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不同土壤基质与施加氮肥对日本落叶松(Larix kaempferi)和长白落叶松(Lolgensis)竞争关系的影响
Alternative TitleEffects of two soil types and N fertilization on competition relationship between Larix kaempferi and L. olgensis
郭庆学
Subtype博士
Thesis Advisor李春阳
2017
Degree Grantor中国科学院大学
Place of Conferral北京
Degree Discipline生态学
Keyword人工林生产力 环境异质性 人工林管理 生态位分化
Abstract作为一种特殊的森林类型,人工林已经是全球森林生态系统的重要组成部分,在木材生产、水土保持和生物多样性等经济生产和生态功能领域,有非常重要的作用。我国是世界上人工林面积最大的国家,长白落叶松和日本落叶松人工林广泛分布在中国东北大部地区。施肥与构建混交林是提高人工林生产力的两种重要措施,本文以日本落叶松和长白落叶松为研究对象,探索不同土壤类型和施肥对两种落叶松竞争关系以及生长的影响,主要得到以下结论。首先,种间竞争与种内竞争对两种落叶松的根系特征影响不同。当两种落叶松混种时,相比种内竞争,二者的比根长(Specific root length: SRL)和比根尖密度(Specific root tip density: SRD)较高,而平均根系直径(Average diameter: AD)有降低的趋势。这表明,种间竞争时两种落叶松感知到异种的存在,通过调整根系结构,增强吸收土壤养分的能力。比如在第一次收获时(2014年8月末),在未施加N肥的落叶松土壤中,种间竞争促使两种落叶松叶片中的N浓度显著高于两物种各自的种内竞争状态,而叶片中C:N值显著的低于种内竞争时。其次,两种落叶松的种间竞争关系表现出时间上的差异。2014年,在未施加N肥的落叶松土壤以及天然次生林土壤中,两种落叶松的叶片、茎枝、根系和总生物量均未表现出明显的差异;施加N肥的落叶松土壤中,日本落叶松的生长受到长白落叶松的促进作用,使得前者的叶片、茎枝、根系和总生物量最高,相对竞争强度(Relative competition intensity: RCI)显著的高于长白落叶松,导致后者的生长受到轻微的抑制,而在其他条件下,二者的相对竞争强度则未有显著差异。然而,在2015年时,施加N肥的落叶松土中,日本落叶松的相对竞争强度降低,相反的,长白落叶松的竞争强度升高,二者的总生物量未有明显的差异;在未施加N肥的落叶松土壤中,长白落叶松的生长受到日本落叶松的促进作用,而在天然次生林土中,日本落叶松的生长受到长白落叶松的促进作用,后者受到抑制。日本落叶松土抑制日本落叶松的生长,在2015年,天然次生林土壤中,日本落叶松(种内和种间竞争)的生物量均显著较高,施N肥能有效的促进日本落叶松的生长。第三,竞争、施肥和土壤影响两种落叶松叶片、茎枝和根系的基本元素(C、N和P)以及非结构性碳水化合物浓度的变化(从2014到2015),比如在未施加N肥的日本落叶松土中,混种的日本落叶松茎枝和叶片的淀粉和总非结构性碳水化合物(Total non-structural carbohydrate, TNC)浓度变化显著的高于其纯种时。竞争、施肥和土壤也影响两物种N库、P库和非结构性碳水化合物的含量以及相对应的积累速率,结果表明,从2014年到2015年,生长受到促进的物种,其C、N和P以及非结构性碳水化合物含量的积累速率显著较高。最后,2015年7月末与8月末,不同条件下两种落叶松的净光合速率(Pn)以及光合N利用效率(Photosynthetic N-use efficiency, PNUE)表现不同,但是两种落叶松光合P利用效率(Photosynthetic P-use efficiency, PPUE)变化最为明显,比如在天然次生林土中,7月份两种落叶松的PPUE未出现明显的变化,但是在8月末份,混种时的日本落叶松PPUE显著高于其纯种条件下。从叶片的超微结构看出,不同的竞争条件下,受到促进的植物,单位视野内,其叶片中含有的叶绿体数量较多,其叶片的叶绿a/b值显著升高,而生长受到抑制的植株,叶绿素a/b值显著降低,这说明,叶绿素a/b可能反映出物种之间的竞争状态。物种、竞争和土壤影响着两种落叶松对15NO3--N和15NH4+-N的选择吸收偏好,在未施N的落叶松土中,与日本落叶松混种后,长白落叶松吸收15NO3--N显著增加,而在次生林土中,混种的日本落叶松对15NO3--N的吸收能力有升高的趋势,但是对15NH4+-N的吸收偏好显著高于其纯种条件下。结果还发现,实验中谷氨酸脱氢酶(GDH)、谷氨酰胺合成酶(GS)的活性以及叶片中各类氨基酸浓度未有明显的变化。总的来说,日本落叶松土对日本落叶松的生长有一定的抑制作用,施加N肥后,日本落叶松的生长受到促进,特别日本落叶松与长白落叶松混种时,两种落叶松的生物量和生长速率均最高,说明在施加N肥的日本落叶松土中,混种是提高落叶松人工林产量的有效措施。
Other AbstractForest plantations provide wood products to meet the social and economic demands and help to restore degraded land and to conserve biodiversity. China has the largest area plantation in the whole world, and Larix kaempferi and L. olgensis plantations are widely spread across the Northeast of China. Establishment of mixed plantation and fertilization are two important management approaches when trying to maintain soil qualities and productivity. In this study, two experiments were conducted. The main results were as follow:Firstly, Inter-specific competition tended to increase the specific root length (SRL) and specific root tip density (SRD), whereas the average root diameter (AD) tended to decline in the unfertilized larch soil (Larch soil N-). It indicated that the two Larix species could detect each other and positively changed root architecture to enhance their ability to absorb nutrient. As a result, after first harvest (late August, 2014), the leaf N concentration of the two species was significantly higher but the leaf C:N was significantly lower when they were in mixed culture.Secondly, the results showed that the relative competitive intensity (RCI) of the two Larix species showed considerable temporal changes from 2014 to 2015. In 2014, in the larch soil N- condition as well as the mixed-forest soil condition, biomass (leaf, shoot, root and total biomass) as well as their RCIs of the two species did not show obvious difference; however, in 2015, in the larch soil N- condition, the RCI of L. olgensis was significantly higher than the L. kaempferi which meant the growth of the former was facilitated by the later; conversely, RCI of the former species were significantly lower than the later in the mixed-forest soil. In 2014, the RCI of L. kaempferi was significantly higher when grown with L. olgensis in the N-fertilized larch soil (Larch soil N+ condition), whereas the RCI of L. olgensis was higher compared to that of L. olgensis in the mixed-forest soil in 2015. The results also indicated that the larch soil inhibit the growth of L. kaempferi but could be alleviated by N fertilization.Thirdly, competition, soil type or N fertilization caused temporal variation in C, N, P and non-structural carbohydrate concentrations from 2014 to 2015. For example, in the larch soil N- condition, concentration variations of shoot starch, soluble sugars and TNC (Total non-structural carbohydrate) as well as leaf starch and TNC of L. kaempferi from mixed culture were higher than those of individuals from L. kaempferi monoculture. The results also showed that the growth of L. kaempferi greatly benefited from the presence of L. olgensis in the mixed-forest soil and it tended to have higher C, N, P and non-structural carbohydrate accumulation ratios.Finally, net photosynthetic rates (Pn) and photosynthetic N-use efficiency (PNUE) of the two species were different between late July and August at 2015. However, the photosynthetic P-use efficiency (PPUE) variation was great. For example, PPUE of the two species in each treatment from the mixed-forest soil did not show any difference at late July, but PPUE of the mixed L. kaempferi was significantly higher than individuals from monoculture in the mixed-forest soil. The leaf mesophyll cell result showed that the benefited species had more chloroplasts. More interestingly, the chlorophyll a/b of the benefited individuals was significantly higher while that of the suppressed individuals was significantly lower. This implied that the chlorophyll a/b may reflect individual competitive status. Competition and soil type affected preference of 15NO3--N and 15NH4+-N. In the larch N- soil condition, the L. olgensis absorbed more 15NO3--N in the presence of L. kaempferi. The results also find out that the activity of glutamate dehydrogenase (GDH) and glutamine synthetase (GS) and concentrations of amino acids among each treatment showed no significant differences.In conclusion, the results indicated that larch soil inhibited the growth of L. kaempferi somewhat, but N fertilization alleviated this negative effect. Furthermore, the two species in mixed culture, especially under N fertilization greatly promoted productivity of the larch forest plantation. 
Pages104
Language中文
Document Type学位论文
Identifierhttp://ir.imde.ac.cn/handle/131551/24606
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
郭庆学. 不同土壤基质与施加氮肥对日本落叶松(Larix kaempferi)和长白落叶松(Lolgensis)竞争关系的影响[D]. 北京. 中国科学院大学,2017.
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