Modelling forest response to increasing CO2 concentration under nutrient-limited conditions

The growth rates of woody plants depend on both the rate of photosynthetic carbon gain and the availability of essential nutrients. Instantaneous carbon gain is known to increase in response to increasing atmospheric CO₂ concentration, but it is uncertain whether this will translate into increased growth in the longer term under nutrient-limited conditions. An analytical model to address this question was developed by Comins & McMurtrie (1993, Ecological Applications 3, 666–681). Their model was further tested and analysed. Manipulation of various assumptions in the model revealed its key assumptions and allowed a more confident prediction of expected growth responses to CO₂ enrichment under nutrient-limited conditions. The analysis indicated that conclusions about the CO₂ sensitivity of production were strongly influenced by assumptions about the relationship between foliar and heartwood nitrogen concentrations. With heartwood nitrogen concentration proportional to foliar nitrogen concentration, the model predicted a strong response of plant productivity to increasing CO₂ concentration, whereas with heartwood nitrogen concentration set constant, the model predicted only a very slight growth response to changing CO₂ concentration. On the other hand, predictions were only slightly affected by: (1) assumptions about the extent of nitrogen retranslocation out of senescing roots and foliage or wood during heartwood formation; (2) the effects of nitrogen status on specific leaf area or (3) leaf longevity; (4) carbon allocation between different plant parts; or (5) changes in the N:C ratio of organic matter sequestered in the passive pool of soil organic matter. Modification of the effect of foliar nitrogen concentration on the light utilization coefficient had only a small effect on the CO₂ sensitivity for pines. However, this conclusion was strongly dependent on the chosen relationship between single-leaf photosynthesis and leaf nitrogen concentration. Overall, the analysis suggested that trees growing under nitrogen-limited conditions can respond to increasing atmospheric CO₂ concentration with considerable increases in growth.     

The influence of nitrogen availability on carbon and nitrogen storage in the biennial Cirsium vulgare (Savi) Ten. I. Storage capacity in relation to resource acquisition, allocation and recycling

Plants of Cirsium vulgare (Savi) Ten. were cultivated under five different nitrogen regimes in order to investigate the effects of nitrogen supply on the storage processes in a biennial species during its first year of growth. External N supply increased total biomass production without changing the relationship between ‘productive plant compartments’ (i.e. shoot plus fine roots) and ‘storage plant compartments’ (i.e. structural root dry weight, which is defined as the difference between tap root biomass and the amount of stored carbohydrates and N compounds). The amount of carbohydrates and N compounds stored per unit of structural tap root dry weight was not affected by external N availability during the season, because high rates of N supply increased the concentration of N compounds whilst decreasing the carbohydrate concentration, and low rates of N supply had the opposite effect. Mobilization of N from senescing leaves was not related to the N status of the plants. The relationship between nitrogen compounds stored in the tap root and the maximum amount of nitrogen in leaves was an increasing function with increasing nitrogen supply. We conclude that the allocation between vegetative plant growth and the growth of storage structures over a wide range of N availability seems to follow predictions from optimum allocation theory, whereas N storage responds in a rather plastic way to N availability.     

Construction and maintenance of the optimal photosynthetic systems of the leaf, herbaceous plant and tree: an eco-developmental treatise

BACKGROUND AND AIMS: The paper by Monsi and Saeki in 1953 (Japanese Journal of Botany 14: 22-52) was pioneering not only in mathematical modelling of canopy photosynthesis but also in eco-developmental studies of seasonal changes in leaf canopies. SCOPE: Construction and maintenance mechanisms of efficient photosynthetic systems at three different scaling levels--single leaves, herbaceous plants and trees--are reviewed mainly based on the nitrogen optimization theory. First, the nitrogen optimization theory with respect to the canopy and the single leaf is briefly introduced. Secondly, significance of leaf thickness in CO2 diffusion in the leaf and in leaf photosynthesis is discussed. Thirdly, mechanisms of adjustment of photosynthetic properties of the leaf within the herbaceous plant individual throughout its life are discussed. In particular, roles of sugar sensing, redox control and of cytokinin are highlighted. Finally, the development of a tree is considered. CONCLUSIONS: Various mechanisms contribute to construction and maintenance of efficient photosynthetic systems. Molecular backgrounds of these ecologically important mechanisms should be clarified. The construction mechanisms of the tree cannot be explained solely by the nitrogen optimization theory. It is proposed that the pipe model theory in its differential form could be a potential tool in future studies in this research area.     

Biomass allocation and leaf chemical defence in defoliated seedlings of Quercus serrata with respect to carbon-nitrogen balance

BACKGROUND AND AIMS: Both nutrient availability and defoliation affect the carbon-nutrient balance in plants, which in turn influences biomass allocation (e.g. shoot-to-root ratio) and leaf chemical composition (concentration of nitrogen and secondary compounds). In this study it is questioned whether defoliation alters biomass allocation and chemical defence in a similar fashion to the response to nutrient deficiency. METHODS: Current-year seedlings of Quercus serrata were grown with or without removal of all leaves at three levels of nutrient availability. KEY RESULTS: Plant nitrogen concentration (PNC), a measure of the carbon-nutrient balance in the plant, significantly decreased immediately after defoliation because leaves had higher nitrogen concentrations than stems and roots. However, PNC recovered to levels similar to or higher than that of control plants in 3 or 6 weeks after the defoliation. Nitrogen concentration of leaves produced after defoliation was significantly higher than leaf nitrogen concentration of control leaves. Leaf mass per plant mass (leaf mass ratio, LMR) was positively correlated with PNC but the relationship was significantly different between defoliated and control plants. When compared at the same PNC, defoliated plants had a lower LMR. However, the ratio of the leaf to root tissues that were newly produced after defoliation as a function of PNC did not differ between defoliated and control plants. Defoliated plants had a significantly lower concentration of total phenolics and condensed tannins. Across defoliated and control plants, the leaf tannin concentration was negatively correlated with the leaf nitrogen concentration, suggesting that the amount of carbon-based defensive compounds was controlled by the carbon-nutrient balance at the leaf level. CONCLUSIONS: Defoliation alters biomass allocation and chemical defence through the carbon-nutrient balance at the plant and at the leaf level, respectively.     

Effects of water stress and high nocturnal temperature on photosynthesis and nitrogen level of a perennial grass Leymus chinensis

Water deficit and high temperature are important environmental factors restricting plant growth and photosynthesis. The two stresses often occur simultaneously, but their interactions on photosynthesis and nitrogen level have been less studied. In the present experiment, we measured photosynthetic parameters, stomatal density, and nitrogen levels, as well as soluble sugar content of leaves of a perennial grass, Leymus chinensis, experiencing two day/night temperature regimes of 30/20 °C and 30/25 °C, and five different soil moisture contents (the soil relative-water content ranged from 80% to 25%). Leaf relative water content, leaf biomass, whole plant biomass, the ratio between the leaf biomass and total plant biomass, and the photosynthetic rate, as well as water-use efficiency decreased at high night temperature, especially under severe water stress conditions. Stomatal index was also increased by soil water stress except very severe water stress, and high nocturnal temperature decreased the leaf stomatal index under soil water stress. Nocturnal warming decreased nitrogen concentration in the leaves and increased it in the roots, particularly when plants were subjected to severe water stress. There were significant positive correlations between the photosynthetic rate and both soluble sugar concentration and nitrogen concentration at low nocturnal temperature. It is suggested that nocturnal warming significantly exacerbates the adverse effects of soil water stress, and their synergistic interactions might reduce the plant productivity and constrain its distribution in the region dominated by L. chinensis, based on predictions of global climate change.     

Variations in the relationship between maximum leaf carboxylation rate and leaf nitrogen concentration

叶片最大羧化速率是表征植物光合能力的关键参数, 受到光照、温度、水分、CO₂浓度、叶片氮含量等多个要素的控制。准确地模拟植物叶片最大羧化速率对环境因子的响应是预测未来植被生产力和碳循环过程的前提。目前大多数陆地碳循环过程模型以Farqhuar光合作用模型为基础模拟植物的光合作用, 关于植物叶片的最大羧化速率与叶氮含量关系的模拟方法却各不相同。该文汇总了1990-2013年国内外植物叶片光合速率观测研究文献中叶片最大羧化速率与叶氮含量的关系式及相关数据, 分析了叶片最大羧化速率与叶氮含量关系随不同植被功能型和时间的变化特征, 以及环境因子变化条件下最大羧化速率与叶氮含量关系的变化特征, 探讨了二者关系变异性的可能原因以及影响因子。结果表明: 1)不同功能型植物叶片的最大羧化速率和叶氮含量的关系存在较大差异, 二者线性关系式的斜率平均值变化范围为16.29-50.25 μmol CO₂·g N^-1·s^-1。落叶植被叶片的最大羧化速率随叶氮含量的变化率和光合氮利用效率一般都高于常绿植被, 其变异主要源于植物的比叶重和叶片内部氮素分配的差异。2)叶片最大羧化速率随叶氮含量的变化存在季节和年际变异。在没有受到水分胁迫的年份中, 叶片最大羧化速率随叶氮含量变化的速率一般在春季或夏季最高, 其季节变异与比叶重和叶氮在Rubisco的分配比例的季节变化有关。受到干旱的影响, 叶片最大羧化速率随叶氮含量的变化率会升高。3)当大气CO₂浓度增加时, 由于叶片中Rubisco含量的降低, 多年生针叶叶片最大羧化速率和叶氮关系斜率值会出现降低; 当供氮水平增加时, 叶片最大羧化速率和叶片氮含量均表现出增加趋势, 二者线性关系的斜率也相应增加。在此基础上, 该文指出在模拟叶片最大羧化速率与叶氮含量的关系时, 应考虑叶片比叶重和叶氮在Rubisco中的分配比例的季节变异、水分胁迫、大气CO₂浓度和供氮水平变化对二者关系的影响。囿于数据的有限性, 今后应进一步加强多因子控制实验研究, 深入探讨叶片最大羧化速率与叶氮含量关系的变异性机理, 并获得更系统的观测数据, 以助生态系统过程模型的改进, 提高模型的模拟精度。     

植物叶片最大羧化速率与叶氮含量关系的变异性

叶片最大羧化速率是表征植物光合能力的关键参数, 受到光照、温度、水分、CO₂浓度、叶片氮含量等多个要素的控制。准确地模拟植物叶片最大羧化速率对环境因子的响应是预测未来植被生产力和碳循环过程的前提。目前大多数陆地碳循环过程模型以Farqhuar光合作用模型为基础模拟植物的光合作用, 关于植物叶片的最大羧化速率与叶氮含量关系的模拟方法却各不相同。该文汇总了1990-2013年国内外植物叶片光合速率观测研究文献中叶片最大羧化速率与叶氮含量的关系式及相关数据, 分析了叶片最大羧化速率与叶氮含量关系随不同植被功能型和时间的变化特征, 以及环境因子变化条件下最大羧化速率与叶氮含量关系的变化特征, 探讨了二者关系变异性的可能原因以及影响因子。结果表明: 1)不同功能型植物叶片的最大羧化速率和叶氮含量的关系存在较大差异, 二者线性关系式的斜率平均值变化范围为16.29-50.25 μmol CO₂·g N^-1·s^-1。落叶植被叶片的最大羧化速率随叶氮含量的变化率和光合氮利用效率一般都高于常绿植被, 其变异主要源于植物的比叶重和叶片内部氮素分配的差异。2)叶片最大羧化速率随叶氮含量的变化存在季节和年际变异。在没有受到水分胁迫的年份中, 叶片最大羧化速率随叶氮含量变化的速率一般在春季或夏季最高, 其季节变异与比叶重和叶氮在Rubisco的分配比例的季节变化有关。受到干旱的影响, 叶片最大羧化速率随叶氮含量的变化率会升高。3)当大气CO₂浓度增加时, 由于叶片中Rubisco含量的降低, 多年生针叶叶片最大羧化速率和叶氮关系斜率值会出现降低; 当供氮水平增加时, 叶片最大羧化速率和叶片氮含量均表现出增加趋势, 二者线性关系的斜率也相应增加。在此基础上, 该文指出在模拟叶片最大羧化速率与叶氮含量的关系时, 应考虑叶片比叶重和叶氮在Rubisco中的分配比例的季节变异、水分胁迫、大气CO₂浓度和供氮水平变化对二者关系的影响。囿于数据的有限性, 今后应进一步加强多因子控制实验研究, 深入探讨叶片最大羧化速率与叶氮含量关系的变异性机理, 并获得更系统的观测数据, 以助生态系统过程模型的改进, 提高模型的模拟精度。     

Effects of root pruning of wheat at different nitrogen levels

In two experiments, wheat plants growing in solutions of different nitrogen concentration were subjected to root pruning. In higher concentrations of nitrogen the growth rate was higher, and the proportional allocation of growth to shoot higher, but pruning did not affect the allocation of growth at either level of nitrogen. This result gives no support to Thornley's source-sink model of the control of shoot: root ratio.     

全球变化下植物的碳氮关系及其环境调节研究进展——从分子到生态系统

在全球变化条件下,温度的升高和降水格局的变化,导致淡水资源更加匮乏。环境因子胁迫,如干旱和高温等,它们单独或联合的作用将导致作物大幅度减产,引发自然生态系统退化。植物的碳氮代谢及其分配相互联系、不可分割,其生物过程及外界环境调节共同决定着植物的净生产力和营养水平。该文试图从分子、组织、器官、个体和生态系统等层面上,就植物的碳氮关系及其环境调节(温度、水分和CO₂浓度等)进行综述,并提出了进一步展开相关研究应重点关注的几个方面。     

植物叶片氮分配及其影响因子研究进展

氮是植物生长的基本限制性因子,它的有效利用可以增加植物的适应性。叶片氮分配是指氮在植物叶片细胞各细胞结构以及游离化合物中所分配的比例。叶片氮的分配方式决定了叶片光合作用的强弱,影响叶片的坚韧程度以及化学防御强度,因此研究氮在植物叶片内的分配方式具有重要意义。阐述了叶片氮分配的方式,分析了影响叶片氮分配的生物和非生物因子(CO2,光,土壤养分),介绍了常用的叶片氮分配的研究方法,并对未来的研究进行了展望。     

Leaf canopy as a dynamic system: ecophysiology and optimality in leaf turnover

BACKGROUND AND AIMS: In a leaf canopy, there is a turnover of leaves; i.e. they are produced, senesce and fall. These processes determine the amount of leaf area in the canopy, which in turn determines canopy photosynthesis. The turnover rate of leaves is affected by environmental factors and is different among species. This mini-review discusses factors responsible for leaf dynamics in plant canopies, focusing on the role of nitrogen. SCOPE: Leaf production is supported by canopy photosynthesis that is determined by distribution of light and leaf nitrogen. Leaf nitrogen determines photosynthetic capacity. Nitrogen taken up from roots is allocated to new leaves. When leaves age or their light availability is lowered, part of the leaf nitrogen is resorbed. Resorbed nitrogen is re-utilized in new organs and the rest is lost with dead leaves. The sink-source balance is important in the regulation of leaf senescence. Several models have been proposed to predict response to environmental changes. A mathematical model that incorporated nitrogen use for photosynthesis explained well the variations in leaf lifespan within and between species. CONCLUSION: When leaf turnover is at a steady state, the ratio of biomass production to nitrogen uptake is equal to the ratio of litter fall to nitrogen loss, which is an inverse of the nitrogen concentration in dead leaves. Thus nitrogen concentration in dead leaves (nitrogen resorption proficiency) and nitrogen availability in the soil determine the rate of photosynthesis in the canopy. Dynamics of leaves are regulated so as to maximize carbon gain and resource-use efficiency of the plant.     

On the vegetative biomass partitioning of seed plant leaves, stems, and roots

A central goal of comparative life-history theory is to derive the general rules governing growth, metabolic allocation, and biomass partitioning. Here, we use allometric theory to predict the relationships among annual leaf, stem, and root growth rates (GL, GS, and GR, respectively) across a broad spectrum of seed plant species. Our model predicts isometric scaling relationships among all three organ growth rates: GL is proportional to GS is proportional to GR. It also provides a conceptual basis for understanding the differences in the absolute amounts of biomass allocated to construct the three organ types. Analyses of a large compendium of biomass production rates across diverse seed plant species provide strong statistical support for the predictions of the theory and indicate that reproductive investments may scale isometrically with respect to vegetative organ growth rates. The general rules governing biomass allocation as indexed by the scaling exponents for organ growth rates are remarkably indifferent to plant size and taxonomic affiliation. However, the allometric "constants" for these relationships differ numerically as a function of phenotypic features and local environmental conditions. Nonetheless, at the level of both inter- and intraspecific comparisons, the same proportional biomass allocation pattern holds across extant seed plant species.     

植物代谢速率与个体生物量关系研究进展

植物的各项生理生态功能（例如,呼吸、生长和繁殖）都与个体生物量成异速生长关系。West, Brown及Enquist基于分形网络结构理论所提出的WBE模型认为：植物的代谢（呼吸）速率正比于个体生物量的3/4次幂。然而,恒定的“3/4异速生长指数”与实测数据、植物生理生态学等研究之间存在矛盾,引发激烈的争论。论文分析了不同回归方法对代谢指数的影响,重点对植物代谢速率与个体生物量异速生长关系研究进展进行了综述,分析并得出了植物代谢指数在小个体时接近1.0,并随着生物量的增加而系统减小,且其密切依赖于氮含量的调控的结论。据此,提出了进一步深入研究植物代谢速率个体生物量关系需要解决的一些科学问题。     

A model of the long-term response of carbon allocation and productivity of forests to increased CO2concentration and nitrogen deposition

We present a simple theoretical analysis of the long term response of forest growth and carbon allocation to increased atmospheric [CO₂] and N deposition. Our analysis is based on a recent model which predicts that plant light-use efficiency increases with [CO₂] but is independent of plant N supply. We combine that model with simple assumptions for nitrogen fluxes in the soil. A quasi-equilibrium analysis of the short term tree and soil pools is then used to develop a simple graphical depiction of the long term carbon and nitrogen supply constraints on total growth, stem growth and foliar allocation. Our results suggest that long-term growth responses to [CO₂] and N deposition depend strongly on the extent to which stem allocation and foliage allocation are coupled. At one extreme (‘no coupling’), when stem allocation is fixed and independent of foliage allocation, there is no response of total growth or stem growth to increased [CO₂] unless N deposition increases. At the other extreme (‘linear coupling’), when stem allocation is proportional to foliage allocation, there is a significant long-term increase in total growth following a doubling of [CO₂], even when N deposition is unchanged, but stem growth decreases because of a long-term decrease in foliage allocation. For both types of coupling, total growth and stem growth increase with increasing N deposition. In the case of linear coupling, however, the N deposition response of stem growth is significantly larger than that of total growth, because of a long-term increase in foliage allocation. We compare our results with those obtained previously from an alternative model of canopy light-use efficiency involving a dependence on the foliar N:C ratio in addition to [CO₂]. Our results highlight the need for more experimental information on (i) the extent to which canopy light-use efficiency is independent of N supply, and (ii) the relationship between foliage allocation and stem allocation.     

Optimization of Plant Root: Shoot Ratios and Internal Nitrogen Concentration

A general theoretical approach is developed to analyze the morphologicaland physiological responses of plants to nitrogen availability.The optimal leaf-nitrogen concentration and corresponding optimalroot: shoot ratio which maximize relative growth rate are foundquantitatively as a function of root—specific activitywhich is assumed to be a function of soil nitrogen availability.The cost of increasing tissue nitrogen concentration is foundto be primarily related to an increase in allocation to roots.Predictions of the analysis are consistent with previous theoriesand general empirical findings, suggesting that plants respondoptimally to soil nitrogen. Relative growth rate is predictedto be a nearly linear function of whole-plant nitrogen concentrationand shoot fraction is a monotonically increasing function oftissue nitrogen concentration when plants respond optimallyto soil nitrogen availability. Plant growth, root:shoot ratios, biomass allocation, nitrogen productivity, optimization     

Growth and nitrogen utilization in seedlings of mountain birch (Betula pubescens ssp. tortuosa ) as affected by ultraviolet radiation (UV-A and UV-B) under laboratory and outdoor conditions

 Growth patterns and nitrogen economy were studied in pot-grown seedlings of mountain birch subjected to different ultraviolet radiation under both laboratory and outdoor conditions at Abisko in northern Sweden. In the laboratory, nutrient supply, temperature, humidity, ultraviolet radiation-A (UV-A, 320–400 nm) and B (UV-B, 280–320 nm) were controlled, while photosynthetically active radiation (PAR, 400–700 nm) and photoperiod varied naturally. Under outdoor conditions nutrient supply was controlled, and the irradiation treatments were ambient and above-ambient UV-B using additional fluorescent lamps. Mountain birch nitrogen economy was affected by increased ultraviolet radiation, as reflected by a changed relationship between plant growth and plant nitrogen both in the laboratory and outdoors. In the laboratory enhanced UV-A decreased leaf area per unit plant biomass (leaf area ratio) but increased biomass productivity, both per unit leaf area (leaf area productivity) and per unit leaf nitrogen (leaf nitrogen productivity). Low levels of UV-B affected growth patterns and nitrogen economy in a similar way to enhanced UV-A. High levels of UV-B clearly decreased relative growth rate and nitrogen productivity, as leaf area ratio, leaf area productivity and leaf nitrogen productivity were all decreased. Under outdoor conditions above-ambient levels of UV-B did not alter growth or biomass allocation traits of the seedlings, whilst nitrogen productivity was increased. Mountain birch seedlings originating from different mother trees varied significantly in their responses to different ultraviolet radiation. Received: 10 April 1997 / Accepted: 19 September 1997     

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.     

A dynamic growth model of vegetative soya bean plants: model structure and behaviour under varying root temperature and nitrogen concentration

A differential equation model of vegetative growth of the soya bean plant (Glycine max (L.) Merrill cv. Ransom') was developed to account for plant growth in a phytotron system under variation of root temperature and nitrogen concentration in nutrient solution. The model was tested by comparing model outputs with data from four different experiments. Model predictions agreed fairly well with measured plant performance over a wide range of root temperatures and over a range of nitrogen concentrations in nutrient solution between 0.5 and 10.0 mmol NO3- in the phytotron environment. Sensitivity analyses revealed that the model was most sensitive to changes in parameters relating to carbohydrate concentration in the plant and nitrogen uptake rate.