129 resultados para Water use efficiency(WUE)
Resumo:
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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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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以栽培一粒(2n)、栽培二粒(4n)和长武134(6n)为材料在大型活动防雨棚条件下研究不同倍性小麦材料在不同密度和水分条件下的产量适应性变化。结果发现,在两种水分条件下随着染色体倍性从2n→6n的增加,产量、千粒重、水分利用效率(WUE)和收获指数均呈增加趋势,在水分胁迫下各材料穗粒数和穗数则呈降低趋势,而在正常供水下穗粒数则呈增加趋势。在水分胁迫下栽培一粒、栽培二粒和长武134最高产量分别出现在中、低、高密度群体,而同一材料不同密度群体间变异系数分别为6.73%,1.98%,9.07%;不同倍性材料千粒重均随着密度增加而减小,而穗数则逐渐增加,二倍体的穗粒数以中密度最高,四倍体的穗粒数随着群体密度的增加而减小,六倍体则相反;三种材料WUE和收获指数分别以低、高、低密度最高。正常供水下随着染色体倍性从2n→6n的增加,三个倍性材料最高产量分别出现在高、低、低密度群体,而同一材料不同密度群体间变异系数分别为6.01%,17.12%,2.46%;千粒重表现为中密度>低密度>高密度,而穗粒数均以低密度群体最高,二倍体和四倍体以高密度群体最低,六倍体则以中密度群体最低,穗数则随着密度群体增加而增加;二倍体WUE以高...
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研究了黄土丘陵区引种草种柳枝稷 ( Panicum virgatum)的光合生理生态特性 ,比较了不同叶位叶片光合速率 ( Pn)、蒸腾速率 ( Tr)、水分利用效率 ( WUE)的日变化以及环境因子的作用。结果表明 ,柳枝稷叶片 Pn日变化曲线为双峰型 ,中午“光合降低”主要是由于叶温过高导致呼吸高引起的净光合速率降低。叶龄增大 ,叶片 Pn日变化相对较平缓 ,其中壮龄叶Pn日变化最为平缓。幼龄叶 Tr的日变化为双峰型 ,随叶龄增大 (叶位下降 )而成为单峰型。WUE的日变化可划分为上午的降低和下午的波动 2个阶段 ,最上充分展开叶 (旗叶 )的WUE始终最高。
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National Natural Science Foundation of China [70673097]
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本文研究了北京山区落叶阀叶林优势种一辽东栎、大叶白蜡、北京丁香、核桃楸、山杏和荆条等乔灌木的稳定碳同位素比率,长期水分利用效率、植物蒸腾特性和土壤植物体的水分运动特点,并从植物解剖学的角度研究了这些植物叶片特点和其水分生理生态特性的关系。 对北京山区落叶阔叶林生态系统这几种乔、灌木植物叶片中的碳稳定性同位素比率(δ¹³C值)测定结果显示,这些植物叶片的δ¹³C值受多种因子的影响,具有较大的种间差异及时空异质性。主要表现在不同植物种叶片δ¹³C值不同,其排列顺序为山杏(- 24.75±0.85%。>大叶白蜡(- 25,94±1.52%。)=荆条(- 26.01±1.63%。)=辽东栎(一26.07±1.17%。)=北京丁香(-26.46±0.80c70。)>核桃楸(-28.11±1.52%。);生长初期叶片δ¹³C值较生长末期高,尤以核桃楸和辽东栎表现明显其生长初期和末期的叶片δ¹³C值皆相差达3‰;生境条件,特别是土壤水分含量和土层厚度,对植物叶片的813C值的高低有较大的影响,生长在于旱生境中的植物具有较高的δ¹³C值。另外,即使是同一株植物,叶片δ¹³C值也因其在冠层中所处的位置不同而异,冠层项部叶片的δ¹³C值高于林冠内部的叶片。北京山区落叶阔叶林优势种的长期水分利用效率与种的特点有关,山杏最高(4.950±0.l71mmolC0_2•mol-1H_2O),核桃楸最低(3.760±0.203mmolC0_2•mol-1H_2O),大叶白蜡、荆条、辽东栎和北京丁香居中(4.346- 4.530 mmolCO_2•mol-1H_2O),大部分植物长期水分利用效率在春季(5月)较高,秋季较低,荆条由于物候期的特殊性在其生长季初期较低,而后逐渐增高。核桃楸在不存在水分亏缺情况下,树干液流速率受微气候因子的影响,液流速率的最大值达1600g•hour 左右。树干液流速率的日进程和大气相对湿度、温度的日进程具有相当好的生态学同步性。通过对核桃楸夜晚树干液流的分析可以得出其有根压存在的结论。 植物叶片和枝条中自由水和束缚水含量主要决定于植物种的特性,枝条的年龄、生境特点,特别是土壤水分特点。在落叶阔叶林I(样地2)中植物叶片自由水含量的排列顺序是:北京丁香>核桃楸>大叶白蜡>辽东栎;而杂灌丛(样地1)中植物叶片自由水含量的排列顺序是:核桃楸>大叶白蜡>山杏>荆条>北京丁香>辽东栎,可见群落类型对植物自由水含量影响是很大的,植物束缚水含量与其自由水含量的格局完全相反,荆条、山杏等植物含量高,核桃揪含量低。枝条水分含量有与叶片水分含量相类似的特点。 北京山区落叶阔叶林优势种的水分生理生态学特性和其叶片的特点有很大的关系,首先是植物叶片的特点总是和其种的特性相联系,主要表现在叶的类型、叶片上毛、气孔密度、着生方式等,如荆条叶片上下表面都密被披针形毛,气孔小,核桃楸气孔较大且凸出,大叶白蜡叶片上的气孔凹陷,辽东栎的气孔呈椭圆状,保卫细胞上有许多白色蜡质结晶。有一些种有环状的气孔外缘。生境的变化对叶片的形态特征有影响,在全光照条件下叶片小而犀,而在庇荫条件下叶片大而薄,在扫描电镜下可见全光照条件下北京丁香叶片基本无毛,庇荫条件下则有短微毛,全光条件下荆条叶片上毛有小乳头状凸起,庇荫条件下没有。本文所研究的植物种气孔都着生在叶片的下表面,气孔密度的大小排列顺序是:辽东栎>山杏>北京丁香>核桃楸>大叶白蜡。经方差分析显示种闷气孔密度存在极显著性差异。对所研究植物的气孔导度和环境因子、叶片解剖特点进行线性回归分析,得到了总体的和各个种的回归模型,结果表明光照强度、气孔密度等对气孔导度影响显著,但因种的不同相互之间存在差别。 植物的蒸腾速率受多因子的影响,主要有种的特点、微气候因子(光照强度、大气相对湿度、叶面温度、叶室温度等)和土壤水分特点。植物的蒸腾速率日进程和微气候因子日进程有相当好的生态学同步性。对辽东栎的蒸腾速率和光照强度的研究发现二者有很好的线性关系。这些植物蒸腾速率都表现出一定规律的日进程和季节进程,大多数植物蒸腾速率在一天中有数次波动,最高峰一般出现在中午12时之前;在整个生长季中,6月底至8月初的蒸腾速率高于其他月份。
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鄂尔多斯高原是一个多层次、复杂的生态地理过渡带,具有复杂多样的环境条件和生态特点。因而,研究该地区植被与环境复杂多变的相互关系有助于我们认识鄂尔多斯高原生态系统的脆弱性和植被对于维持该区生态系统结构和功能过程的重要性。 降水、温度及其组合特点决定了鄂尔多斯高原特殊的生态环境格局,分析其演变趋势有助于理解鄂尔多斯高原生态环境的演变过程。从旬、月、生长季和非生长季及年四个不同的时间分辨率对鄂尔多斯高原八个气象台站近30年的气象资料进行分析。结果表明30年来年平均气温、2、9、12月均温明显增高;年降水未发生显著变化,降水分配模式变化明显。采用与植物生长密切相关的气候因子对整个鄂尔多斯高原的气候特征进行综合分析,将鄂尔多斯高原划分为三个综合气候类型,即:半湿润、低蒸发型,半干旱、半湿润、中等蒸发型和偏干旱、高蒸发型。根据不同的气候类型,确定适宜恢复的植被类型,对目前该区域进行的生态恢复工作具有重要指导意义。 在鄂尔多斯高原从东向西的降水梯度上,选取三个实验点测定柠条的光合、蒸腾和水分利用效率等主要生理生态指标,分析了同种植物对不同环境的反应。鄂托克旗柠条光合速率最高,东胜东部柠条次之,杭锦旗柠条光合速率最低而蒸腾速率最大,东胜东部柠条和鄂托克旗柠条的蒸腾速率相差不大。 植被能够降低其下垫面及其附近的地表温度,从而影响地表蒸发。以鄂尔多斯高原典型植物油蒿和柠条作为研究对象,采用先进的非接触红外测温法,并提出度量植株温度效应的影响指数,对鄂尔多斯高原两种典型植物植株附近地表温度进行了比较分析,结果表明,柠条对地表温度的影响较油蒿明显。
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陆地样带是国际地圈——生物圈计划(IGBP)研究中最引入注目的创新之一。目前,国际上已经设立了15条陆地样带,研究内容涉及环境梯度分析、气候变化对植被初级生产力的影响及环境变化、土地利用等与植被变化的对应关系等。沿该陆地样带分布较广的关键种生理适应性等方面对影响其生理功能形态结构的研究较少,特别是茎、叶等组织功能研究较少。 中国东北样带(NECT)是全球陆地样带的重要组成部分,多年来已开展了大量深入系统的研究工作,已成为我国生态学、地学等学科的重要研究平台。本研究以中国东北样带中西段广泛分布的重要关键种——羊草(Leymus chinensis)为研究对象,分析了羊草茎、叶显微结构的生态可塑性及其与水分利用效率的关系,进而阐述了羊草适应不同生境条件,特别是适应水分变化的机制,为揭示羊草及其种群、群落乃至以羊草为优势种或建群种的草地生态系统在全球变化背景下的发展趋势提供理论依据。 基于2001年7~8月第3次中国东北样带考察资料,采用高精度Olympus显微镜及C同位素分析技术(δ13C判别值),结合在野外取样过程中测定的样地土壤含水量和海拔高度,以及近十年各样地年降水量和年均温度气象资料,分析了羊草茎、叶显微结构和水分利用变化与环境因子的关系,以及以羊草为建群种或共建种的无牧和放牧样地群落生物量、物种多样性和植物功能型组成变化与环境因子的关系。 结果表明:羊草叶片表面及内部主要显微结构特征参数各样地间有不同程度的差异,其中气孔密度与降水量呈线性正相关。代表气孔开张程度的气孔长度和宽度变化与土壤含水量呈线性相关。叶表面角质层厚度与海拔高度变化关系较大,并以上表面角质层厚度变化最为明显,主要受海拔高度升高引起的紫外线照射增强的影响。运动细胞带宽度占叶面积比虽然与各环境因子关系不很密切,但温度变化的影响较突出,这一显微结构调整与气孔变化构成干旱——高温调节机制。叶片表面毛茸的变化也是非常显著的,但与各环境因子关系密切程度均不大,可以肯定的是在土壤水分状况较好的生境下羊草叶片表面毛茸密度及长度明显增加,而一些干旱生境中常表现为毛茸较少、较短,个别样地基本没有发育较好的毛茸。总体上看,羊草叶片对干旱化的形态结构调整以气孔密度和开张程度的变化最大,是羊草叶片调节水分利用效率的重要适应性生态可塑性调整。 与叶片相比,羊草茎横切面结构特征的变化与各环境因子关系的显著性不是很强,但各样地间的差异是比较显著的,许多结构调整可能与土壤养分条件的变化有一定关系,如茎秆粗度变化、基本薄壁组织厚度和中央空腔(髓腔)直径的变化等,但本研究未能涉及这方面内容,有待于进一步研究。 羊草水分利用效率与降水量和土壤含水量呈显著的负相关关系,即随降水量和土壤含水量增大羊草水分利用效率明显降低,蒸腾耗水增大,这一生理变化与显微结构的调整关系密切,特别是气孔密度与气孔宽度在水分较差生境中明显减小,从而有利于适应干旱环境,减少耗水量。表现比较突出的是非地带性林西样地,其降水量处于10个样地的中等偏低水平,但其δ13C判别值较低,达-26.063‰,与降水量较大的长岭、双辽样地几乎相当,并比相邻的林东和克旗样地明显低,其气孔密度、开张程度及叶脉后生导管直径均较高(大),但其土壤水分状况是最好的样地之一,尽管取样时不幸遇到雨天,但从其群落类型——羊草杂类草草甸,并伴生许多中、湿生种类上看,其生境的湿润程度是毋庸置疑的。这一非地带性样地中羊草结构的变化从另一侧面反映了羊草显微结构调整对水分环境的适应。δ13C判别值是一个非常敏感的参数,在分析植物水分利用效率及其相关领域的研究中应深入利用。 群落植物功能群组成与环境因子及群落初级生产力关系研究结果表明,丛生禾草生长型功能群、旱生和中旱生植物水分生态类型功能群具有明显的地带性变化规律,并与群落生物量变化关系密切,变异性较低,占群落生物量比例较大,可考虑作为植物功能型组合对无牧样地植被变化进行评估和预测。在放牧影响下,C4植物光合类型功能群呈现明显的地带性变化,并在群落中所起的作用明显增强,亦可考虑作为评估和预测植被变化的植物功能型组合。无牧样地与放牧样地研究结果均表明,按Raunkiaer划分的地面芽、地下芽、地上芽和一年生植物生活型功能群,其地带性变化不明显,或变异率高,或占群落生物量比例小,不宜作植物功能群组合对植被变化进行评估和预测。
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随着三峡大坝建设,在2003年6月三峡库区蓄水到135 m水位,之后人为调节使其在137-139 m范围内波动变化。从2003年7月开始,我们对库区植物的水分关系及其对三峡水位上升的可能反应进行了系统研究。 在库区中残存的次生松栎混交林,我们从江边沿海拔梯度设置了3块样地:Riparian,Mid-hill和Top-hill样地,垂直高度相差约20 m。从2003年7月到2004年7月,我们比较了岸边样地内与高处两样地内植物的木质部水分稳定同位素D和18O值,植物清晨和中午水势,叶片碳稳定同位素值13C,以及2004年7月测定的气体交换。岸边 植物木质部水分同位素值显著高于江水的同位素值,而与高处两样地内植物木质部水分同位素相近。岸边植物与高 处植物具有相近的清晨水势和中午水势,也表明对岸边植物来说,江水并不是它们重要的水分来源。同样,岸边植 物叶片 13C值与高处同种植物的值也不存在显著差异。我们研究的3种植物清晨水势都与土壤含水量正相关,尤 其浅层土壤更为显著。研究结果表明岸边植物几乎没有利用江水,而同高处两样地内植物一样,都是以利用渗入到土壤中的降雨为主。 松栎混交林中,马尾松与槲栎相比,净光合速率和气孔导度,叶片含N量,以及清晨水势低于槲栎,而中午水势,叶片13C值高于槲栎。两种树木叶片13C值与含N量都成正相关关系。槲栎叶片13C值与清晨水势成负相关,而马尾松针叶13C值与清晨水势相关性并不显著。 在岸边的松栎混交幼林与成年林相比,幼树的清晨水势略高于对应的成年树,叶片13C值低于成年树;幼树的光合速率和气孔导度略高于成年树,而瞬时水分利用效率低于成年树,但差异均不显著。马尾松幼苗为实生苗,与成年树相比,更偏向于利用浅层土壤水;而槲栎幼树多为从原来被砍伐的树上萌生的,木质部水分同位素与成年树相近。 从2004年5月到10月,我们又对大坝下游江段岸边植物(马尾松,枫杨和柑桔)的水分关系进行了研究。木质部水分同位素比率表明,岸边植物木质部水分同位素比率与高处植物具相近的值,且显著高于江水同位素值。研究表明岸岸边和高处植物以降雨或靠降雨补充的浅层地下水为主要水分来源,即便岸边植物也几乎不利用江水。岸边植物与高处植物具有相近的清晨水势和中午水势,光合速率和气孔导度,以及叶片C值等,也进一步说明岸边植物与高处植物具有相近的水分生理特征。 叶片13C可以指示植物的一些生理过程,我们对松栎混交林中不同生活型植物的13C值进行了分析,乔木层叶片 13C值比较高,其中常绿针叶的值又高于落叶阔叶树木的值;林下灌木,非禾本科草本,及藤本植物的13C值都明显低于乔木层。 三峡大坝改变河水对植物生理生态过程的影响是一个长期的过程,库区水位上涨是否影响到岸边植物的生理过程及生长等,需要进一步作长期、定位和连续的观测研究。
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水分是半干旱草原生态系统植物多样性和初级生产力的限制因素。近年来由于人为干扰和气候干旱,特别是过度放牧,导致半干旱地区草原严重退化,草地生产利用和生态服务功能日益衰减,自然灾害频繁发生。草原退化演替主要表现为不同植物种的消长和替代过程。在内蒙古草原,过度放牧导致多年生根茎禾草和丛生禾草被豆科灌木或具根茎和不定根的杂类草替代,丰富的多年生杂类草被一、二年生植物替代。目前,对于上述功能群之间的替代机制,以及草原退化的机理尚不十分清楚。本研究在中国科学院内蒙古草原生态系统定位研究站选取了79年围封的大针茅(Stipa grandis)样地、其外围的自由放牧样地及中德合作项目“ Matter Fluxes in grassland of Inner Mongolia as influenced by stocking rate (MAGIM) ”的传统平地放牧处理小区为研究对象,从植物-水分关系的角度,利用氢和碳稳定性同位素技术,研究主要植物种和功能群水分来源和水分利用效率对放牧的响应,进而揭示放牧干扰下草原退化的机理。本研究取得了如下主要研究结果和结论: 1. 在干旱年份,雨水仅湿润了大针茅草原0-20cm的表层土壤。土壤中保存的冬季降水贡献了植物可利用水分的30%,而在正常年份只有10%。干旱季节羊草(Leymus chinensis)和大针茅能够利用土壤60cm以下保存的多年降水,而糙隐子草(Cleistogenes squarrosa)和冰草(Agropyron cristatum)只能利用生长季的雨水。长期过度放牧使冰草和糙隐子草的相对生物量和相对多度均显著增加,而羊草的相对生物量和相对多度显著降低。这种由放牧引起的群落结构与组成的改变,降低了植被对冬季降水的利用,加剧了干旱对生态系统的影响。 2. 依据羊草草原群落植物的生活型和光合类型划分为:C4禾草、C4杂类草、C4灌木、C3禾草、C3杂类草、C3鳞茎类植物和C3灌木七个功能群,各功能群间的水分利用效率差异显著。不同功能群植物在对土壤水分的利用上存在着明显的生态位分离。C4禾草和C3鳞茎类植物主要利用表层水分,而C4灌木则利用深层土壤水分,C3其它功能群介于二者之间。在无放牧和低载畜率下,各功能群植物均主要吸收浅层土壤水分,功能群间竞争相对强烈。在较高载畜率下,C3禾草转为利用深层土壤水分,功能群间出现水分生态位的分离,部分地缓解了植物对土壤表层水分的竞争。沿着载畜率梯度,群落对浅层土壤水分的利用呈现出先增加,尔后下降的趋势。 3. 在放牧干扰下,羊草草原群落中主要植物种稳定性碳同位素值的变化范围为-13.100/00~-27.590/00。不同功能群植物间的水分利用效率有着显著的差异:C4禾草> C4杂类草> C4灌木> C3禾草、C3杂类草> C3鳞茎类植物> C3灌木。C3鳞茎类植物和C4杂类草的水分利用效率随载畜率增加而降低。C3和C4禾草的水分利用效率随载畜率的增加表现为先增加,尔后下降的趋势。群落水平的水分利用效率主要由占生物量90%的禾草的水分利用效率决定。随着载畜率增加,群落水分利用效率有逐渐降低的趋势。
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采用裂区试验设计,对黄土塬区补充灌溉及氮磷配施条件下麦田土壤水分动态、作物产量及水分利用效率等进行研究。结果表明:1)冬小麦对土壤水分的利用深度随小麦生长发育逐渐加深,在越冬前期和孕穗期分别达1.2和2.2 m土层以下,不同处理土壤含水量在小麦生育前期差异不明显,孕穗后氮磷配施处理的土壤含水量显著低于不施肥处理;2)试验条件下,补充灌溉后同样施肥处理的作物产量与雨养相比,虽有增加但不显著;不论是雨养水平,还是补充灌溉水平,氮磷配施均表现出显著的增产效果,从低氮低磷到高氮高磷,增产幅度在134%到240%之间;3)氮磷配施能显著提高冬小麦水分利用效率,而补充灌溉后水分利用效率降低3%~30%,但未达显著水平;4)不同氮磷配施的增产效应高于补充灌溉,补充灌溉与高氮高磷处理有显著的水肥协同效应,能显著提高作物产量并保持较高的水分利用效率。
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为了探讨不同灌溉制度对玉米根系生长和水分利用效率的影响及基因型间差异,在大型活动防雨棚和棚外田间条件下,利用一组玉米遗传材料杂交种户单四号、父本803和母本天四进行了研究。结果发现玉米杂交种在根系生长、分布和水分利用效率上表现出显著的杂种优势。在充分灌溉条件下,玉米杂交种在浅层的根长密度大于亲本,但在水分亏缺条件下,玉米杂交种根长密度在整个剖面上都显著大于亲本;同一玉米基因型在不同的灌溉制度下根长密度在土壤剖面的分布也不同,拔节期不灌溉条件下玉米根系在深层土壤中的分布较充分灌溉条件下大,保证了玉米对深层土壤水分的充分吸收,而后期灌水延缓了表层根系生长的衰退,产生明显的补偿效应;拔节期干旱而抽雄期和灌浆期灌水显著提高了3种基因型玉米的水分利用效率。通过合理灌溉优化玉米根系分布特性以提高玉米吸水能力和水分利用效率,是节水栽培上的可行途径。
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利用室内控制试验研究了根据不同深度土壤水分传感器灌溉处理对冬小麦生物学性状及水分利用率等的影响。结果显示,以10 cm探头控制灌溉最为省水,同时冬小麦生物学性状及水分利用率等最佳。由于试验冬小麦处于生育前期,随着小麦生育期延伸,当根系超过30 cm深度时,根系吸水的深度增加,探头的埋设深度需要田间试验进行更详细的研究。