Competition for nutrients between Calluna vulgaris (L.) Hull and Molinia caerulea (L.) Moench

Summary A combination of a removal and fertilization experiment in the field and a replacement experiment in containers in a cold frame was carried out to investigate the role of nutrient supply in the interactions between Calluna vulgaris and Molinia caerulea. It is concluded that the growth of Calluna as well as that of Molinia increased with increasing nutrient availability. However, the increase in biomass of Molinia was much larger than that of Calluna. It is also concluded that increased nutrient availability favours the competitive vigour of Molinia in interaction with Calluna.     

Effects of prescribed burning on plant available nutrients in dry heathland ecosystems

Heathland management is an important tool with which to modify ecosystem impacts caused by atmospheric nutrient deposition. Since changes in nutrient availability as a result of management measures affect the outcomes of heathland succession and species competition, studies on this issue are important from both a nature conservation and management point of view. This study reports the effects of prescribed burning on nutrient availability in dry heathland soils and the nutrient content of the two competing heathland species Calluna vulgaris and Deschampsia flexuosa, with particular reference to N and P. We hypothesise that winter prescribed burning leads to additional N availability, which enhances the importance of P in the context of nutrient limitation in heathland ecosystems. In the nature reserve “Lueneburg Heath” (NW Germany) we examined the availability of nutrients in the humus horizons and in the leachate as well as the relevant C:element ratios in Calluna and Deschampsia before and after a burning experiment. Our results show that prescribed burning resulted in drastically increased NH₄⁺ availability in the O-horizon. We observed only short-term effects (for NO₃⁻, PO₄³⁻, Mg) and insignificant effects on the availability of other nutrients (K, Ca). As a consequence of an increased nutrient availability in the humus horizons and a limited nutrient uptake by plants after burning, leaching increased significantly for N, Ca, K, and Mg after burning treatment. No significant changes were found in the foliar C:N ratios for either species after prescribed burning, although Deschampsia showed an increased deficiency for all the other nutrients, particularly for P, as expressed by increased foliar C:P and N:P ratios. By contrast, the nutrient content of Calluna did not change significantly, suggesting that prescribed burning favours the competitive capacity of Calluna as against Deschampsia. We assume that water shortage as a result of changes in the microclimate was mainly responsible for the deterioration of the nutrient content of Deschampsia. This gives Calluna a competitive advantage, enabling it to out-compete Deschampsia on burned heathlands, with respect to the key factor P-limitation.     

浙江天童常绿阔叶林、常绿针叶林与落叶阔叶林的C:N:P化学计量特征

以浙江天童常绿阔叶林、常绿针叶林和落叶阔叶林为对象, 通过对叶片和凋落物C:N:P比率与N、P重吸收的研究, 揭示3种植被类型N、P养分限制和N、P重吸收的内在联系。结果显示: 1)叶片C:N:P在常绿阔叶林为758:18:1, 在常绿针叶林为678:14:1, 在落叶阔叶林为338:11:1; 凋落物C:N:P在常绿阔叶林为777:13:1, 常绿针叶林为691:14:1, 落叶阔叶林为567:14:1; 2)常绿阔叶林和常绿针叶林叶片与凋落物C:N均显著高于落叶阔叶林; 叶片C:P在常绿阔叶林最高, 常绿针叶林中等, 落叶阔叶林最低, 常绿阔叶林和常绿针叶林凋落物C:P显著高于落叶阔叶林; 叶片N:P比也是常绿阔叶林最高、常绿针叶林次之, 落叶阔叶林最低, 但常绿阔叶林凋落物N:P最低; 3)植被叶片N、P含量间(N为x, P为y)的II类线性回归斜率显著大于1 (p < 0.05), 表明叶片P含量的增加可显著提高叶片N含量; 凋落物N、P含量的回归斜率约等于1, 反映了凋落物中单位P含量与单位N含量间的等速损耗关系; 4)常绿阔叶林N重吸收率显著高于常绿针叶林与落叶阔叶林, 落叶阔叶林P重吸收率显著高于常绿阔叶林和常绿针叶林。虽然植被的N:P指示常绿阔叶林受P限制, 落叶阔叶林受N限制, 常绿针叶林受N、P的共同限制, 但是N、P重吸收研究结果表明: 受N素限制的常绿阔叶林具有高的N重吸收率, 受P限制的落叶阔叶林并不具有高的P重吸收率。可见, 较高的N、P养分转移率可能不是植物对N、P养分胁迫的一种重要适应机制, 是物种固有的特征。     

天童常绿阔叶林演替系列植物群落的N：P化学计量特征

土壤氮磷养分对植物生长的限制性可通过植被的N：P化学计量特征来反映。该研究以常绿阔叶林演替系列为对象,将N：P作为诊断指标,揭示常绿阔叶林次生演替过程中植物群落的N：P化学计量特征和养分限制作用。结果显示：1）物种水平的N：P大小不一,但演替系列总体的变化特征表现出了较高的一致性。2）在群落水平上,次生演替初期的灌草丛N：P极小（7．38）,远远低于14,当演替进入灌丛阶段,N：P显著增高到19．96,在进入演替中期的针叶林（14．29）和针阔混交林（14．21）时,N：P显著下降到14～16之间,演替中后期的木荷（Schima superba）群落（18．77）和栲树（Castanopsis fargesii）群落（20．13）的N：P发生了显著的升高过程。根据以往对N：P临界值的确定,可以认为,常绿阔叶林次生演替初期的植物群落生产力主要受到氮素的限制作用;演替中期的针叶林和针阔混交林主要受氮磷的共同限制,但以氮素的限制作用更为强烈;演替中后期植物群落主要受到土壤磷素的限制作用。     

Effect of nutrient availability on the C, N, and P elemental ratios in the cyanobacterium Microcystis aeruginosa

To clarify whether nutrients limit the growth of Microcystis aeruginosa (Kütz.) Kütz during the growing season in Lake Yogo, we examined the cellular ratios of carbon (C), nitrogen (N), and phosphorus (P) in the populations of M. aeruginosa from August to December 2001. We also measured cellular C, N, and P ratios of M. aeruginosa under batch culture conditions. The cellular levels of N and P of M. aeruginosa in natural population changed more than twofold. The atomic N: C ratio of natural populations of Microcystis fluctuated from 0.11 to 0.26. The atomic P: C ratio fluctuated from 0.0080 to 0.024. The N: C, P: C, and N: P ratios of exponentially growing M. aeruginosa in N-and P-rich medium were 0.19, 0.013, and 15 on average. The growth of M. aeruginosa was suppressed below the N: C ratio of 0.13 under the N-free condition and below the P: C ratio of 0.0026 in the P-free condition. In the natural population, the N: C ratio was low on August 1-2 (0.11) and the P: C ratio was low (less than 0.011) until September. The Microcystis population on August 1-2 was N limited, judging from the results of the culture experiment. In other periods, the population seemed to be supplied with a sufficient amount of N. Although the P: C ratio was low (approximately 0.01) during August and September, it was several times larger than the value of the reduction of growth rate that occurred in culture. P limitation did not occur during the study period. N became more of a limiting factor than P for the formation of blooms of Microcystis. No blooms were observed in August and September, in spite of the increase of cellular levels of N. The formation of Microcystis blooms in Lake Yogo seems to be affected by artificial manipulations such as pumping from Lake Biwa and outflow.     

施肥与刈割对内蒙古草地生态化学计量特征的影响

为明确施肥和刈割对蒙古半干旱草地四种植物化学计量特征的影响,该文于2014年8月以内蒙古半干旱草地的四种植物冷蒿(Artimesia frigida)、菊叶委陵菜(Potentilla tanacetifolia)、羊草(Aneurotepidimu chinense)、扁蓿豆(Melissitus ruthenica)为研究对象,利用方差分析及多重比较分析对照、施氮肥、割草、割草和施氮肥处理下的植物碳、氮、磷含量的差异以及不同处理下碳氮比和氮磷比的变化。结果表明:施氮肥与割草均未影响植物碳含量。施氮肥显著提高了植物氮含量(P<0.05),进而降低了碳氮比(P<0.05),提高了氮磷比(P<0.05)。割草处理后扁蓿豆的氮含量显著提高了0.18%(P<0.05),羊草的氮含量显著降低了0.13%(P<0.05),冷蒿与菊叶委陵菜的氮含量无显著变化。割草显著提高了冷蒿和菊叶委陵菜的磷含量(P<0.05),但是未影响羊草和扁蓿豆的磷含量。该研究证实了氮添加或割草会影响部分草地物种的化学计量特征,但是氮添加或割草对植物化学计量特征的影响存在种间差异。     

Nutrient leaching in dry heathland ecosystems: effects of atmospheric deposition and management

Atmospheric nutrient deposition has contributed to widespread changes in sensitive seminatural ecosystems throughout Europe. For an understanding of underlying processes it is important to quantify input–output flows in relation to ongoing atmospheric inputs and current management strategies. In this study we quantified losses of N, P, Ca, Mg, and K via leaching in heathland ecosystems (Lüneburger Heide, NW Germany) as a function of current deposition rates and different management measures (mowing, prescribed burning, choppering, sod-cutting) which aim to prevent shrub and tree encroachment. Leaching was only moderately related to atmospheric input rates, indicating that leaching was mostly affected by internal turnover processes. Leaching significantly increased for most of the nutrients after the application of management measures, particularly in the choppered and sod-cut plots. However, atmospheric nutrient inputs exceeded leaching outputs for most of the nutrients, even in the plots subjected to management. Despite high deposition rates (20–25 kg N ha⁻¹ year⁻¹), retention of atmospheric N input ranged between 74% and 92% in the control plots. In the treated plots, N retention decreased to 59–80%. However, in the study area mean N leaching in the controls has almost doubled since 1980 and currently amounts to 3.7 kg ha⁻¹ year⁻¹, indicating an early stage of N saturation. Our study provides evidence that leaching did not compensate for atmospheric nutrient deposition, particularly as regards N. Management, thus, will be an indispensable tool for the maintenance of the low-nutrient status as a prerequisite for the long-term preservation of heathland ecosystems.     

天童常绿阔叶林演替系列植物群落的N∶P化学计量特征

 土壤氮磷养分对植物生长的限制性可通过植被的N∶P化学计量特征来反映。该研究以常绿阔叶林演替系列为对象,将N∶P作为诊断指标,揭示 常绿阔叶林次生演替过程中植物群落的N∶P化学计量特征和养分限制作用。结果显示：1)物种水平的N∶P大小不一,但演替系列总体的变化特 征表现出了较高的一致性。2）在群落水平上,次生演替初期的灌草丛N∶P极小(7.38),远远低于14,当演替进入灌丛阶段,N∶P 显著增高到 19.96,在进入演替中期的针叶林(14.29)和针阔混交林(14.21)时,N∶P显著下降到 14～16之间,演替中后期的木荷(Schima superba)群落 (18.77)和栲树(Castanopsis fargesii)群落(20.13)的N∶P发生了显著的升高过程 。根据以往对N∶P临界值的确定,可以认为,常绿阔叶林次 生演替初期的植物群落生产力主要受到氮素的限制作用;演替中期的针叶林和针阔混交林主要受氮磷的共同限制,但以氮素的限制作用更为强 烈;演替中后期植物群落主要受到土壤磷素的限制作用。     

Nitrification in Dutch heathland soils

A survey was conducted over a range of 17 Dutch heathland locations, subdivided into 41 sites dominated by either dwarf-shrubs (Calluna vulgaris or Erica tetralix) or grass species (Deschampsia flexuosa or Molinia caerulea). Among the habitats of the dominant plant species relatively little differences in general soil properties were observed. The P status of Deschampsia sites was relatively high as well as the NO₃ ^–-N concentrations in the 0–10 cm layer (FH included) at the grass-dominated sites. At sites with a dead or degenerating dwarf-shrub vegetation, NH₄ ⁺-N concentrations reached very high levels.Net production of nitrate was observed during incubation of intact 0–10 cm soil cores (FH-layer included) in the laboratory for all sites, even though in some instances, particularly at Calluna and Erica sites, no nitrate was initially measured. Generally, a higher nitrification rate was found for the grass-dominated sites, and for Deschampsia in particular. The net production of nitrate was highly significantly correlated with net N mineralization, being a reasonable predictor of nitrification in a simple regression model (R²=0.47; P<0.001). Net nitrification was also significantly correlated with the NO₃ ^–-N initially present at the start of the growing season (R=0.65; P<0.001) and with the labile organic P content of the soil (R=0.65; P<0.001). By including initial NO₃ ^–-N and labile organic P, together with net N mineralization and pH, in a multiple regression model, net nitrate production could be predicted with a much higher precision (R²=0.75; P<0.001). Although apparent nitrification was not significantly correlated with pH, the latter contributed significantly to the multiple regression equation for the prediction of the former.The influence of the labile organic P pool may act via its positive correlation with microbial biomass, thus more or less reflecting the potential mineralization/nitrifying capacity of a particular site.     

放牧对全球草地生态系统碳氮磷化学计量特征影响的Meta分析

为明确全球尺度下放牧管理措施对草地生态系统碳(C)、氮(N)、磷(P)化学计量特征的影响,提高草地生态系统管理水平,本研究选取国内外83篇中英文文献进行Meta分析,并通过亚组分析探讨了放牧家畜组合(羊单牧、牛单牧和牛羊混牧)和放牧强度(轻度、中度、重度)对草地生态系统叶片、凋落物、根系,以及土壤C、N、P化学计量特征的影响。结果表明: 放牧会显著降低叶片和凋落物C含量、C/N、C/P,增加N、P含量及N/P;显著降低根系和土壤C、N含量,C/P和N/P,增加P含量和C/N。叶片、凋落物化学计量特征变化对牛、羊单独放牧响应更为明显,而根系、土壤化学计量特征变化则对混牧响应更为明显,重度放牧会对草地生态系统化学计量特征产生更大的影响。放牧会降低土壤N含量,增加P含量,表明放牧对草地N、P含量的影响路径不同。进一步研究N、P含量变化对放牧活动不平衡响应机制,将放牧方式、强度的影响纳入草地生态系统预测、管理模型,能够有效提高草地生态系统管理水平。     

GROWTH RATE VARIATION IN THE N:P REQUIREMENT RATIO OF PHYTOPLANKTON1

Theoretical considerations predict that the cell N:P ratio at transition from nitrogen limitation to phosphorus limitation of phytoplankton growth (critical ratio, R_c) varies, as a function of population growth rate. This prediction is confirmed by experimental, data from the literature along with new experimental data for the marine, prymnesiophyte Pavlova lutheri (Droop) Green. R_c passes through a maximum at intermediate growth rates for the three phytoplankton species for which data, are available, but there is significant interspecific variability in its value. There is no theoretical or experimental evidence to support the idea that the ratio of subsistence N and P cell quotas is equal to R_c over the range of growth rates, or that the subsistence quota ratio equals the ratio of the N and P cell quotas minus a storage fraction. Examination of N:P composition ratios can be used to determine which nutrient is limiting, but cannot be used to determine relative growth rates or competitive advantage between species limited by the same nutrient. Growth rates are determined by environmental conditions and by the cell quota of the limiting nutrient, without reference to the cell quota of the non-limiting nutrient.     

Effects of nutrient (N,P, C) enrichment,grazing and depth upon littoral periphyton of a softwater lake

Three field experiments were performed in Lake Lacawac, PA to determine the importance of potentially limiting nutrients relative to other factors (grazing, depth) in structuring shallow water algal periphyton communities. All three experiments measured periphyton growth (as chlorophyll-a, AFDM or biovolumes of the algal taxa) on artificial clay flower pot substrates which released specified nutrients to their outer surfaces.Control of standing crop by nutrient supply rate vs. grazing was examined in Expt. I. Substrates releasing excess N and P, together with one of 4 levels of C (as bicarbonate) were placed either inside or outside exclosures designed to reduce grazer densities. Chlorophyll-a rose from 1.1–25.6 µg.cm^–2, and some dominant taxa (e.g., Oedogonium, Nostoc, Anacystis) were replaced by others (e.g., Scenedesmus, Cryptomonas) as bicarbonate supply increased. Reductions in invertebrate density did not significantly affect chlorophyll-a at any of the nutrient levels.Reasons for the species shift were further evaluated in Expt. II, using a minielectrode to measure the elevation of pH within the periphyton mat through photosynthetic utilization of bicarbonate. The pH adjacent to pots diffusing N, P and large quantities of bicarbonate, and supporting high chlorophyll-a densities of 32 µg cm^–2, averaged 10.0 compared to 6.3 in the water column. Pots diffusing only N and P supported 0.7 µg chlorophyll-a cm^–2 and elevated pH to 8.2. We suspect that bicarbonate addition favored efficient bicarbonate users (e.g., Scenedesmus), while inhibiting other taxa (e.g., Oedogonium) because of the attendant high pH.Expt. III was designed to test effects of depth (0.1 m vs. 0.5 m) and N (NH₄ ⁺ vs. NO₃ ^–) upon the growth response to bicarbonate observed in Expts. I and II. Similar standing crop and species composition were noted on pots at 0.1 m vs. 0.5 m. Enrichment with NH₄ ⁺ vs. NO₃ ^– also appeared to have little effect upon the periphyton community.Shallow water periphyton communities in Lake Lacawac, when supplied with sufficient N and P, appear to show a distinctive response to increasing bicarbonate concentration and pH which is robust to moderate variation in grazer densities, distance from the water surface, and the form of N enrichment.     

遮光处理对观赏栀子氮、磷、钾分配与生长的影响

为探究观赏栀子在不同遮光条件下生长和光合及氮、磷、钾(N、P、K)分配的规律,该研究对3种观赏栀子设置5种不同的遮光处理(0%、60%、70%、80%、90%),通过对观赏栀子的生长和光合及各器官N、P、K含量进行统计与分析,探究了不同遮光处理对观赏栀子栽植效果的影响。结果表明:(1)60%和0%遮光率的大花栀子、80%和70%遮光率的雀舌栀子、0%遮光率的花叶栀子均长势较好。(2)随着遮光率的增加,雀舌栀子和花叶栀子的净光合速率、胞间CO₂浓度、蒸腾速率逐渐降低,气孔导度、水分瞬时利用率在遮光率60%下达到最大值。(3)大花栀子总N量最高,器官含N量高低排序为叶、根、茎; 雀舌栀子总P量最高,器官含P量排序为根、茎、叶; 大花栀子在90%遮光率下含K量最高。综合考虑认为,雀舌栀子更耐荫蔽,适合在80%和70%的遮光条件下生长; 大花栀子次之,适合在60%和0%遮光条件下生长; 花叶栀子最不耐荫,适合在0%遮光条件下生长。     

Phylogenetic ecology of foliar N and P concentrations and N:P ratios across mediterranean‐type ecosystems

Aim Interpretations of global‐scale patterns in foliar N and P concentrations and N:P ratios across climatic gradients and biomes often ignore complications imposed by taxonomic and phylogenetic structure in the nutrient concentrations of the constituent taxa. We analysed foliar N and P concentrations and N:P ratios of species from similar climate zones, but with contrasting soil fertility, to determine the relative importance of phylogeny and geographic region in driving foliar nutrient concentrations. Location Mediterranean climate regions. Methods Mean foliar N and P concentrations of 564 species from five mediterranean climate regions were compiled. Regional comparisons of foliar N and P concentrations and N:P ratios were performed using a phylogenetically independent contrasts (PIC) procedure. We also evaluated phylogenetic structuring in these variables, determining for each clade whether (1) the mean trait value and (2) the variance in trait values deviate significantly from chance expectation. Results Foliar N and P concentrations were found to vary between regions, a result confirmed using PIC. Tests of phylogenetic structure identified lineages having unusually high or low N and P concentrations, these being largely consistent amongst two of the regions. There was a general pattern of conservatism in foliar N and P concentrations and N:P ratios with localized instances of overdispersion. Main conclusions Our findings identify soil fertility as a strong environmental filter which has led to the dominance of low‐nutrient adapted lineages in the South African Cape and Western Australia. There is also a pronounced clade‐specific ‘stoichiometric homeostasis’ in foliar N and P concentrations and N:P ratios and this strong phylogenetic conservatism, together with the narrow range of foliar concentrations, is an evolutionary outcome of factors associated with developmental and physiological nutrient requirements specific to each lineage. Our results reinforce the need to include phylogenetic considerations in discussions of the biological stoichiometry of plants if we are to expand our current knowledge of foliar N and P concentrations and N:P patterns from an ecological to a biogeographic scale.     

Variation in nitrogen and phosphorus concentrations of wetland plants

The use of nutrient concentrations in plant biomass as easily measured indicators of nutrient availability and limitation has been the subject of a controversial debate. In particular, it has been questioned whether nutrient concentrations are mainly species' traits or mainly determined by nutrient availability, and whether plant species have similar or different relative nutrient requirements. This review examines how nitrogen and phosphorus concentration and the N:P ratio in wetland plants vary among species and sites, and how they are related to nutrient availability and limitation. We analyse data from field studies in European non-forested wetlands, from fertilisation experiments in these communities and from growth experiments with wetland plants. Overall, the P concentration was more variable than the N concentration, while variation in N:P ratios was intermediate. Field data showed that the N concentration varies more among species than among sites, whereas the N:P ratio varies more among sites than among species, and the P concentration varies similarly among both. Similar patterns of variation were found in fertilisation experiments and in growth experiments under controlled nutrient supply. Nutrient concentrations and N:P ratios in the vegetation were poorly correlated with various measures of nutrient availability in soil, but they clearly responded to fertilisation in the field and to nutrient supply in growth experiments. In these experiments, biomass N:P ratios ranged from 3 to 40 and primarily reflected the relative availabilities of N and P, although N:P ratios of plants grown at the same nutrient supply could vary three-fold among species. The effects of fertilisation with N or P on the biomass production of wetland vegetation were well related to the N:P ratios of the vegetation in unfertilised plots, but not to N or P concentrations, which supports the idea that N:P ratios, rather than N or P concentrations, indicate the type of nutrient limitation. However, other limiting or stressing factors may influence N:P ratios, and the responses of individual plant species to fertilisation cannot be predicted from their N:P ratios. Therefore, N:P ratios should only be used to assess which nutrient limits the biomass production at the vegetation level and only when factors other than N or P are unlikely to be limiting.     

Contrasted Responses of Two Understorey Species to Direct and Indirect Effects of a Canopy Gap

Positive associations between adult trees and understorey species have been explained either by direct or indirect facilitation. We tested both models by comparing the performance of two understorey species with contrasted stress-tolerance abilities Galium odoratum and Deschampsia flexuosa. Individuals of both species were transplanted in the four combinations of two treatments (gap and removal of an herbaceous competitor, Molinia caerulea). Our experiment demonstrated that direct facilitation of adult trees may explain the restricted occurrence of the shade-demanding Galium within closed forest communities. In contrast, the shade-tolerant Deschampsia was subjected to additional competition within the forest, likely because adult trees had a higher negative effect on light availability and a similar negative effect on nitrogen availability within the forest than did Molinia in the gaps.     

N limitation increases along a temperate forest succession: evidences from leaf stoichiometry and nutrient resorption

温带森林演替加剧了氮限制：来自叶片化学计量和养分重吸收的证据 森林生产力和碳汇功能在很大程度上取决于土壤氮和磷的有效性。然而,迄今为止,养分限制随森林演替的时间变化仍存在争议。叶片化学计量和养分重吸收是预测植物生长养分限制的重要指标。基于此,本研究测定了温带森林4个演替阶段所有木本植物叶片和凋落叶中氮和磷的含量,并分析了演替过程中非生物因子和生物因子如何影响叶片化学计量和养分重吸收。研究结果表明,在个体尺度上,叶片氮磷含量在演替末期显著增加,而叶片氮磷比无显著变化;氮的重吸收效率随演替显著增加,然而磷的重吸收效率先增加后减少;氮重吸收效率与磷重吸收效率的比值仅在演替末期显著增加。此外,植物氮素循环对土壤养分的响应比磷素循环更弱。在群落尺度上,叶片氮磷含量随森林演替呈现先降低后升高的趋势,主要受香农-维纳多样性指数和物种丰富度的影响;叶片氮磷比随演替而显著变化,主要由胸径的群落加权平均值决定;氮的重吸收效率增加,主要受物种丰富度和胸径的影响,而磷的重吸收效率相对稳定。因此,氮重吸收效率与磷重吸收效率的比值显著增加,表明随着温带森林演替,氮限制加剧。这些结果可能反映了较高生物多样性群落中物种间对有限资源的激烈竞争,强调了生物因子在驱动森林生态系统养分循环中的重要性,为中国温带和北方森林可持续经营的施肥管理提供了参考。