IMHE OpenIR  > 山地表生过程与生态调控重点实验室
干热河谷土壤典型因子对辣木补偿生长的影响
Alternative TitleEffect of Typical Soil Factors on the Compensatory Growth of Moringa oleifera in an Arid-Hot Valley
赵广
Subtype博士
Thesis Advisor刘刚才
2018
Degree Grantor中国科学院大学
Place of Conferral北京
Degree Discipline土壤学
KeywordN添加 P添加 AM真菌 侵染程度 去叶
Abstract植物叶片遭受虫害和动物采食是自然界普遍发生的事件,探明植物采食后再生生长的生理生态特征、阐明植物再生的条件或者效应,不仅具有重要生态学理论意义,而且对于合理利用植物的再生机制、提高和维持植物生产力、确定合理利用程度具有重要指导作用。金沙江干热河谷区是典型的低生产力地区,土壤养分和水分条件是限制植物生长和生态系统恢复的主要限制因素,而越来越多的研究发现AM真菌对于维持干热河谷脆弱生态系统稳定性存在重要作用。本研究以金沙江干热河谷地区引进的辣木(Moringa oleifera)为对象,通过研究土壤养分、水分和AM真菌侵染对辣木各器官氮磷含量、光合特性、生物量分配、非结构性碳水化合物和叶片功能性状等的影响,探讨土壤养分、水分和AM真菌对辣木再生过程中的固碳能力、物质和能量分配以及植物性状适应性等方面的生理生态影响机制。研究结果对于揭示植物补偿生长发生的养分效应和AM真菌与被采食植物间的反馈调节机制具有重要理论意义,同时也能为探明如何最大程度利用辣木再生生长潜能和干热河谷地区虫害防治,提供科学依据。主要结论如下:(1)氮、磷添加对辣木补偿生长的影响:辣木生物量整体上随着生长进行而逐渐升高。去枝50%在处理前期显著降低了总生物量,但后期植株生物量存在明显回升现象。氮、磷添加对辣木生长均有一定促进作用,但磷素添加对生物量的影响未达到显著性水平(P>0.05),而氮素则显著提高了植株生物量。各种处理下,施肥处理明显降低了植株去枝后地上生物量和地下生物量的直线拟合斜率,即去枝后辣木倾向于将更多的物质和能量资源投入到地上组织建成。去枝50% 处理在无养分添加处理中降低了辣木株高-地茎异速生长指数;氮添加或氮、磷同时添加,去枝对辣木植株株高-地茎异速生长指数的影响不显著。叶片比叶面积(SLA)随着植株生长呈下降趋势。无养分添加处理下(N-P-),去枝降低比叶面积,而养分添加的3个处理中(N+P-、N-P+和N+P+),去枝对比叶面积的无显著影响。无养分添加处理下(N-P-),去枝处理中叶绿素含量先降低后升高,此后去枝处理的叶绿素总含量均高于对照组。氮添加和氮、磷同时添加(N+P-和N+P+),去枝处理在75 d时对叶绿素总含量有明显下降作用。去枝对N-P-处理中叶片N含量的无显著影响,但各时期均值均高于对照处理。氮添加和氮、磷同时添加处理下,叶片N含量存在升高趋势。去枝50%在氮添加和氮磷同时添加处理中升高了叶片N含量,但未达到显著性水平(P>0.05)。(2)土壤水分胁迫对辣木补偿生长的影响水分添加和N添加处理下,辣木植株生物量虽然有显著升高,但并未出现超补偿生长;无水分和N添加,去叶处理下的辣木植株根茎叶生物量反而高于对照处理,去叶后发生超补偿生长。水分胁迫虽然对辣木植物各器官的生长发育产生抑制作用,但是在N添加处理中却能促进受到叶损失的辣木进行补偿性生长;N添加能显著提高辣木叶和茎的生物量,但对根的作用不显著,也不能促进植物进行超补偿生长。去枝在无N添加下能显著提高辣木分枝数,水分胁迫和去枝对辣木分枝数表现为“协同作用”,而N添加则表现为“拮抗作用”。植株各组织N含量整体表现为叶>茎>根;去叶处理下,植株叶片和茎的N含量在75% FC 中均降低,在N-W+下升高。根系N含量在N-W-、N-W+和N+W-处理下有上升趋势。辣木各组织磷含量整体表现为叶=茎>根;去叶对辣木在各处理下磷素含量变化不显著。去叶整体上提高氮/磷比,但是这种促进作用在恶劣土壤环境下更显著,即无N添加和水分胁迫中(N-W-),辣木各器官的氮/磷比在去叶后存在明显升高现象。95% FC中,无N添加的去叶处理会降低叶片光合速率, N添加在后期提高光合速率。75% FC中,去叶对无N添加下叶片光合速率的影响不显著,仍和未去叶处理维持在同一水平;N添加后,去叶处理后期显著提高了叶片光合速率。N添加后,水分胁迫(75% FC)使辣木叶绿素a和叶绿素b含量分别下降了9.4%―9.8%和8.24%―15.4%。土壤水分条件对叶和茎的可溶性糖与淀粉含量无显著影响。无N添加处理中,去叶会显著升高可溶性糖含量(P<0.05),而N添加处理中则降低。低养分环境下去叶更能促进叶片淀粉含量升高。去叶对茎可溶性糖含量的影响受到N添加的影响:无N添加处理中,去叶会显著降低茎的可溶性糖含量;N添加处理中,去叶显著升高了可溶性糖含量,降低淀粉浓度。去叶对根系淀粉含量的影响受到土壤水分状况的影响:正常土壤水分处理下(95% FC),N添加会降低根系淀粉含量,而去叶却未产生显著性影响;土壤水分胁迫处理中,N添加对根系淀粉的无显著性作用,而去叶却显著降低根系淀粉含量。(3)AM真菌和P添加对辣木补偿生长的影响从生物量上看,AM真菌对辣木生长和补偿生长的作用受到P添加的影响,对生物量分配的影响主要表现为降低茎/叶比、根/叶比和根/茎比;去叶整体上会降低根冠比,但在AMF+处理中,去叶对根冠比的降低作用被弱化。各器官N和P含量整体均表现为叶片>茎>根。去叶后,叶片N含量存在下降趋势,P添加显著提高了各器官P的含量。叶片P含量在去叶处理下均出现下降趋势,茎的P含量在P添加处理中去叶后也下降。AM真菌侵染和土壤P含量丰富时有助于稳定植株体内的氮/磷比。P添加在AMF-中提高了叶片光合速率,但在AMF-处理中却产生抑制作用。去叶和AM真菌整体上升高叶片瞬时水分利用效率,而P添加则降低水分利用效率。AMF-处理中,去叶显著降低了P-处理中的叶绿素a含量;但在AMF+处理中,去叶对无P添加处理的叶片叶绿素a含量无显著影响,表明AM真菌对低磷环境下去叶带来的叶绿素a含量降低有明显的补偿作用。此外,AM真菌对叶绿素b含量有明显促进效果。去叶均降低了比叶面积,叶片干物质含量在去叶后均出现升高趋势。AM真菌侵染显著提高了辣木叶片可溶性糖含量,这种促进作用在低磷环境下更显著;去叶后,叶片可溶性糖含量存在上升趋势。AM真菌对叶片淀粉浓度的的影响未达到显著性水平,而P添加显著降低叶片淀粉含量。AM真菌侵染对茎的可溶性糖含量没有显著影响,但茎的淀粉含量在在AM真菌侵染下有明显降低现象,表明真菌侵染会影响茎对淀粉的贮存。AM真菌会降低根系可溶性糖和淀粉含量,且去叶后AM真菌会显著降低根系淀粉含量(P<0.05)。(4)AM真菌侵染程度对辣木补偿生长的影响不去叶处理中,AM真菌侵染程度升高促进辣木叶片生长,但对根系生长存在限制作用,降低全植株生物量。中度去叶处理中(50%),AM真菌侵染程度增加可以提高根系生物量和根冠比,其中MR处理对叶再生有明显促进作用。重度去叶处理中(100%),AM真菌侵染增加能促进叶和茎的再生,但对根生长产生限制。AM真菌侵染程度升高显著降低了根冠比,表明AM真菌在一定程度上抑制植物根系生长。叶绿素a和叶绿素b含量均随着AM真菌侵染程度的上升先升高后下降。AM真菌侵染能提高辣木叶的比叶面积,但是去叶50%后辣木比叶面积均降低,且AM真菌侵染程度越高,比叶面积越小;当去叶100%时,比叶面积虽有上升,但随侵染程度升高仍保持相同变化规律。接种AM真菌会降低根系淀粉浓度。无去叶处理和50%去叶处理中,AM真菌侵染程度对根系淀粉浓度没有显著影响,但去叶100%处理时,根淀粉浓度随着菌根侵染程度的升高而逐渐降低。这些结果表明,AM真菌对辣木补偿生长的影响表现出的促进或抑制效应主要受叶片损失程度影响。
Other AbstractInsect herbivores and animal feeding are common occurrences in the natural world. Investing physiological characteristics of regenerating plants after eating, and clarifying the conditions or effects of plant regeneration, not only have important ecological theoretical significance, but also are meaningful for making rational use of plant’s regeneration mechanism, improving and maintaining the productivity of plants, and determining the degree of rational utilization. The dry-hot valley of Jinsha River is a typical low-productivity area. Soil nutrients and water conditions are the main limiting factors in the area that limit plant growth and ecosystem restoration, and studies have increasingly found that AM fungi play an important role in maintaining the stability of this fragile ecosystem. In this study, Moringa oleifera was selected as object of the study in the dry-hot valley, and nitrogen, phosphorus content, photosynthetic characteristics, biomass allocation, non-structural carbohydrates, and leaf functional traits were studied under different soil nutrient, water status, and AM fungi infection rate under pruning or defoliation treatment. The physiological effects of soil nutrient, water status, and AM fungi on the carbon fixation, material and energy allocation, and adaptability of plant traits were invested. The research results have important theoretical significance for revealing the nutrient effect of plant compensatory growth and the feedback regulation mechanism between AM fungi and the plant being fed. Meanwhile, it can also make a proposal on how to make the best use of regeneration potential of Moringa oleifera and how to prevent insect pests in dry and hot valleys. The main conclusions are as follows:(1) Effects of nitrogen and phosphorus addition on the compensatory growth of Moringa oleifera :The biomass gradually increased as the growing season progressed. Pruning of 50% significantly reduced the total biomass in the early stage of treatment, but there was a significant increase in the biomass in the late stage of treatment. The addition of nitrogen and phosphorus had a certain promotion effect on the growth of Moringa oleifera, but the effect of phosphorus addition on the biomass did not arrive a significant level (P>0.05), while nitrogen significantly increased the biomass under different treatment. Under various treatments, the fertilization treatment significantly reduced the straight-line fitting slope of above-ground biomass and underground biomass after pruning. That is to say, Moringa oleifera tends to put more material and energy resources into the aboveground part to build the plants after pruning. Underground tissues are limited in growth and development under adequate nutrient conditions. Pruning of 50% of the treatment reduced the Moringa oleifera plant height-basal diameter allometric index in the absence of nutrient treatment, indicating that pruning caused the plant to allocate more resources to the growth of the ground stem. Adding nitrogen or adding nitrogen and phosphorus at the same time, the removal of shoots had no significant effect on the allometry index of plant height and stem of Moringa oleifera plant.The leaf specific area (SLA) showed a decreasing trend with plant growth. No nutrient addition treatment (N-P-) and removal of shoots reduced specific leaf area, while nutrient additions of three treatments (N+P-, N-P+, and N+P+) had no significant effect on SLA. Under the treatment of N-P-, the chlorophyll content of the pruning treatment decreased firstly and then increased. Nitrogen addition and nitrogen and phosphorus were added at the same time (N+P- and N+P+). Pruning treatment significantly decreased the total chlorophyll content at 75 days. The removal of shoots had no significant effect on N content of leaves in N-P- treatment, but the mean value was higher in each period than the control. With the addition of nitrogen and the addition of both nitrogen and phosphorus, the N content in the leaves increased. At the same time, the content of N in leaves increased by 50% in the addition of nitrogen and nitrogen and phosphorus, but did not reach the level of significance.(2) The effect of soil water stress on compensatory growth of Moringa oleiferaUnder the condition of water addition and N addition, although the biomass of Moringa oleifera increased significantly, there was no over-compensation growth; without water and N addition (W-N-), the biomass of roots, leaves and leaves of Moringa oleifera under defoliation treatment were higher than the control treatment. After the leaves removing, Moringa oleifera have overcompensation growth. Although water stress limited the growth and development of various organs of Moringa oleifera, it could promote the compensatory growth of Moringa oleifera in the leaves during N addition. N addition can significantly increase the biomass of Moringa oleifera leaves and stems. However, the effect on the roots is not significant, nor does it promote the over compensation of plants. The branch removal without adding N could significantly increase the number of branches of Moringa oleifera. The water stress and pruning showed a “synergistic effect” on the number of branches of Moringa oleifera, and N addition showed “antagonism effect”.Nitrogen content of each tissue in the plant ranked as leaf>stem>root as a whole. Under the removal of leaves, nitrogen content in leaves and stems of plants decreased in 75% FC and increased under N-W+. The content of root N increased in N-W-, N-W+ and N+W-treatments. Phosphorus content in all tissues of Moringa oleifera was displayed as leaf=stem>root. The change of phosphorus content in Moringa oleifera was not significant under all treatments. The total N/P ratio was increased under leaves removing, but this promotion was more pronounced in the harsh soil environment. In the absence of N addition and water stress (N-W-), the N/P ratio of Moringa oleifera was obviously increased in the leaves. In 95% FC, the leaf removal without N addition reduced the photosynthetic rate of the leaves, and N addition increased the photosynthetic rate later. In 75% FC, the effect of leaf removal on the photosynthetic rate of leaves without N addition was not significant, and remained at the same level as the treatment without N removal. After N addition, the photosynthetic rate of leaves was significantly increased at the late leaf removal stage. After N was added, the water stress (75% FC) decreased the chlorophyll a and chlorophyll b contents by 9.4%-9.8% and 8.24%-15.4%, respectively.Soil moisture conditions had no significant effect on soluble sugar and starch content in leaves and stems. In the non-N addition treatment, the removal of leaves significantly increased the soluble sugar content (P<0.05), which was decreased during the N treatment. The lower nutrient environment further promoted the increase of leaf starch content. The effect of removing leaves on the soluble sugar content of the stem were influenced by N addition: in the absence of N, the removal of leaves significantly reduced the soluble sugar content of the stem, and in the N addition treatment, the removal of leaves significantly increased the soluble sugar content and decreased the starch concentration. The effect of defoliation on starch in root was influenced by soil water status: under normal soil moisture treatment (95% FC), N addition decreased root starch content, but no significant effect was observed in the leaves under defoliation; effect of N addition on root starch was not significant in soil water stress treatment, while the removal of leaves significantly reduced root starch content.(3) Effect of AM fungi and P addition on the compensatory growth of Moringa oleiferaFrom the biomass point of view, the effect of AM fungi on growth and compensatory growth of Moringa oleifera was affected by P addition, and the effect on biomass allocation was mainly reflected in reduced stem/leaf ratio, root/leaf ratio, and root/stem ratio; Overall, the root-shoot ratio will be reduced, but in the AMF+ treatment, the effect of removing the leaves from the root to shoot ratio is weakened.The contents of N and P in all organs were as follows: leaf> stem> root. After leaf removal, there was a downward trend in leaf N content. P addition significantly increased the content of P in various organs. The content of P in leaves showed a downward trend under the treatment of removing leaves, and the content of P in stems also decreased after the removal of leaves in P addition treatment. AM fungi infection and enrichment of soil P contributed to stabilizing the N/P ratio in the plant. Addition of P in AMF- enhanced leaf photosynthetic rate, but inhibited in AMF-treatment. Defoliation and AM fungi generally increased leaf instantaneous water use efficiency, whereas P addition decreased water use efficiency. In the AMF-treatment, the removal of leaves significantly reduced the content of chlorophyll a in the P- treatment; however, in the AMF+ treatment, the removal of leaves did not have a significant effect on the content of chlorophyll a in leaves without P treatment, indicating that the arbuscular mycorrhizal fungi have a negative impact on the low-phosphorus environment. The resulting reduction in chlorophyll a content has a significant compensatory effect. In addition, arbuscular mycorrhizal fungi significantly promoted the chlorophyll b content. Defoliation reduced the specific leaf area, and the dry matter content of leaves showed a tendency to increase after leaves removing.The infection of AM fungi significantly increased the soluble sugar content in leaves of Moringa oleifera, and this promotion was more pronounced in the low-phosphorus environment. There was an upward trend in soluble sugar content after leaf removal. The effect of AM fungi on leaf starch concentration did not reach a significant level, while P addition significantly reduced the starch content of leaves. AM fungi had no significant effect on the soluble sugar content of the stem, but the starch content of the stem was significantly decreased under the infection of the AM fungi, indicating that the fungal infection will affect the stem storage of starch. AM fungi decreased root soluble sugar and starch content, and AM fungi significantly reduced root starch content after leaf removal (P<0.05), indicating that AM fungi increased the consumption of stored carbon sources in plant roots. (4) Effect of AM fungal infection degree on the compensatory growth of Moringa oleiferaIn the 0% treatment of defoliation, the increase in the degree of infection by AM fungi promoted the growth of Moringa oleifera leaves, which limited the growth of roots and reduces the whole plant biomass. In moderate leaf removal treatment (50%), the increase of arbuscular mycorrhizal infection increased root biomass and root-shoot ratio, and MR treatment significantly promote leaf regeneration. In severe leaf removal treatments (100%), increasing arbuscular mycorrhizal infection promoted leaf and stem regeneration, but limited root growth. The increase in the degree of infection of AM fungi significantly decreased the ratio of root to shoot, indicating that AM fungi inhibited the growth of plant roots to a certain extent.The contents of chlorophyll a and chlorophyll b increased firstly and then decreased with the increase of the degree of infection of AM fungi. AM fungi infection could increase the specific leaf area of Moringa oleifera, but the specific leaf area of Moringa oleifera decreased after 50% leaf removal, and the higher the degree of infection of AM fungi have the smaller SLA; when the leaf was removed to 100%, the SLA increased, but maintaining the same changes with the increase in the degree of infection. Inoculation of arbuscular mycorrhizal fungi reduces root starch concentration. In the absence of leaf removal and 50% removal of leaves, the degree of AM fungi infestation had no significant effect on the starch concentration in the roots, but the root starch concentration decreased gradually with increasing mycorrhizal infection when 100% of leaves were removed. These results indicate that the promotion or inhibition effect of AM fungi on the compensatory growth of Moringa oleifera is mainly affected by the degree of leaf loss. 
Pages132
Language中文
Document Type学位论文
Identifierhttp://ir.imde.ac.cn/handle/131551/24776
Collection山地表生过程与生态调控重点实验室
Affiliation中国科学院成都山地灾害与环境研究所
First Author Affilication中国科学院水利部成都山地灾害与环境研究所
Recommended Citation
GB/T 7714
赵广. 干热河谷土壤典型因子对辣木补偿生长的影响[D]. 北京. 中国科学院大学,2018.
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