Allocation of carbon and nitrogen as a function of the internal nitrogen status of a plant: Modelling allocation under non-steady-state situations

In this paper we model allocation of carbon and nitrogen to roots and leaves as a function of the nitrogen status of a plant. Under steady-state conditions, allocation of carbon and nitrogen to leaves is exponentially (positively) correlated with plant nitrogen concentration, whereas allocation to roots is correlated negatively, also in an exponential manner.Allocation functions derived under steady-state conditions are used to simulate biomass partitioning under non-steady-state nutrient conditions. Upon nitrogen deprivation, measured and simulated values are rather similar with time, suggesting that allocation functions derived under steady-state conditions also hold under non-steady-state conditions.     

Monitoring cellular C:N ratio in phytoplankton by means of FTIR‐spectroscopy

Statistical growth rate modelling can be applied in a variety of ecological and biotechnological applications. Such models are frequently based on Monod or Droop equations and, especially for the latter, require reliable determination of model input parameters such as C:N quotas. Besides growth rate modelling, a C:N quota quantification can be useful for monitoring and interpretation of physiological acclimation to abiotic and biotic disturbances (e.g., nutrient limitations). However, as high throughput C:N quota determination is difficult to perform, alternatives need to be established. Fourier‐transformed infrared (FTIR) spectroscopy is used to analyze a variety of biochemical, chemical, and physiological parameters in phytoplankton. Hence, a quantification of the C:N quota should also be feasible. Therefore, using FTIR spectroscopy, six phytoplankton species from among different phylogenetic groups have been analyzed to determine the effect of nutrient limitation on C:N quota patterns. The typical species‐specific response to increasing nitrogen limitation was an increase in the C:N quota. Irrespective of this species specificity, we were able to develop a reliable multi‐species C:N quota prediction model based on FTIR spectroscopy using the partial least square regression (PLSR) algorithm. Our data demonstrate that the PLSR approach is more robust in C:N quota quantification (R² = 0.93) than linear correlation of C:N quota versus growth rate (R² ranges from 0.74 to 0.86) or biochemical information based on FTIR spectra (R² ranges from 0.82 to 0.89). This accurate prediction of C:N values may support high throughput measurements in a broad range of future approaches.     

Carbon and nitrogen allocation to male and female reproduction in Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca, Pinaceae)

We measured carbon (respiration, photosynthesis, and production) and nitrogen allocation to male and female cones of Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca) to quantify gender-specific: (1) resource allocation to reproduction, and (2) contribution to carbon costs of reproduction via photosynthesis. We also measured foliar photosynthesis and nitrogen concentration ([N]) near and far from female cones to examine the relationship between reproduction and foliar physiology. Over one growing season, male cones required only 8% of all carbon allocated to reproduction, with females consuming the remaining 92%. Female cones, however, had maximum instantaneous refixation rates of 54%, which, integrated over the season, offset 6% of their total carbon requirements, while male cones were completely dependent on vegetative tissues for carbon. Male cones received 22% of all nitrogen allocated to reproduction and female cones received the remaining 78%. Foliage near female cones had elevated photosynthesis during the early stages of cone development and consistently lower [N] than foliage far from cones. Although female cones may photosynthesize, the annual sum of carbon fixed by reproductive structures is minor in comparison to the total carbon allocated to production and respiration.     

Optimal nitrogen distribution within a leaf canopy under direct and diffuse light

Nitrogen distribution within a leaf canopy is an important determinant of canopy carbon gain. Previous theoretical studies have predicted that canopy photosynthesis is maximized when the amount of photosynthetic nitrogen is proportionally allocated to the absorbed light. However, most of such studies used a simple Beer's law for light extinction to calculate optimal distribution, and it is not known whether this holds true when direct and diffuse light are considered together. Here, using an analytical solution and model simulations, optimal nitrogen distribution is shown to be very different between models using Beer's law and direct–diffuse light. The presented results demonstrate that optimal nitrogen distribution under direct–diffuse light is steeper than that under diffuse light only. The whole‐canopy carbon gain is considerably increased by optimizing nitrogen distribution compared with that in actual canopies in which nitrogen distribution is not optimized. This suggests that optimization of nitrogen distribution can be an effective target trait for improving plant productivity.     

我国南亚热带几种人工林生态系统碳氮储量

以我国南亚热带格木(Erythrophleumfordii)、红椎(Castnopsis hystrix)和马尾松(Pinus massoniana)人工林为研究对象,对其碳氮储量及分配格局进行研究.结果表明,不同树种体内碳的分布与器官年龄的关系不明显,而氮的分布与年龄的关系则较为密切,表现为幼嫩器官中的氮含量大于老化器官,而老化器官的C/N比值大于幼嫩器官.格木人工林生态系统内各组分的氮含量均高于其他两种人工林生态系统,并且3种人工林生态系统碳氮在土壤表层具有明显的富集作用.格木、红椎和马尾松人工林碳储量分别为236.22、267.84、200.57 t/hm2,氮储量分别为17.91、12.38、10.59 t/hm2.乔木层碳储量分别占42.57％、36.31％和40.28％,0-100 cm土壤碳储量分别占55.77％、62.52％和57.83％;氮储量则是土壤占绝对优势,分别为92.00％、93.72％和95.53％.说明3种人工林生态系统碳氮储量主要分布在土壤中,且红椎人工林生态系统具有较高的固碳能力.     

Carbon fixation and carbon availability in marine phytoplankton

It is widely believed that inorganic C does not limit the rate of short-term photosynthesis, the net productivity, or the maximum biomass, of marine phytoplankton. This lack of inorganic C restriction is less widely believed to hold for phytoplankton in many low alkalinity freshwaters or for seaweed in nutrient-enriched rock pools. These views are examined in the context of the physical chemistry of the inorganic C system in natural waters and of the ways in which various taxa of phytoplankton deal with inorganic C and discriminate between ¹²C and ¹³C. Using this information to interpret data obtained in the ocean or in freshwater suggests that short-term photosynthesis, production rate, and achieved biomass, of phytoplankton are rarely limited by inorganic C supply but, rather, that the widely suggested factors of limited light, nitrogen or phosphorus supply are the resource inputs which restrict productivity. Global change, by increasing atmospheric CO₂ partial pressure and global mean temperatures, is likely to increase the mean CO₂ concentration in the atmosphere, but the corresponding change in the oceans will be much less. There are, however, genotypic differences in the handling of inorganic C among the diversity of marine phytoplankton, and in impact on use of limiting nutrients, so increases in the mean CO₂ and HCO₃ ^- concentrations in surface ocean waters could cause changes in species composition. However, the rarity of inorganic C limitation of marine phytoplankton short-term photosynthesis, net productivity, or the maximum biomass, in today's ocean means that global change is unlikely to increase these three values in the ocean.     

Kinetics of intracellular carbon allocation in a marine diatom

To test models of intracellular carbon flow we measured the labelling kinetics (from ¹⁴CO₂) of major classes of cell polymers (carbohydrate, protein, lipid) and of dissolved organic carbon produced by the marine diatom Thalassiosira pseudonana Hustedt, grown at rates of 0.2 to 2.0·day^?1 under nitrogen or light limitation. Compartmental analysis indicated that tracer carbon quickly entered respiratory and excretory streams, accumulating in the cells at the rate of net production after only 25–50% of cell generation (doubling) time. Respiration rates were low (≤ 0.1 · day^?1) and suggested that illuminated cells in steady-state growth made only minor use of oxidative respiration to support cell synthesis. The tracer was quick to enter all polymers; compartmental analysis indicated that polymer labelling rates were close to the rates of mass synthesis after several hours of incubation with ¹⁴C. Polymer labelling also showed a reallocation of photosynthate from protein to carbohydrate within a few hours of perturbation (shift-down) of nutrient supply in a N-limited chemostat. In steady-state growth, the protein: carbohydrate ratio increased directly with N-limited growth rate but attained its maximum under extreme light-limitation. Carbon flow into the metabolic processes of respiration, excretion and polymer synthesis appeared to be mediated by a small and rapidly cycled pool of substrates under all steady-state growth conditions.     

Redfield revisited: variability of C:N:P in marine microalgae and its biochemical basis

Cytological events immediately following plasmogamy in Durvillaea potatorum are described. Eggs contain several types of cytoplasmic vesicle differing in size and appearance. Histochemical tests and measurements are used to characterise and distinguish different types of vesicle containing phenolic compounds, lipids and polysaccharides. Within 2 min of plasmogamy, small phenolic vesicles located just below the egg membrane undergo mass synchronised exocytosis. The contents of these vesicles are discharged as phenolic bodies on the outside of the membrane. Secretion of phenolic bodies precedes secretion of the primary zygote wall by several minutes. Limited secretion of phenolics also occurs in unfertilised eggs. Peripheral phenolic vesicles are distinguishable from physodes, which also contain phenolic compounds but which are significantly larger and tend to be localised around the egg nucleus. The possible functional significance of the phenolic bodies is discussed. Coated pits and vesicles are common in zygotes, and their presence is evidence for endocytosis.     

水曲柳和落叶松人工林乔木层碳、氮储量及分配

采用树木解析和连续土芯法,估测了20年生水曲柳和落叶松人工林乔木层各部分生物量和生产量,以及两种林分各部分的碳、氮含量及储量.结果表明:水曲柳和落叶松乔木林生物量分别为6815.10和9295.95 g·m^-2;两树种树干生物量占总生物量的比例均最高,分别为57.32%和58.01%;细根生物量最低,分别为2.67%和1.80%.水曲柳和落叶松的年生产量分别为1618.16和2102.45 g·m^-2·a^-1,其中树干年生产量最高,分别占总生物量的39.34%和46.70%;细根的年生产量较低,分别占总生物量的12.06%和5.25%.水曲柳各器官碳含量低于落叶松,氮含量则高于落叶松;水曲柳林碳储量低于落叶松,而两树种氮储量差别不大.水曲柳分配到地上部分的生物量、生产量以及碳、氮比例均小于落叶松,反映了落叶松在构建地上部分相对于水曲柳的高效性;由于树种之间以及同一树种不同器官之间的碳、氮含量差别显著,精确估计森林碳、氮储量时应分树种和器官进行测定.     

施氮对几种草地植物生物量及其分配的影响

为了研究施氮对不同草地植物生物量及其分配的影响,以及温带草地生态系统碳交换过程对氮素的响应,在内蒙古太仆寺旗农田-草地生态系统野外站,以4种草地植物:紫花苜蓿(Medicago sativa L.)、高丹草(Sorghum bicolor L.) 、羊草(Leymus chinensis T.)和小叶锦鸡儿(Caragana microphylia L.)为材料,进行了3种氮素水平 的盆栽控制实验。研究结果表明:施氮显著促进了4种植物地上生物量的积累,紫花苜蓿在中氮水平地上生物量最大,较对照增加了24.8%,高丹草、羊草、小叶锦鸡儿在高氮水平地上生物量最大,分别较对照增加了45.6%、39.3%和72.2%。4种植物在中氮水平地下生物量最大,而细根(直径≤2mm)生物量随施氮量的增加显著减少。羊草根茎生物量及其分配比例随施氮量的增加而增大。施氮显著降低了4种植物的根冠比,紫花苜蓿的根冠比在中氮水平时最小,为1.62,高丹草、羊草、小叶锦鸡儿的根冠比在高氮水平时最小,分别为0.57、1.02和0.41。随施氮量的增加,植物地下部分特别是细根的分配比例显著降低,地上部分分配比例显著增加。不同植物对施氮水平的响应不同,相比豆科植物,施氮显著促进禾本科植物生物量积累,并使其生物量分配格局发生显著改变。     

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.     

Carbon and nitrogen allocation shifts in plants and soils along aridity and fertility gradients in grasslands of China

Plant carbon (C) and nitrogen (N) stoichiometry play an important role in the maintenance of ecosystem structure and function. To decipher the influence of changing environment on plant C and N stoichiometry at the subcontinental scale, we studied the shoot and root C and N stoichiometry in two widely distributed and dominant genera along a 2,200‐km climatic gradient in China's grasslands. Relationships between C and N concentrations and soil climatic variables factors were studied. In contrast to previous theory, plant C concentration and C:N ratios in both shoots and roots increased with increasing soil fertility and decreased with increasing aridity. Relative N allocation shifted from soils to plants and from roots to shoots with increasing aridity. Changes in the C:N ratio were associated with changes in N concentration. Dynamics of plant C concentration and C:N ratios were mainly caused by biomass reallocation and a nutrient dilution effect in the plant‐soil system. Our results suggest that the shifted allocation of C and N to different ecosystem compartments under a changing environment may change the overall use of these elements by the plant‐soil system.     

The impact of irradiance on optimal and cellular nitrogen to phosphorus ratios in phytoplankton

Phytoplankton acclimates to irradiance by regulating the cellular content of light‐harvesting complexes, which are nitrogen (N) rich and phosphorus (P) poor. Irradiance is thus hypothesised to influence the cellular N : P ratio and the N : P defining the threshold between N and P limitation (the ‘optimal’ N : P). We tested this hypothesis by first addressing the response of the optimal N : P to irradiance in a controlled experiment with Chlamydomonas reinhardtii. Then, we did a meta‐analysis of experimental data on optimal and cellular N : P ratios across light gradients to test the generality of an N : P to light response within species. In both the experiment and the meta‐analysis, N : P ratios decreased with irradiance, indicating that factors affecting underwater irradiance, like depth and the composition of the water, may influence the relative N : P requirement. The effect of irradiance did not differ between optimal and cellular N : P ratios, but observations of optimal N : P were on average 2.8 times higher than observations of cellular N : P.     

Assimilation and allocation of carbon and nitrogen of thermal and nonthermal Agrostis species in response to high soil temperature

We studied whether changes in the assimilation and allocation of carbon and nitrogen are associated with plant tolerance to high soil temperatures. Two Agrostis species, thermal Agrostis scabra, a species adapted to high-temperature soils in geothermal areas in Yellowstone National Park (USA), and two cultivars of a cool-season species, Agrostis stolonifera, L-93 and Penncross, were exposed to soil temperatures of 37 or 20 degrees C, while shoots were exposed to 20 degrees C. Net photosynthesis rate, photochemical efficiency, NO(3) (-)-assimilation rate and root viability decreased with increasing soil temperatures in both species. However, the decreases were less pronounced for A. scabra than for both A. stolonifera cultivars. Carbon investment in growth of plants exposed to 37 degrees C decreased more dramatically in both A. stolonifera cultivars than in A. scabra. Nitrogen allocation to shoots was greater in A. scabra than in both creeping bentgrass cultivars at 37 degrees C soil temperature. Our results demonstrate that plant tolerance to high soil temperature is related to efficient expenditure and adjustment of C- and N-allocation patterns between growth and respiration.     

On evolution of andromonoecy and 'overproduction' of flowers: a resource allocation model

The conditions for the evolution of andromonoecy and male-function-controlled overproduction of fertile flowers in hermaphrodites are considered using the evolutionarily stable strategy (ESS) approach. Andromonoecy and male-function-controlled floral display are promoted if further increase in pollen amount per flower is disadvantageous and/or increase in the number of polliniferous units is advantageous; the costs of attractive organs and pollen per flower are relatively low while the cost of ovules per flower and the cost per fruit are relatively high; the probability of setting a fruit from a pistillate flower is high; male fertility increases with the resources devoted to flowering; and if selfing is relatively low. Assumptions and predictions of the model are discussed.     

四川巴郎山齿果酸模叶片氮素及其分配的海拔响应

在卧龙自然保护区, 按海拔梯度选择了齿果酸模(Rumex dentatus)的4个分布地点(2350、2700、3150和3530 m), 对各研究地点的齿果酸模进行了叶片光合、扩散导度、叶片碳稳定同位素组成(δ¹³C)、氮素含量、光合氮利用效率(PNUE)、比叶面积(SLA))等参数的测量, 以期揭示该植物叶片氮素、氮素分配情况及其他生理生态参数随海拔的响应趋势, 进而明确氮素及其分配在齿果酸模响应和适应海拔梯度环境的生物学过程中的作用。结果表明: 随着海拔的升高, 齿果酸模的叶片单位面积氮含量(N_area)随之增加, 进而光合能力随之增加。随着海拔升高而增加的扩散导度也在一定程度上促进了这一趋势, 这可能是落叶草本植物对于高海拔低温所导致的叶寿命缩短的适应结果。沿着海拔梯度, 植物叶片氮素和扩散导度均通过羧化位点与外界CO₂分压比(P_c/P_a)而间接影响叶片δ¹³C值, 且相比之下, 以氮素为基础的羧化能力对于P_c/P_a的作用更大些, 进而导致齿果酸模叶片δ¹³C随海拔增加; 随着海拔的升高, 齿果酸模叶片将更多的氮素用于防御性结构组织的建设, 这也是SLA和PNUE降低的主要原因; 在光合系统内部, 随着海拔的升高, 植物光合组织增加了用于捕光系统氮素的比例, 使得植物可以更好地利用随海拔升高而增强的光照资源, 进而促进了光合能力的增加。可见, 氮素及其在叶片各系统间(尤其是在光合系统与非光合系统间)的分配方式是齿果酸模适应和响应海拔梯度环境的关键。     

Herbivore-induced shifts in carbon and nitrogen allocation in red oak seedlings

* A dual-isotope, microcosm experiment was conducted with Quercus rubra (red oak) seedlings to test the hypothesis that foliar herbivory would increase belowground carbon allocation (BCA), carbon (C) rhizodeposition and nitrogen (N) uptake. Plant BCA links soil ecosystems to aboveground processes and can be affected by insect herbivores, though the extent of herbivore influences on BCA is not well understood in woody plants. * Microcosms containing 2-yr-old Q. rubra seedlings and soil collected from the Coweeta Hydrologic Laboratory (NC, USA) were subjected to herbivory or left as undamaged controls. All microcosms were then injected with 15N-glycine and pulsed with 13CO2. * Contrary to our hypothesis, herbivore damage reduced BCA to fine roots by 63% and correspondingly increased allocation of new C to foliage. However, 13C recoveries in soil pools were similar between treatments, suggesting that exudation of C from roots is an actively regulated component of BCA. Herbivore damage also reduced N allocation to fine roots by 39%, apparently in favor of storage in taproot and stem tissues. * Oak seedlings respond to moderate insect herbivore damage with a complex suite of allocation shifts that may simultaneously increase foliar C, maintain C rhizodeposition and N assimilation, and shift N resources to storage.     

东北东部山区11种温带树种粗木质残体分解与碳氮释放

采用长期定位跟踪实测方法,比较分析了我国东北温带森林4个水热状况不同的立地条件(红松(Pinus koraiensis)人工林、硬阔叶林、蒙古栎(Quercus mongolica)林和林外空旷地)下11个温带树种粗木质残体(CWD)分解初期3年中的碳氮动态及其影响因子。测定树种包括:白桦(Betula platyphylla)、山杨(Populus davidiana)、紫椴(Tilia amurensis)、胡桃楸(Juglans mandshurica)、蒙古栎、色木槭(Acer mono)、春榆(Ulmus japonica)、红松、黄檗(Phellodendron amurense)、兴安落叶松(Larix gmelinii)和水曲柳(Fraxinus mandshurica)。结果表明:在分解过程中,所有树种CWD的碳浓度没有明显变化(p0.05),但其干重、碳密度、氮浓度和氮密度均随分解进程不同程度地减小,碳氮比(C/N)则增大,而且树种间差异显著(p0.001)。针叶树种的CWD分解速率显著地低于阔叶树种,其中白桦的3年CWD干重损失率(65%)约为兴安落叶松(22%)的3倍。径级大的CWD分解较慢。CWD分解与碳氮释放均与CWD的初始N含量呈正相关,而与初始C/N呈负相关。4个立地条件下CWD的干重和碳氮含量的变化差异不显著,均表现出一致的变化趋势。该研究指出,在分解初期的前3年中,CWD基本上是一个碳源和氮源。