A review of belowground biomass allocation and its response to global climatic change in grassland ecosystems北大核心CSCD

Biomass allocations between aboveground and belowground organs provide pivotal information for connecting aboveground productivity and belowground carbon sequestration. As accurate measurement of belowground biomass is essential for determining the biomass allocation, we first reviewed the methods in quantifying belowground biomass and their merits. We then presented the major advances on plant biomass allocations between aboveground and belowground organs, as well as the potential drivers such as precipitation, warming, atmospheric CO2 concentration, and nitrogen deposition. We finally provided a list of challenges in studying belowground biomass allocation for the future. This review has important implications for studies on carbon cycling in grassland ecosystems under the changing climate.     

Biomass estimation models and allocation patterns of 14 shrub species in Mountain Luya,Shanxi, China

Aims Shrub species have evolved specific strategies to regulate biomass allocation among various organs or between above- and belowground biomass and shrub biomass model is an important approach to estimate biomass allocation among different shrub species. This study was designed to establish the optimal estimation models for each organ (leaf, stem, and root), aboveground and total biomass of 14 common shrub species in Mountain Luya, Shanxi Province, China. Furthermore, we explored biomass allocation characteristics of these shrub species by using the index of leaf biomass fraction (leaf to total biomass), stem biomass fraction (stem to total biomass), root biomass fraction (root to total biomass), and root to shoot mass ratio (R/S) (belowground to aboveground biomass).
Methods We used plant height, basal diameter, canopy diameter and their combination as variables to establish the optimal biomass estimation models for each shrub species. In addition, we used the ratios of leaf, stem, root to total biomass, and belowground to aboveground biomass to explore the difference of biomass allocation patterns of 14 shrub species.
Important findings Most of biomass estimation models could be well expressed by the exponential and linear functions. Biomass for shorter shrub species with more stems could be better estimated by canopy area; biomass for taller shrub species with less stems could be better estimated by the sum of the square of total base diameter multiply stem height; and biomass for the rest shrub species could be better estimated by canopy volume. The averaged value for these shrub species was 0.61, 0.17, 0.48, and 0.35 for R/S, leaf biomass fraction, stem biomass fraction, and root biomass fraction, respectively. Except for leaf biomass fraction, R/S, stem biomass fraction, and root biomass fraction for shrubs with thorn was significantly greater than that for shrubs without thorn.     

Variation in biomass allocation of Nitraria tangutorum during different phenological phases

植物生物量的分配模式是植物对环境适应的结果, 并伴随着植物生活史的每一个阶段, 与植物的生长和发育息息相关。目前关于植物生物量分配的大小依赖性已有相关报道, 但关于其对物候期的响应尚鲜有报道。该研究以乌兰布和沙漠地区白刺(Nitraria tangutorum)为研究对象, 通过对其2016与2017年连续2个生长季里盛花期、盛果期与营养生长期3个物候期的根、压条、新枝、老枝、叶、繁殖器官等部分的生物量测定, 采用标准化主轴回归方程的斜率和截距的显著性比较, 分别探讨了白刺在不同物候期的异速生长的大小依赖程度和相对生物量分配比例, 特别是地上与地下部分之间、支持与同化器官之间, 在不同物候期的生物量分配规律。结果表明: 白刺的繁殖生长对生物量分配模式造成的影响主要体现在相对生物量分配比例(36.00%)而不是个体大小的依赖性程度上(16.67%), 且对新枝的影响较大, 使其不同物候期的大小依赖性程度发生了改变, 但是变化趋势不一致, 同时繁殖生长增加对叶片的相对生物量分配比例, 减少对老枝的相对生物量分配比例, 但并没有改变他们的大小依赖性程度。白刺生长过程中的地下部分生物量分配率随个体生物量的累积均增大, 而繁殖分配会在一定程度内减弱这种速率。白刺随着个体生物量的增大其生物量向支持器官分配率也越大, 但随着生长时间推移, 更倾向于将生物量分配给同化吸收器官。     

Biomass allocation and carbon density of Sophora moorcroftiana shrublands in the middle reaches of Yarlung Zangbo River,Xizang, China

Aims The expansion of shrublands is considered as one of the key reasons leading to the increase of carbon density in terrestrial ecosystems in China. In the present study, our aims were to explore the biomass allocation and carbon density of Sophora moorcroftiana shrublands in Xizang.
Methods We sampled the biomass of S. moorcroftiana shrubs from 18 sites in the middle reaches of Yarlung Zangbo River, Xizang. Using concentrations of different organs, we estimated the carbon density of different layers in S. moorcroftiana shrublands.
Important findings The plant cover rather than biomass volume (the product of cover and height) provided the best fit for aboveground biomass. The average of the total biomass was 5.71 Mg·hm^-2, ranging from 2.32 to 8.96 Mg·hm^-2. The average biomass of shrub layer, the main component of shrub ecosystem, was 4.08 Mg·hm^-2, accounting for 71% of the total biomass. The belowground biomass of shrub and herb layers was 2.08 and 0.86 Mg·hm^-2, respectively, which was higher than the corresponding aboveground biomass. The average biomass carbon density was 2.48 Mg·hm^-2. Shrub vegetation in the eastern part of the middle reaches has lower carbon density than that in the western part. The relatively high biomass allocation to roots to increase water and nutrient undertake as well as physical support for plants is an important strategy of S. moorcroftiana to cope with the arid environment on the Qinghai-Xizang Plateau. Moreover, the lower carbon density in the eastern part of the middle reaches might be due to the dry environment resulted from high temperature and evapotranspiration and enhanced human activities at low altitudes. The continuous decrease of evapotranspiration under scenarios of future climate change may lead to increase in carbon density in S. moorcroftiana shrublands.     

Estimation of biomass allocation and carbon density of Rhododendron simsii shrubland in the subtropical mountainous areas of China

Aims As an important potential carbon sink, shrubland ecosystem plays a vital role in global carbon balance and climate regulation. Our objectives were to derive appropriate regression models for shrub biomass estimation, and to reveal the biomass allocation pattern and carbon density in Rhododendron simsii shrubland.
Methods We conducted investigations in 27 plots, and developed biomass regression models for shrub species to estimate shrub biomass. The biomass of herb and litterfall were obtained through harvesting. Plant samples were collected from each plot to measure carbon content in different organs.
Important findings The results showed that the power and linear models were the most appropriate equation forms. The D and D²H (where D was the basal diameter (cm) and H was the shrub height (m)) were good predictors for organ biomass and total biomass of shrubs. All of the biomass models reached extremely significant level, and could be used to estimate shrub biomass with high accuracy. It was more difficult to predict leaf and annual branch biomass than stem biomass, because leaf and annual branch were susceptible to herbivores and inter-plant competition. The mean biomass of the shrub layer was 20.78 Mg·hm^-2, in which Rhododendron simsii and Symplocos paniculata biomass accounted for 93.63%. Influenced by both environment and species characteristics, the biomass of the shrub layer organs was in the order of stem > root > leaf > annual branch. The root:shoot ratio of the shrub layer was 0.32, which was less than other shrubs in subtropical regions. The relative higher aboveground biomass allocation reflected the adaptation of plants to the warm and humid environment for more photosynthesis. The mean total community biomass was 26.26 Mg·hm^-2, in which shrub layer, herb layer and litter layer accounted for 79.14%, 7.62% and 13.25%, respectively. Litter biomass was relatively high, which suggested that this community had high nutrient return. There were significant correlations among aboveground biomass, belowground biomass and total biomass of shrub layer and herb layer. The mean biomass carbon density of the community was 11.70 Mg·hm^-2 and the carbon content ratio was 44.55%. The carbon density was usually obtained using the conversion coefficient of 0.5 in previous studies, which could overestimate carbon density by 12.22%.     

Biomass allocation patterns of Loropetalum chinense

Aims Biomass is the most fundamental quantitative character of an ecosystem. Biomass allocation patterns reflect the strategies of plants to adapt various habitat conditions and play a vital role in evolution, biodiversity conservation and global carbon cycle. Loropetalum chinense shrub is one of the most dominant shrub types in subtropical China. The objectives of this study were to quantify the allometric relationships and the biomass allocation pattern among organs, and to investigate the effects of body size, shrub regeneration origin and site factors on allometry and biomass allocation.
Methods Individual samples of L. chinense were harvested from shrublands in subtropical China and were further divided into leaves, stems and roots. The allometric relationships between different organs were modeled with standard major axis (SMA) regression and the biomass allocation to different organs was quantified. The effects of body size, shrub regeneration origin and other habitat factors on allometry and allocation were examined using Pearson’s correlation analysis and multiple linear regressions.
Important findings The isometric scaling relationships between shoot and root changed to allometric relationships with increasing basal diameter. The scaling relationships between leaf and stem and between leaf and root were isometric for smaller diameter classes, while for larger diameter classes they were allometric. These relationships were significantly different among shrub regeneration origin types. The scaling relationships between different organs were not affected by habitat factors; while the coverage of shrub layer and slope affected biomass allocation due to their influences on the allometric relationships between different organs at the initial stage of growth. The mean dry mass ratios of leaf, stem, root and the mean root to shoot ratio were 0.11, 0.55, 0.34 and 0.65, respectively. With the increase of basal diameter class, stem mass ratio (0.50-0.64) increased, while leaf mass ratio (0.12-0.08) and root mass ratio (0.38-0.28) decreased, and consequently root to shoot ratio (0.91-0.43) also decreased. In secondary shrublands, the leaf mass ratio was 0.12 and the root mass ratio was 0.33, while these values were 0.07 and 0.36 respectively in natural shrublands. The ratio of aboveground allocation was significantly correlated to shrub layer coverage (r = 0.44, p < 0.05). Leaf mass ratio was significantly correlated to slope (r = -0.36, p < 0.05) and root mass ratio was significantly correlated to mean annual temperature (r = 0.34, p < 0.05). Results showed that with the increase of body size, the scaling relationships between different organs of L. chinense changed from isometric to allometric, and more biomass was allocated to aboveground part, and concretely, to stems. Human disturbance affected biomass allocation by its influences on the allometric relationships between different organs, and by increasing biomass allocation to leaves and decreasing allocation to roots. Reduced light resource promoted the biomass allocation to aboveground part, and higher slope resulted in decreased biomass allocation to leaves, while higher mean annual temperature promoted biomass allocation to roots. The variation in annual precipitation had no significant influences on biomass allocation. The biomass allocation strategies of L. chinense partially support the optimal partitioning theory.     

Response of plant biomass to nitrogen addition and precipitation increasing under different climate conditions and time scales in grassland

生产力是草地生态系统重要的服务功能, 而生物量作为生态系统生产力的主要组成部分, 往往同时受到氮和水分两个因素的限制。在全球变化背景下, 研究草地生态系统生物量对氮沉降增加和降水变化的响应具有重要意义, 但现有研究缺乏对其在大区域空间尺度以及长时间尺度上响应的综合评估和量化。本研究搜集了1990-2017年间发表论文的有关模拟氮沉降及降水变化研究的相关数据, 进行整合分析, 探讨草地生态系统生物量对氮沉降和降水量两个因素的变化在空间和时间尺度上的响应。结果表明: (1)氮添加、增雨处理以及同时增氮增雨处理都能够显著地提高草地生态系统的地上生物量(37%, 41%, 104%)、总生物量(32%, 23%, 60%)和地上地下生物量比(29%, 25%, 46%)。单独增雨显著提高地下生物量(10%), 单独施氮对地下生物量影响不显著, 但同时增雨则能显著提高地下生物量(43%); (2)氮添加和增雨处理对草地生态系统生物量的影响存在明显的空间变异。在温暖性气候区和海洋性气候区的草地生态系统中, 氮添加对地上、总生物量及地上地下生物量比的促进作用更强, 而在寒冷性气候区和温带大陆性气候区的草地生态系统中, 则增雨处理对地下、总生物量的促进作用更强; (3)草地生态系统生物量对氮添加和增雨处理的响应也存在时间格局上的变化, 地下生物量随着氮添加年限的增加有降低的趋势, 地上、总生物量及地上地下生物量比则有增加的趋势。增雨年限的增加对总生物量没有明显的影响, 但持续促进地上生物量和地下生物量, 增加地上地下生物量比, 可见长期增氮、长期增雨对地上生物量的促进作用更明显。     

Plant leaf traits,height and biomass partitioning in typical ephemerals under different levels of snow cover thickness in an alpine meadow

Aims In the cold life zones, snow cover is a comprehensive environmental factor that directly influences soil temperature, soil water content, light and nutrient availability. Plants in these zones develop a series of unique mechanisms involving phenological characteristics, reproductive strategies, physiology and morphology to adapt to environmental changes. This paper is focused on the responses of plant leaf traits, height and biomass partitioning to variations in snow cover thickness, in order to better understand the responses of plant functional traits and specific adaptation strategies under global climate change scenarios. Methods Three transects were established along a gradient of snow cover in an alpine meadow of Mt. Kaka, in the eastern Qinghai-Xizang Plateau. Primula purdomii, Pedicularis kansuensis and Ranunculus tanguticus, which are three widely distributed and dominant ephemerals, were sampled and studied, particularly at their blooming stages. Plant height, specific leaf area (SLA) and biomass partitioning were measured accordingly. Important findings The values of SLA in Pedicularis kansuensis and R. tanguticus were relatively greater under better soil conditions; it was smaller in Primula purdomii with thick snow cover. The relationship between aboveground biomass and belowground biomass in Primula purdomii was allometric at sites with both thick and thin snow cover. No significant relationships were found between aboveground biomass and belowground biomass in Pedicularis kansuensis and R. tanguticus at some individual sites. However, when samples of the three species were pooled, the relationships between aboveground biomass and belowground biomass were allometric at all sites, which did not support isometric scaling hypothesis. In addition, on sites with either thick or thin snow cover, aboveground biomass had greater rate of accumulation than belowground biomass; whereas on sites with medium snow cover, the rate of biomass accumulation was greater for belowground component than aboveground component. Functional traits and biomass variables were better correlated in Primula purdomii and Pedicularis kansuensis than in R. tanguticus.     

Aboveground Tree Growth Varies with Belowground Carbon Allocation in a Tropical Rainforest Environment

Young secondary forests and plantations in the moist tropics often have rapid rates of biomass accumulation and thus sequester large amounts of carbon. Here, we compare results from mature forest and nearby 15–20 year old tree plantations in lowland Costa Rica to evaluate differences in allocation of carbon to aboveground production and root systems. We found that the tree plantations, which had fully developed, closed canopies, allocated more carbon belowground - to their root systems - than did mature forest. This increase in belowground carbon allocation correlated significantly with aboveground tree growth but not with canopy production (i.e., leaf fall or fine litter production). In contrast, there were no correlations between canopy production and either tree growth or belowground carbon allocation. Enhanced allocation of carbon to root systems can enhance plant nutrient uptake, providing nutrients beyond those required for the production of short-lived tissues such as leaves and fine roots, and thus enabling biomass accumulation. Our analyses support this deduction at our site, showing that enhanced allocation of carbon to root systems can be an important mechanism promoting biomass accumulation during forest growth in the moist tropics. Identifying factors that control when, where and for how long this occurs would help us to improve models of forest growth and nutrient cycling, and to ascertain the role that young forests play in mitigating increased atmospheric carbon dioxide.     

Effects of enclosure on carbon density of plant-soil system in typical steppe and desert steppe in Nei Mongol,China

草地生态系统是巨大的碳库, 在全球碳循环中起着重要的作用。该研究以内蒙古中温带草地区典型草原和荒漠草原为研究对象, 测定了两种草原类型围封与放牧后地上生物量碳密度、地下生物量碳密度和土壤碳密度, 探讨围封对两种草原类型植被-土壤系统碳密度的影响。结果表明: (1)围封显著地增加了典型草原地上和地下生物量的碳密度, 对荒漠草原地上生物量碳密度增加影响显著, 对地下生物量碳密度增加影响不显著; (2)围封显著地增加了典型草原土壤碳密度, 使荒漠草原土壤碳密度有增加的趋势, 但影响不显著; (3)典型草原围封样地地下生物量和土壤碳密度的垂直分布显著高于放牧样地, 而荒漠草原围封样地地下生物量和土壤碳密度的垂直分布与放牧样地的差异不显著; (4)围封分别提高了典型草原和荒漠草原植被-土壤系统碳密度的2.2倍和1.6倍, 典型草原和荒漠草原分别有超过65%和89%的碳储存在土壤中, 两种草原类型的地下生物量碳库均占总生物量碳库的90%以上。研究结果表明围封能够有效地增加草原生态系统的碳储量。     

Distribution of biomass in relation to environments in shrublands of temperate China

Aims Shrubland is one of the most widely distributed vegetation types in northern China. Previous studies on pattern and dynamics of plant biomass have been focused on forest and grassland ecosystems, while relevant knowledge on shrubland ecosystems is lacking. It is important to include shrublands in northern China to improve the accuracy in estimating the terrestrial ecosystem biomass in China.
Methods Based on investigations and samplings from 433 shrubland sites, we explored the distribution and allocation patterns of biomass in relation to climatic and soil nutrient factors of shrublands of temperate China.
Important findings The average shrubland biomass density in northern China is 12.5 t·hm^-2. It decreases significantly from temperate deciduous shrubland in northeast to desert shrubland in northwest. The average biomass density of temperate deciduous shrubland, alpine shrubland, and desert shrubland is 14.4, 28.8, and 5.0 t·hm^-2, respectively. Within temperate deciduous shrublands, plant biomass is lower in North China than in Northeast China. The average aboveground and belowground biomass density of shrub layer is 4.5 and 5.4 t·hm^-2, respectively; while that of grass layer is 0.8 and 1.8 t·hm^-2, respectively. Environmental factors affect biomass allocation across different plant organs. The belowground-aboveground biomass ratio of shrub exhibits no significant changes with environmental variables. The leaf-stem ratio increases with annual precipitation, and leaf biomass is low in arid region.     

Hierarchical plant responses and diversity loss after nitrogen addition: testing three functionally-based hypotheses in the Inner Mongolia grassland

Background

Numerous studies have shown that nitrogen (N) deposition decreases biodiversity in terrestrial ecosystems. To explain the N-induced species loss, three functionally based hypotheses have been proposed: the aboveground competition hypothesis, the belowground competition hypothesis, and the total competition hypothesis. However, none of them is supported sufficiently by field experiments. A main challenge to testing these hypotheses is to ascertain the role of shoot and root competition in controlling plant responses to N enrichment. Simultaneously examining both aboveground and belowground responses in natural ecosystems is logistically complex, and has rarely been done.

Methodology/Principal Findings

In a two-year N addition experiment conducted in a natural grassland ecosystem, we investigated both above- and belowground responses of plants at the individual, species, and community levels. Plants differed significantly in their responses to N addition across the different organizational levels. The community-level species loss was mainly due to the loss of perennial grasses and forbs, while the relative abundance of plant species was dependent mainly on individual-level responses. Plasticity in biomass allocation was much smaller within a species than between species, providing a biological basis for explaining the functionally based species loss. All species increased biomass allocation to aboveground parts, but species with high belowground allocations were replaced by those with high aboveground allocations, indicating that the increased aboveground competition was the key process responsible for the observed diversity loss after N addition in this grassland ecosystem.

Conclusions/Significance

Our findings shed new light on the validity of the three competing hypotheses concerning species loss in response to N enrichment. They also have important implications for predicting the future impacts of N deposition on the structure and functioning of terrestrial ecosystems. In addition, we have developed a new technique for ascertaining the roles of aboveground and belowground competition in determining plant responses to N fertilization.     

Response and correlation of above- and below-ground functional traits of Leymus chinensis to nitrogen and phosphorus additions

AimsLeymus chinensis is a constructive and dominant species in typical steppe of northern China. The structure and functions of L. chinensis grassland ecosystem has been degenerated seriously due to long-term overgrazing in recent decades. As an effective measure to restore the degraded grasslands, the effects of nutrient addition on plant growth and ecosystem structure and functioning have been paid more attention in manipulation experimental research. The effects of nutrient addition, especially P addition on the above- and below-ground functional traits of L. chinensis have rarely been studied; particularly the underpinning mechanisms remain unclear. Our objective is to examine the responses and adaptive mechanisms of L. chinensis to different levels of N and P additions. MethodsWe conducted a culture experiment in the greenhouse, with three levels of N (50, 100 and 250 mg N·kg^-1) and P (5, 10 and 25 mg P·kg^-1) addition treatments. The above- and below-ground biomass, leaf traits (e.g., specific leaf area, leaf N and P contents) and root traits (e.g., specific root length, root N and P contents) of L. chinensis were determined in this study.Important findings Our results showed that: 1) the aboveground biomass and total biomass of L. chinensis were mostly affected by N addition, while the belowground biomass was mainly affected by P addition. N addition greatly enhanced the aboveground biomass of L. chinensis, while P addition reduced the belowground biomass at the moderate and high N levels. The root-shoot ratio of L. chinensis was influenced by both N and P additions, and root-shoot ratio decreased with increasing N and P levels. N and P additions promoted more biomass and N and P allocations to aboveground and leaf biomass. 2) Leymus chinensis showed different responses and adaptive mechanisms to P addition at low and high N levels. At low N level, L. chinensis exhibited high photosynthetic rate and specific root length (SRL) to improve photosynthetic capacity and root N acquisition, which promoted aboveground biomass. High root P content was favorable for belowground biomass. At high N level, P addition did not significantly affect plant growth of L. chinensis, even reduced its belowground biomass. Leymus chinensis showed high specific leaf area (SLA) and SRL to improve light interception and N acquisition in order to maintain stable aboveground biomass. 3) P addition greatly impacted below-ground than above-ground functional traits. SLA exhibited a weakly positive correlation with SRL, indicating L. chinensis exhibited relatively independence of resource acquirement and utilization between leaf and root functional traits.     

Combining tree-ring analyses on stems and coarse roots to study the growth dynamics of forest trees: a case study on Norway spruce (Picea abies [L.] H. Karst)

We show the potential of a new method combining tree-ring analyses on stems and on coarse roots of individual trees in order to advance the understanding of growth dynamics in forest trees. To this end, we studied the root–shoot allometry of trees and its dependence on site conditions. Along a gradient in water supply in Southern Germany from dry to moist sites we selected 43 Norway spruce trees (Picea abies [L.] H. Karst.) aged 65–100 years. Increment cores were taken from stem and main roots revealing aboveground and belowground growth course over the last 34 years. Annual growth rates in roots and stems and their allometric relationships were applied as surrogate variables for tree resource allocation to aboveground and belowground organs. The mean sensitivities of both stem and root chronologies were found to be site-specific, and increased from the moist through the dry sites. No temporal offset between aboveground and belowground growth reactions to climate conditions was found in Norway spruce at any of the sites. These results suggest that the root–shoot allometry depends on the specific site conditions only at the driest site, following the optimal biomass partitioning theory (the more restricted the water supply, the more organic matter allocation into the belowground organs).     

Climate factors affect forest biomass allocation by altering soil nutrient availability and leaf traits

Biomass in forests sequesters substantial amounts of carbon; although the contribution of aboveground biomass has been extensively studied, the contribution of belowground biomass remains understudied. Investigating the forest biomass allocation is crucial for understanding the impacts of global change on carbon allocation and cycling.Moreover, the question of how climate factors affect biomass allocation in natural and planted forests remains unresolved. Here, we addressed this question by coll...     

白刺不同物候期的生物量分配规律

植物生物量的分配模式是植物对环境适应的结果, 并伴随着植物生活史的每一个阶段, 与植物的生长和发育息息相关。目前关于植物生物量分配的大小依赖性已有相关报道, 但关于其对物候期的响应尚鲜有报道。该研究以乌兰布和沙漠地区白刺(Nitraria tangutorum)为研究对象, 通过对其2016与2017年连续2个生长季里盛花期、盛果期与营养生长期3个物候期的根、压条、新枝、老枝、叶、繁殖器官等部分的生物量测定, 采用标准化主轴回归方程的斜率和截距的显著性比较, 分别探讨了白刺在不同物候期的异速生长的大小依赖程度和相对生物量分配比例, 特别是地上与地下部分之间、支持与同化器官之间, 在不同物候期的生物量分配规律。结果表明: 白刺的繁殖生长对生物量分配模式造成的影响主要体现在相对生物量分配比例(36.00%)而不是个体大小的依赖性程度上(16.67%), 且对新枝的影响较大, 使其不同物候期的大小依赖性程度发生了改变, 但是变化趋势不一致, 同时繁殖生长增加对叶片的相对生物量分配比例, 减少对老枝的相对生物量分配比例, 但并没有改变他们的大小依赖性程度。白刺生长过程中的地下部分生物量分配率随个体生物量的累积均增大, 而繁殖分配会在一定程度内减弱这种速率。白刺随着个体生物量的增大其生物量向支持器官分配率也越大, 但随着生长时间推移, 更倾向于将生物量分配给同化吸收器官。     

黄河口新生湿地碱蓬生物量及氮累积与分配对外源氮输入的响应

2014年4—11月,选择黄河入海口北部滨岸高潮滩的碱蓬湿地为研究对象,基于野外原位氮输入模拟试验,研究了不同氮输入梯度下(N0,无氮输入;N1,低氮输入;N2,中氮输入;N3,高氮输入)碱蓬不同器官生物量以及氮累积与分配特征的差异。结果表明,尽管不同氮输入处理并未改变碱蓬地上生物量的季节变化模式,但在不同程度上均促进了地上生物量的增长(平均增幅为19.71%—62.29%)且整体表现为N3N2N1N0;不同氮输入处理亦延长了碱蓬的生长高峰期,N1、N2和N3处理下地上生物量达到最大值的时间相对于N0处理推迟20 d左右。与地上生物量不同,不同氮输入处理改变了地下生物量的季节变化模式,特别是N2和N3处理均对生长初期的地下生物量产生了明显促进作用,且其初期地下生物量达到较高值的时间相对于N1和N0处理提前20—50d。不同氮输入处理下的枯落物量在产生初期和中期均增幅不大,末期则骤然增加且整体表现为N3N2N1N0。不同氮输入处理下碱蓬各器官的全氮(TN)含量总体上均表现为叶茎根,叶是氮的主要累积器官。尽管不同氮输入处理并未改变碱蓬不同器官的氮累积与分配格局以及地上与地下之间的养分供给关系,但其为适应不同养分条件而调整自身养分供给与分配的特性在N2处理下表现的尤为明显。研究发现,N2处理下碱蓬种子的发育时间相对于N0、N1和N3处理可能会提前约1个月,原因可能与N2氮输入水平可显著影响碱蓬体内的碳分配比以及碱蓬对适量氮养分输入环境的特殊适应对策有关。随着黄河口新生湿地氮养分供给的不断增加,当未来碱蓬湿地氮养分状况达到较高水平(特别是中等水平)时,其生物量、生长节律(特别是种子发育时间)以及不同器官氮累积与分配状况可能将发生明显改变。     

黑河中游荒漠草地地上和地下生物量的分配格局

草地生态系统中地上和地下生物量的分配方式对于研究生态系统碳储量和碳循环有着重要的意义。为了解黑河中游荒漠草地的地上和地下生物量分配格局, 从群落和个体两个水平对黑河中游的地上和地下生物量进行了调查。结果表明: 群落水平上地上生物量介于3.2-559.2 g·m^-2之间, 地下生物量介于3.3-188.2 g·m^-2之间, 个体水平上地上生物量介于6.1-489.0 g·株^-1之间, 地下生物量介于2.4-244.2 g·株^-1之间, 群落水平上的根冠比(R/S)为0.10-2.49, 个体水平上为0.07-1.55, 地下生物量均小于地上生物量, 群落水平上R/S值大于个体水平。群落和个体水平地上和地下生物量的拟合斜率分别为1.1001和0.9913, 与1没有显著差异, 说明地上与地下生物量呈等速生长关系。群落和个体水平土壤表层0-20 cm和0-30 cm的根系生物量分别占全部根系生物量的89.81%、96.95%和81.42%、93.62%, 表明地下生物量主要集中在0-20 cm和0-30 cm土壤表层。     

The role of belowground competition and plastic biomass allocation in altering plant mass–density relationships

Metabolic scaling theory (MST) predicts a ‘universal scaling law’ for plant mass–density relationships, but empirical observations are more variable. Possible explanations of this variability include plasticity in biomass allocation between the above‐ and belowground compartment and different modes of competition, which can be asymmetric or symmetric. Although complex interactions of these factors are likely to occur, so far the majority of modelling and empirical studies has focussed on mono‐factorial explanations. We here present a generic individual‐based model, which allows exploring the plant mass–density relationship in realistic settings by representing plasticity of biomass allocation and different modes of competition in the above‐ and belowground compartment. Plants grew according to an ontogenetic growth model derived from MST. To evaluate the behavior of the simulated plants related to the allocation patterns and to validate model predictions, we conducted greenhouse experiments with tree seedlings. The model reproduced empirical patterns both at the individual and population level. Without belowground resource limitation, aboveground processes dominated and the slopes of mass–density relationships followed the predictions of MST. In contrast, resource limitation led to an increased allocation of biomass to belowground parts of the plants. The subsequent dominance of symmetric belowground competition caused significantly shallower slopes of the mass–density relationship, even though the growth of individual plants followed MST. We conclude that changes in biomass allocation induced by belowground resource limitation explain the deviations from the mass–density relationship predicted by MST. Taking into account the plasticity of biomass allocation and its linkage to the above‐ and belowground competition is critical for fully representing plant communities, in particular for correctly predicting their response of carbon storage and sequestration to changing environmental conditions.     

地上竞争与地下竞争对科尔沁沙地榆树幼苗生长的影响

榆树疏林草原对科尔沁沙地植被恢复和景观保护有着重要意义.本文采用双因素两水平控制试验,从幼苗生物量、地下/地上生物量、茎高、根茎比、叶片数等方面,研究了草-树地上、地下竞争对科尔沁沙地榆树幼苗生长的影响.结果表明:对于1年生榆树幼苗,单株平均生物量表现为无竞争＞地上竞争＞全竞争＞地下竞争;地下/地上生物量表现为地下竞争＞全竞争＞无竞争＞地上竞争;幼苗高度表现为地上竞争＞无竞争＞全竞争＞地下竞争;根茎比表现为地下竞争＞全竞争＞无竞争＞地上竞争;叶片数表现为地上竞争＞无竞争＞地下竞争＞全竞争.地下竞争对1年生榆树幼苗生长影响显著,而地上竞争对榆树幼苗生长无显著影响.地上竞争与地下竞争对2年生榆树幼苗生长的影响均不显著.科尔沁沙地草本植物对榆树幼苗生长的影响主要通过地下竞争的方式实现,但地下竞争并没有改变榆树幼苗的资源分配方式.随榆树幼苗龄级的增长,草本植物竞争作用的影响逐渐减弱.