较高的主要有丁香酸、伞花内脂和香豆酸。 2. 生姜茎和叶不同浓度的水浸液均显著抑制了生姜幼苗的株高、每株叶片数和叶面积,其抑制程度随着水浸液浓度的增加而有所增强,而生姜幼苗每株分枝数差异不显著;同时生姜水浸液也极大程度地影响了生姜幼苗的生物量(包括地下生物量、地上生物量和总生物量,均为鲜重)。在同一浓度下,茎水浸液对生姜幼苗形态指标及生物量指标均显示出最强的抑制作用,叶水浸液次之,根茎水浸液最弱。与对照相比,低浓度的生姜根茎水浸液提高了生姜幼苗叶片内四种抗氧化酶(SOD、POD、CAT、APX)活性,高浓度的根茎水浸液抑制了四种抗氧化酶活性,而茎和叶水浸液均随着浓度的增加而抑制了四种抗氧化酶活性,三种水浸液均随着浓度的增加降低了生姜幼苗叶片内叶绿素的含量,而增加了生姜幼苗叶片的相对电导率和丙二醛含量。同时,三种水浸液均随着浓度的增加降低了生姜幼苗的光合参数(包括胞间CO2浓度、气孔导度、蒸腾速率及净光合速率)。 3. 三种生姜水浸液对所测六种土壤酶活性均产生了不同程度的影响,其中影响最大的是酸性磷酸酶和蔗糖酶,在10 g l-1 时就达到了显著水平,并且所有酶均有随着水浸液浓度增加而增大的趋势;相同部位的水浸液随着浓度的增加,细菌和真菌的数量呈增加趋势,而放线菌的数量呈减少趋势;三种生姜水浸液均随着浓度的增加降低了土壤中有机质的含量,加剧了土壤中硝态氮含量的积累,根茎水浸液对土壤有效磷、速效钾和铵态氮均显示出低浓度提高其含量而高浓度降低其含量的趋势,而茎和叶水浸液则随着浓度的增加均降低了其含量。 4. 与生姜单作相比,所有间作系统均在旺盛生长期和收获期不同程度地提高了土壤酶活性,同时也增加了土壤细菌数量及土壤微生物总数但不显著;所有间作系统在旺盛生长期和收获期均不同程度地影响了土壤真菌及放线菌数量(增加或减少),所有间作系统间的多样性指数差异不显著,除了旺盛生长期四种作物(生姜-大豆-四季葱-大蒜)的间作模式显著降低了多样性指数,其值仅为生姜单作的33.18%;生姜与大豆间作不仅提高了19.6%的生姜产量而且获得了较好的经济效益,并且,所有间作系统均显著抑制了生姜姜瘟病的发生。 5. 不同栽培模式不同程度地影响了收获期生姜的株高、分枝数、根茎产量及内在品质。其中处理2显著地促进了生姜的分枝(10.5%),同时处理2、3和4也促进了生姜的生长(株高分别增加了15.0%、11.4%和14.0%),并且这三个处理提高了生姜的产量;处理2和3能有效提高生姜块茎中维生素C(分别较单作生姜显著提高了3.29%和4.05%)、处理3显著提高了可溶性糖(8.2%)、姜辣素(4.6%)和蛋白质等有益物质的含量,降低硝酸盐有害物质的含量(处理2显著降低了14.0%),改善了姜块的外观和内在品质。并且,生姜与大豆间作具有最高的纯收入和产投比,分别较生姜单作提高了24.80%和8.8%。Recently, allelopathy has been more and more paid attentions by national and foreign scholars with profound research on reasons of crop replanted (continuous planted) obstacle. Ginger rhizome is valuable all over the world either as a spice or herbal medicine and ginger replanted obstacle is also paid attentions. Systematic research on ginger allelopathy will contribute to understanding and ultimate solving problem of ginger replanted obstacle. The effects of ginger aqueous extracts with different parts and concentrations on seed germination and early seedling growth of soybean and chive were studied in this article to testify that ginger existed allelopathy. Furthermore, ginger autotoxicity was also studied by pot experiment in greenhouse (namely research on effects of ginger aqueous extracts with different parts and concentrations on morphological indexes, physiological and biochemical indexes, photosynthesis, soil enzymes, soil microbial diversity and soil nutrients) to reveal mechanism of ginger degeneration and senescence, provide scientific basis for selecting appropriate intercropping species and put forward scientific resolvent for ginger replanted obstacle. The main results were as follows: 1. All aqueous extracts at all concentrations inhibited seed germination, seedling growth, water uptake and lipase activity of soybean and chive compared with the control, and the degree of inhibition increased with the incremental extracts concentration. The degree of toxicity of different ginger plant parts can be classified in order of decreasing inhibition as stem>leaf>rhizome. The results of this study suggested that rhizome, stem and leaf of ginger contained water soluble allelochemicals which could inhibit seed germination and seedling growth of soybean and chive. The rhizome is the main harvested part of ginger. The residue (mainly stems and leaves) of the ginger plant should be removed from the field so as to diminish its inhibitory effect. The main allelopathic components of three kind of aqueous extracts were as follows: Rhizome extract chiefly contained syringic acid and vmbelliferone and stem extract mainly contained frulic acid whose content was the highest (73.4 ug/g). The other six substances were detected except of frulic acid, but only contents of syringic acid, vmbelliferone and p-coumaric acid were higher. 2. Stem and leaf aqueous extracts of ginger with different concentrations significantly inhibited plant height, leaf numbers per plant and leaf area, and the degree of inhibition increased with the incremental extracts concentration. However, tiller number per plant of ginger seedling showed no significant difference. At the same time, ginger aqueous extracts also influenced biomass including under-ground biomass, above-ground biomass and total biomass (fresh weight) to a large extent. Under the same concentration, stem aqueous extract showed the mostly inhibitory effect on morphological indexes and biomass indexes of ginger seedling. Rhizome aqueous extract showed the leastly inhibitory effect and leaf aqueous extract was intervenient. Enhanced concentration of ginger aqueous extracts significantly reduced total chlorophyll content, accompanying with increases in memberane permeability (REL) and lipid peroxidation (MDA). Compared with the control, rhizome ginger aqueous extract of lower concentration (10 g l-1) increased the activities of major antioxidant enzymes (superoxide dismutase, SOD; peroxidase, POD; catalase, CAT; ascorbate peroxidase, APX) of ginger leaf tissue and higher concentration inhibited the activities of four antioxidant enzymes. However, stem and leaf aqueous extract inhibited the activities of four antioxidant enzymes with increase in concentration. Meanwhile, enhanced concentration of ginger aqueous extracts significantly reduced photo-parameters of ginger seedling (including CO2 concentration, stoma conductivity, net photosynthesis rate and transpiration rate). 3. Rhizome, stem and leaf ginger aqueous extract showed different effect on six soil enzyme activities, and acid phosphatase and invertase showed significant effect when aqueous extract concentration got 10 g l-1. Furthermore, six soil enzyme activities increased with increase in aqueous extract concentration. Bcterial and fungi number tended to increase while antinomyces tented to decrease with the increase in aqueous extract concentration of identical part. Ginger aqueous extracts reduced soil organic matter content with increased concentration, accompanying with NO3-—N accumulation in soil. Rhizome aqueous extract showed the same tendency for available P, available K and NH4+—N, namely lower concentration increased their contents in soil and higher concentration reduced their contents. While stem and leaf aqueous extracts reduced their contents with the increamental concentration. 4. All intercropping systems increased soil enzyme activities to different extent both at VGS and at HS compared to solo ginger. All intercropping systems increased the colony numbers of soil bacteria and total of soil microbe but not significantly either at VGS or at HS. All intercropping systems increased the colony numbers of soil fungi and actinomytes to a different extent (increase or decrease) both at VGS and at HS. For DI, difference between all cultivation patterns and S-G was not significant either at VGS or at HS except that G-S-C-G whose value was only 33.18% of S-G at VGS significantly decreased. G-S not only increased ginger yield by 19.6% but also obtained better economic benefit. Furthermore, all intercropping systems significantly inhibited occurrence of bacterial wilt of ginger. 5. Different cultivated pattern influenced plant height, tiller numbers, rhizome yields and intrinsic quality of ginger. Treatment 2 significantly facilitated tiller occurring (10.5%). Treatment 2, 3 and 4 promoted ginger growth (plant height respectively increased 15.0%、11.4% and 14.0%) and enhanced rhizome yields. Treatment 2 and 3 effectively increased vitamin C content (significantly increased 3.29% and 4.05% compared to solo ginger). Treatment 3 significantly increased contents of beneficial substances such as soluble sugar (8.2%), gingerols (4.6%) and protein. Treatment 2 significantly decreased contents of deleterious substance namely nitrate (14.0%) and improved appearance and intrinsic quality of ginger rhizome. Furthermore, treatment 2 (ginger/soybean intercropping) could obtain better economic benefit and showed the highest net income and ratio of benefit and cost whose values respectively increased by 24.80% and 8.8% compared to solo ginger.
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土壤是人类赖以生存的自然环境和农业生产的重要资源,目前土壤受到干旱和盐胁迫的危害越来越严重。杨树具有适应性强、生长快和丰产等特性,本论文以青杨组杨树为模式植物,研究杨树对土壤干旱和盐胁迫的生态生理及蛋白质组学反应,研究成果可为我国干旱半干旱地区营造人工林、防止沙漠化提供理论依据,也为恢复与重建盐污染地区退化生态系统提供科学指导。主要研究结果如下: 1 青杨不同种对逐步干旱胁迫的响应差异 将来自喜马拉雅山东缘高海拔的康定杨和低海拔的青杨枝条扦插在温室中,用来检测它们对逐步干旱胁迫的响应。研究结果表明来自不同海拔的杨树对逐步干旱胁迫的适应性反应是不一样的。株高、叶片发育、叶片相对含水量、丙二醛、过氧化氢等指标的显著性变化在青杨中比在康定杨中来得早些,而且随着干旱胁迫程度的增加,这些参数的变化越来越明显,尤其是当青杨受到严重干旱胁迫的时候;而可溶性蛋白、可溶性糖、游离脯氨酸、抗氧化酶活力变化在康定杨中来得早一些。与青杨相比,在干旱胁迫下,康定杨仍能保持较好的植株生长和叶片发育;康定杨也能在逐步干旱条件下积累更多的可溶性蛋白、可溶性糖、游离脯氨酸及抗氧化酶活力,但是在丙二醛和过氧化氢含量方面增加的更少些。而且,我们的研究结果表明高海拔的康定杨有更强的耐干旱能力,杨树对干旱胁迫的适应能力与干旱发生的速度、强度、持续时间及两种杨树的海拔有关。 2 干旱胁迫下青杨不同种的蛋白质组学分析 来自青杨和康定杨雌株的枝条扦插在温室中,用来研究它们对干旱胁迫的蛋白质组学反应。采用TCA-丙酮/酚提取法提取总蛋白,并进行双向电泳分析。在每个处理的重复图像中都能检测到1,000 个以上的蛋白点。在青杨中有58 个蛋白在干旱处理后发生显著变化,其中22 个蛋白通过肽指纹图谱成功鉴定。康定杨中有69 个蛋白的表达量发生了显著变化,其中有25 个蛋白通过肽指纹图谱成功鉴定。这些被鉴定的蛋白主要参与了光合作用、氧化还原平衡、信号传导、能量代谢、蛋白质合成等过程。尽管被鉴定的蛋白只占叶片总蛋白的很少一部分,但这些被鉴定的干旱响应蛋白可能对维持植株内部平衡方面有重要作用。 3 青杨的盐胁迫响应 青杨植株分别用 0、50 和100 mM NaCl 溶液进行处理。叶片相对含水量、叶绿素a、b 含量、CO2 同化速率和气孔导度的降低表明叶绿体受到了盐胁迫的影响。过氧化氢、丙二醛含量及电导率的升高表明细胞受到了伤害。可溶性糖、游离脯氨酸含量及抗氧化酶含量的上升增加了植株耐盐胁迫的能力。在每个处理的重复图像中都能检测到1,000 个以上的蛋白点。其中有38 个盐响应蛋白被成功鉴定,有16 个蛋白(点4、10、11、14、15、21、24、26、27、28、33、34、35、36、37 和38)出现在盐胁迫的植株中;3 个蛋白(点10、11 和35)只出现在重度盐胁迫处理中;而1 个蛋白(点1)只出现在对照处理中。2 个蛋白(点1 和2)表达量下降,其余蛋白点表达量都增加。被鉴定的蛋白一部分参与了生理生化反应,而另一部分则在信号传导、蛋白质合成等方面有重要作用。盐胁迫下的生理生化变化及蛋白质组学的联合研究有利于青杨对盐胁迫的适应性分析。 Soil is the indispensable environment for human survival and important resource for agriculture development. Nowadays soil is threatened by drought stress and salt stress. Poplars (Populus spp.) possess some characters such as strong acclimilation, fast growth and great production of biomass. In this study, different species of Populus section Tacamahaca spach were used as model plants to investigate the ecophysiological and proteomic responses to drought stress and salt stress. Our results can provide theoretical evidence for the afforestation and prevention of desertification in the arid and semi-arid areas, and also can supply scientific direction for the reconstruction and rehalibitation of ecosystems contaminated by salinity. The results are as follows: 1 Adaptive responses to progressive drought stress in two contrasting poplar species originating from different altitudes Cuttings of Populus kangdingensis C. Wang et Tung and Populus cathayana Rehd., originating from high and low altitudes in the eastern Himalaya, respectively, were examined during one growing season in a greenhouse to determine the effects of progressive drought stress. The results manifested that the adaptive responses to progressive drought stress were different in these two species from different altitudes. Significant changes in height increment, leaf development, relative water content (RWC), malondialdehyde (MDA) and hydrogen peroxide (H2O2) appeared earlier in P. cathayana than in P. kangdingensis, whereas changes in soluble protein, soluble sugar, free proline and antioxidant enzymes appeared earlier in P. kangdingensis. In addition, changes in these parameters became more and more significant when the drought stress progressed, especially under severe drought stress in P. cathayana. Compared with P. cathayana, P. kangdingensis was able to maintain a superior height increase and leaf development under drought stress. Also, P. kangdingensis possessed greater increments in soluble protein, soluble sugar, free proline and antioxidant enzymes, but lower increments in MDA and H2O2 than did P. cathayana when the cuttings were exposed to progressive drought stress. Our results suggest that P. kangdingensis originating from the high altitude has a better drought tolerance than does P. cathayana originating from the low altitude. Furthermore, this study manifested that acclimation to drought stress are related the rapidity, severity, duration of the drought event and the altitude of two contrasting species. 2 Proteomic responses to drought stress in two contrasting poplar species originating from different altitudes The cuttings from a female clone of P. kangdingensis and P. cathayana were used to determine proteomic response to drought stress, respectively. Total proteins of the leaves were extracted by a combination of TCA-acetone and phenol, and separated by two-dimensional gel electrophoresis. More than 1,000 protein spots were reproducibly detected on each gel. 58 differentially expressed spots were detected under drought stress in P. cathayana and 22 drought-responsive proteins were identified by peptide mass fingerprint. 69 differentially expressed spots were detected under drought stress in P. kangdingensiss and 25 drought-responsive proteins were identified by peptide mass fingerprint. The identified proteins are involved in several processes, i.e., signal transduction, protein processing, redox homeostasis, CO2 fixation and energy metabolism. Although the proteins identified in this investigation represent only a very small part of the poplar leaf proteins, some of the novel drought-responsive proteins identified here may be involved in the establishment of homeostasis in response to drought stress in the woody plants. 3 Responses to salt stress in P. cathayana Cuttings from a female clone of P. cathayana were treated by Hoagland’s solution: 0, 50, 100 mM NaCl, respectively. Salinity significantly decreased the relative water content of leaves, the contents of chlorophyll a and chlorophyll b, CO2 assimilation rate (A) and stomatal conductance (gs) in both salt stress treatments,which suggested the chloroplast was affected by salt stress. The observed increases of H2O2 and malondialdehyde contents and electrolyte leakage suggested that salinity caused cellular damage, whereas the increases in compatible solutes and in the activities of antioxidant enzymes enhanced the salt tolerance. More than 1,000 protein spots were reproducibly detected on each gel, and 38 salt-responsive proteins were successfully identified by peptide mass fingerprint (PMF). 16 spots (spot 4, 10, 11, 14, 15, 21, 24, 26, 27, 28, 33, 34, 35, 36, 37 and 38) absent in the control sample were induced by the salt treatment, and three spots (spot 10,11 and 35) were present only in the severely salt-stressed treatment. The %vol of the differentially expressed proteins generally increased with progressing salt stress, except for the decreased %vol of two proteins (spot 1 and 2) under salt stress and the presence of spot 1 only in the control sample. Some of the novel salt-responsive proteins identified here may be involved in physiological, biochemical response to salt stress in P. cathayana, the other identified proteins play a role in numerous cellular functions, including signal transduction and protein processing. An integrated physiological, biochemical and proteomic approach was used here to systematically investigate salt acclimation in poplar.
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人类活动引起全球大气中温室气体(CO2、CH4、NOx)浓度不断增加,致使地球表面温度在过去的100 年中已经增长了0.74 ± 0.18℃,预计到本世纪末将会增加1.1-6.4℃。此外,氮沉降也是当今社会的重要环境问题,随着经济发展的全球化, 高氮沉降也呈现出全球化趋势。全球气候变暖和氮沉降给陆地生态系统的地上、地下生物学和生物地球化学过程所带来巨大影响越来越引起人们的关注。 本文以川西亚高山针叶林的两个重要树种云杉和油松幼苗为研究对象,采用红外辐射增温(空气增温2.1℃,土壤增温2.6℃)和根部施氮(施氮量25 g N m-2yr-1)的方法,从生长形态、光合作用、抗氧化能力和矿质营养等方面研究这两种幼苗对气候变暖和氮沉降的响应。该实验为室外控制实验,包括四个处理:(1)不增温+不施氮(UU);(2) 不增温+施氮(UF);(3) 增温+不施氮(WU);(4) 增温+施氮(WF)。本研究旨在从生理生化、物质代谢 、生长及形态等不同水平上研究模拟增温和施氮对两种树苗的联合效应,提高我们对全球变化下亚高山针叶林早期更新过程的理解,同时也为森林管理提供科学依据。具体研究结果如下: 单独增温处理显著提高了云杉和油松幼苗的地茎、叶重、茎重、根重以及总生物量;单独施氮处理也增加了两种幼苗的株高和总生物量。而增温和施氮联合作用对两种幼苗生长的影响并不相同,联合作用对云杉幼苗生长指标的正效应显著低于单独施氮处理,但是联合作用比单独增温或施氮更大程度的促进了油松幼苗生物量的积累。 单独增温和施氮都有利于提高云杉和油松叶片中叶绿素含量、净光合速率(A)、最大净光合速率(Amax)、表观量子效率(Φ)、最大光能转化效率(Fv/Fm)和量子产量(Y)。与对两种幼苗生长指标的影响相似,加氮和增温共同作用下油松幼苗的以上光合指标比在单独增温或施氮处理下有更大程度的提高;而联合作用下云杉幼苗叶绿素含量、净光合速率、最大净光合速率、表观量子效率、最大光能转化效率以及量子产量比单独施氮处理明显地降低。 增温和施氮都显著地降低了云杉和油松幼苗针叶组织中活性氧和丙二醛的积累。交互作用降低了云杉幼苗叶片的抗氧化酶活性、脯氨酸和ASA 的含量,却显著提高了油松幼苗SOD、POD、APX 等抗氧化酶的活性,并且对油松幼苗脯氨酸和ASA 积累的促进作用比单一因子更加明显。因此,增温和施氮共同作用下油松幼苗叶片中O2-产生速率、H2O2 及MDA 含量明显降低,而云杉叶片中只有O2-产生速率出现降低趋势。 增温和施氮都降低了云杉体内的P、Ca、Mg 元素的含量,增加了Cu、Zn、Mn 在各器官内的积累。对油松幼苗而言,增温和加氮单独作用也显著降低了Ca 含量增加了Cu、Zn、Mn 的积累,但是不同于云杉幼苗的是P、Mg 也显著增加。增温和施氮联合作用对云杉幼苗体内元素的影响与单一施氮处理或增温处理相似,不同的是比单一因子作用更为明显降低了P、Ca、Mg 含量,增加了植株中N、Cu、Zn、Mn 的含量,但是油松矿质元素含量在联合作用下并没有产生类似于云杉幼苗的双因子叠加效应。 总之,尽管单独增温或者施氮都有利于云杉和油松幼苗生长指标、光合能力以及抗氧化能力的提高。但是,增温和施氮对云杉幼苗生长生理的促进效应非但没有在交互作用下有更大的提高,反而低于单独氮处理。与此不同的是,增温和施氮联合作用比单因子作用更有利于油松幼苗生长及生理指标的提高。 With the continued increase in atmospheric concentrations of greenhouse gases (CO2、CH4、NOx), the mean global surface temperature has increased by about 0.74 ± 0.18℃ over the past century and is predicted to rise by as much as 6.4℃ during this century. Besides global warming, nitrogen deposition is another serious environmental problem caused by human activities, and high nitrogen load has become globalization as a result of global economy development. Global climate warming and nitrogen deposition have induced dramatic alternations in above - and below- ground biology and biogeochemistry process in terrestrial ecosystems, and more and more attention has been invited to those problems. This experiment mainly studies two important species Picea asperata and Pinus tabulaeformis in subalpine coniferous forest of western Sichuan, China. Infared heaters are induced to increase both air and soil temperature by 2.1℃ and 2.6 ℃, respectively. Ammonium nitrate solution (for a total equivalent to 25 g N m-2 year-1) is added to soil surface. There are four treatments in this study: (1) unwarmed unfertilized (UU); (2) unwarmed fertilized (UF); (3) warmed unfertilized (WU); (4) warmed fertilized (WF). This study is conducted to determine the influences of experimental warming and nitrogen fertilization on physiolchemistry, nutrition metabolism, growth and morphology in the two coniferous species seedlings. The current study is favorable for increasing our understanding on the early phase of regeneration behavior in subalpine coniferous forest, and it also provide scientific direction for forest management under future global changes. The results are as follows: Artificial warming alone significantly increased basal diameter, leaf mass, stem mass, root mass and total biomass for Picea asperata and Pinus tabulaeformis seedlings, and single nitrogen fertilization are also favorable for growth of the two species and stimulate plant hight and total biomass. The two species seedlings respond differently to the combination of elevated temperature and nitrogen addition. Warming combined with nitrogen fertilization weakens the positive effects of nitrogen addition for growth of Picea asperata seedlings. However, the combination of elevated temperature and nitrogen fertilization further increase biomass accumulation of Pinus tabulaeformis seedlings. Both elevated temperature alone and nitrogen fertilization alone can increase photosynthetic pigments contents, net photosynthetic rate (A), maximum net photosynthetic rate (Amax), apparent quantity yield (Φ), maximum photochemical efficiency of photosystem II (Fv/Fm) and effective quantum yield (Y). Similarly with growth parameters, the combination of warming and nitrogen addition induced more increment of these above photosynthetic parameters for Pinus tabulaeformis seedlings. However, these photosynthetic parameters of Picea asperata seedlings under the combination of warming and nitrogen addition are lower than those under nitrogen fertilization alone. The levels of active oxygen species (AOS) and malodiadehyde (MDA) in needles of the two coniferous species seedling are obviously decreased by experimental warming or additional nitrogen. Warming combined with nitrogen fertilizer reduces the activities of SOD, CAT and APX, and the contents of proline and ASA of Picea asperata seedlings, but the combination significantly increases activities of these antioxidant enzymes in needlels of Pinus tabulaeformis seedlings and further improves the accumulation of proline and ASA compared to either artificial warming or nitrogen addition. Therefore, the rate of O2 - production, the contents of H2O2 and MDA in needles of Pinus tabulaeformis seedlings are remarkably reduced by the combination of warming and nitrogen addition, but the combination only significantly decreased the rate of O2 - production of Picea asperata seedlings. Elevated temperature or nitrogen fertilization decrease the contents of P, Ca, Mg but increase Cu, Zn, Mn contents for Picea asperata seedlings. For Pinus tabulaeformis seedlings, elevated temperature alone and nitrogen fertilization alone decreased Ca, but increased P, Mg, Cu, Zn, Mn contents. The effects of the combination of warming and nitrogen addition on these element contents in needles of Picea asperata seedlings are added or multiplied the effects of warming and nitrogen addition alone, resulting in less contens of P, Ca, Mg and more contents of Cu, Zn, Mn than either elevated temperature or nitrogen fertilization. Howere, these adding or multipluing single-factor effects on contents of these elements are not observed in the case of Pinus tabulaeformis seedlings. In conclusion, growth parameters, photosynthetic capacities and antioxidant abilities of Picea tasperata and Pinus abulaeformis seedlings are improved by experimental warming or nitrogen fertilization. Interestingly, the positive effects of warming and nitrogen addition on growth and physiological performances are not multiplied by the combination of elevated temperature and nitrogen fertilization, even dempened for Picea asperata seedlings. However, for Pinus tabulaeformis seedlings, growth and physiological performances are further improved by the combination.
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SPAC系统中水总是从水势高处流向水势低处,土壤水势的高低是影响植物根系吸水速率的主要因素。相对于土壤含水量来说,用土壤水势作作物非充分灌溉的下限指标更具科学性,更利于水分下限指标的推广应用。试验结果表明,对于整个生育期来说,-200 kPa是比较适宜的大棚番茄土壤水势下限。在这一指标的推广过程中,须先测定当地土壤水分特征曲线,并从曲线上找出该适宜水势对应的土壤含水量,通过控制土壤含水量达到控制土壤水势的目的,从而实现精量灌溉。
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在旱棚控制条件下 ,用五因素五水平二次回归正交旋转组合设计 ,研究了水肥施用分配 ,补水量 ,施N量 ,施K量 ,有机肥施用量的综合作用效应 .结果表明 ,五因素对肥料N利用率的作用顺序为施N量 >补水量>水肥施用分配 >有机肥施用量 >施K量 .水肥施用分配相对于不同水、化肥 (N、K )、有机肥量的选择 ,可以大大地提高肥料N利用率 ;量少应重前施用 ,量多重后施或均施为好 .补水量与施N量的耦合规律为 :高水高肥效果最佳 ,肥料N利用率可达 5 7 83 % ,中水中N及低水低N次之 ,为 2 9 17%~ 40 99% ;高水低N或低水高N导致最低的利用率 ,为 2 2 87%或 2 2 5 1%
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延安日光温室蔬菜生产从初步发展走向稳定健康发展 ,基础设施建设、栽培技术等取得了一定成绩 ,但技术力量仍感不足 ,农药、化肥污染愈加严重。提出今后进一步发展的对策
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揭示黄瓜产生连作障碍的机理,为设施蔬菜的可持续发展和减轻连作障碍提供参考依据。【方法】在日光温室内进行田间试验,以露地玉米地土壤为对照,研究不同连作年限(4,5,8和12年)下黄瓜产量和品质及土壤酶活性的变化。【结果】随着黄瓜连作年限的增加,黄瓜产量、可溶性固形物和维生素C含量均下降,硝酸盐含量上升,土壤呼吸强度降低,土壤脲酶活性、蔗糖酶活性、碱性磷酸酶活性均呈先上升后降低的趋势;随着黄瓜生长季节的变化,土壤脲酶活性呈先上升后降低再升高的趋势,土壤蔗糖酶活性、碱性磷酸酶活性和土壤呼吸强度均呈先上升后降低的趋势。【结论】随着连作年限的延长,设施黄瓜的产量和品质均下降,土壤脲酶、蔗糖酶和碱性磷酸酶活性均呈先升高后降低的趋势。
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大气温室效应气体N2O、CO2增多与全球气温变暖有着密切的关系,由于农业活动导致的碳排放量占碳总排放量的25%,因此研究农田土壤有机碳的影响因素,对增加农田碳素固定和保持,减少由于不合理的土地使用而导致大量CO2的排放,维持农业和生物圈生态系统的可持续发展有着重要意义。本文分析了温度、水分、土地开垦、休闲和撩荒、耕翻、轮作、秸秆还田、肥料管理等对土壤有机碳的影响。减少翻耕次数,增加秸秆还田,优化氮、磷、钾等养分用量及配比,是提高农田,尤其是旱地农田土壤有机碳含量,培肥、改良土壤的重要途径。
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对陕北黄土高原日光温室大棚蔬菜黄瓜、大青菜、油麦菜和西芹进行了叶面喷施富万钾有机钾肥的田间试验研究。结果表明,在日光温室大棚蔬菜上施用有机钾肥,黄瓜植株高度、叶片数、茎粗、叶面积、瓜长均增加。大青菜的茎粗增长44%。油麦菜的株高、茎粗、地上植株、地下根重、总生物量均明显增加。西芹的株高、茎粗、地上植株、地下根重以及总生物量增加。富万钾有机钾肥能明显促进蔬菜生长,提高产量。
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采用田间实验研究了施用氮肥对设施条件下大青菜生长发育和产量,以及对土壤硝酸盐含量累积分布的影响。结果表明,在施氮肥0~500 kg hm-2范围内,氮肥施用量越大,产量越高,但到达一定量时,便不能再促进产量的提高,反而会引起减产。土壤铵态氮含量随着土壤剖面向下不断下降,表层的铵态氮含量较高,土壤硝态氮含量在整个土壤剖面的分布变化较小,但也是从表层向下处于一种缓慢减少的趋势,铵态氮的累积主要集中在上部土壤剖面,硝态氮的累积分布在整个土壤剖面,土壤硝态氮含量随施肥量增加而增加;氮肥施用量对硝酸盐含量有很大影响。在施氮肥0~700kg hm-2范围内,氮肥施用量越多,大青菜硝酸盐含量越高。
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To explore the changes of soil chemical properties in vegetable greenhouse,a comparative study was carried out with the samples gathered from vegetable greenhouse fields and their adjacent upland fields in Damintun Town,Xinming County,Liaoning Province.The results showed that compared with upland fields,the contents of soil organic carbon and total nitrogen in greenhouse fields increased significantly.At the depth of 0~30 cm,soil organic carbon in greenhouses of 1-,4-and 10-year increased by 31.09%,35.44%,and 66.80%,respectively,compared with the upland soil.Soil nitrate content at the depth of 0~30 cm in greenhouse fields was 5.05~12.49 times as much as that in upland fields.The nitrate content in different soil layers increased with the increasing age of greenhouse field.,e.g.,at the depth of 20~30 cm,soil nitrate content was significantly higher in 10-year than in 1-and 4-year greenhouse field,with an increase of 65.73% and 50.89%,respectively,and 6.55 times as much as that in upland field,which indicated that soil nitrate transported downwards,and obviously enriched in deeper soil layers under heavy application of fertilizer.Also with the increasing age of greenhouse field,soil pH decreased,while soil soluble salts accumulated.