The seasonal dynamics of amino acids and other nutrients in Alaskan Arctic tundra soils

Past research strongly indicates the importance of amino acids in the N economy of the Arctic tundra, but little is known about the seasonal dynamics of amino acids in tundra soils. We repeatedly sampled soils from tussock, shrub, and wet sedge tundra communities in the summers of 2000 and 2001 and extracted them with water (H₂O) and potassium sulfate (K₂SO₄) to determine the seasonal dynamics of soil amino acids, ammonium (NH₄⁺), nitrate (NO₃^–), dissolved organic nitrogen (DON), dissolved organic carbon (DOC), and phosphate (PO₄^2–). In the H₂O extractions mean concentrations of total free amino acids (TFAA) were higher than NH₄⁺ in all soils but shrub. TFAA and NH₄⁺ were highest in wet sedge and tussock soils and lowest in shrub soil. The most predominant amino acids were alanine, arginine, glycine, serine, and threonine. None of the highest amino acids were significantly different than NH₄⁺ in any soil but shrub, in which NH₄⁺ was significantly higher than all of the highest individual amino acids. Mean NO₃^– concentrations were not significantly different from mean TFAA and NH₄⁺ concentrations in any soil but tussock, where NO₃^– was significantly higher than NH₄⁺. In all soils amino acid and NH₄⁺ concentrations dropped to barely detectable levels in the middle of July, suggesting intense competition for N at the height of the growing season. In all soils but tussock, amino acid and NH₄⁺ concentrations rebounded in August as the end of the Arctic growing season approached and plant N demand decreased. This pattern suggests that low N concentrations in tundra soils at the height of the growing season are likely the result of an increase in soil N uptake associated with the peak in plant growth, either directly by roots or indirectly by microbes fueled by increased root C inputs in mid-July. As N availability decreased in July, PO₄^2– concentrations in the K₂SO₄ extractions increased dramatically in all soils but shrub, where there was a comparable increase in PO₄^2– later in the growing season. Previous research suggests that these increases in PO₄^2– concentrations are due to the mineralization of organic phosphorus by phosphatase enzymes associated with soil microbes and plant roots, and that they may have been caused by an increase in organic P availability.     

An Exotic Tree Alters Decomposition and Nutrient Cycling in A Hawaiian Montane Forest

We evaluated the effects of the exotic tree Fraxinus uhdei on decomposition dynamics and nutrient turnover in a montane Hawaiian rainforest. We used reciprocal transplants of litterbags between forests dominated by Fraxinus and by the native Metrosideros polymorpha to distinguish between endogenous (litter quality) and exogenous (for example, microclimate, nutrient availability, microbial and invertebrate communities) effects of Fraxinus on mass loss and nutrient dynamics of decomposing litter. Fraxinus produced greater quantities of litter that was thinner, had higher N and P concentrations, and lower concentrations of lignin and soluble polyphenols. Microbes decomposing Fraxinus litter produced fewer enzymes involved in N and P acquisition and more of those involved in cellulose degradation. Differences in litter quality and microbial activity resulted in a strong effect of litter type on rates of mass loss, whereby Fraxinus litter decomposed and released nutrients at nearly twice the rate of Metrosideros litter (k=0.82 versus 0.48), regardless of site of decomposition. Although site of decomposition had no effect on rates of litter mass loss, Fraxinus litter decomposed under a Fraxinus canopy mineralized approximately 20% less P after one year than Fraxinus litter decomposed under a Metrosideros canopy. Furthermore, Fraxinus litter decomposed under a Fraxinus canopy immobilized greater amounts of N and P in the early stages of decay, suggesting that the large amounts of N and P in Fraxinus litterfall have raised nutrient availability to decomposers in the forest floor. Greater immobilization of N and P under a Fraxinus canopy may act as a governor on rates of nutrient cycling, limiting the degree to which Fraxinus invasion accelerates N and P cycling in this system.     

定位施肥对紫色菜园土磷素状况的影响

基于紫色菜园土壤莴笋-白菜轮作3a的12季连续定位施肥试验,研究施肥对土壤磷素状况的影响。结果表明,在紫色土上以化肥为基础增施有机肥(泥炭或菜籽粕)既提高土壤磷含量,又增加无机磷组分中有效磷源和缓效磷源的比例,且不会增加磷素淋失的风险,是所有处理中最优者;增磷、增钾处理虽然提高了土壤磷含量,但未能提高速效磷源与缓效磷源的比例;增氮、增硼及常规施肥降低土壤磷含量和提高无效磷源的比例;无肥处理磷含量最低,无效磷源比例最高。紫色菜园土无机磷约为有机磷的4—6倍,无机磷各组分以钙磷为主,占I-P总量的60.89%—67.92%,显示紫色土风化程度较低。各形态磷素呈Ca10-P>Ca8-P>Fe-P≈Al-P≈OC-P>Ca2-P序列变化,其中Ca2-P和Al-P总共仅占无机磷总量的11.47%—19.43%,表明紫色土中对植物最有效的磷源不足;而Ca10-P和OC-P共占无机磷总量的42.48%—59.55%,意味着紫色土无机磷有一半左右是以作物不可利用的形态存在的。紫色土全磷与无机磷、有机磷、Ca2-P、Ca8-P、Al-P、Fe-P、OC-P及Ca10-P呈显著正相关;无机磷与多种形态磷呈显著正相关(除有机磷、Ca2-P、有效磷外);有机磷与全磷、Ca2-P显著正相关;有效磷与全磷、Ca2-P、Al-P、Fe-P显著正相关;无机磷组分间也都存在显著正相关关系(OC-P与Ca10-P间有极显著正相关,但它们与其它无机磷组分无相关性)。表明在土壤磷的整个循环系统中,不同组分无机磷与有机磷之间处于一个动态平衡中,它们之间存在着相互影响和制约。土壤pH与各形态磷关系密切。对土壤有效磷与无机磷组分间的关系进行系统分析,简单相关分析结果土壤有效磷与Ca2-P、Fe-P呈极显著正相关,与Al-P呈显著正相关,与Ca8-P、OC-P、Ca10-P正相关但不显著;通径分析结果对土壤有效磷直接影响较大的无机磷组分是Al-P、Ca8-P与Ca2-P,其中Ca8-P的直接影响为负效应;逐步回归分析结果与通径分析一致。综合分析得出,Ca2-P、Al-P是最有效的磷源,Fe-P、Ca8-P次之,Ca10-P与OC-P是非有效磷源。莴笋和白菜产量与紫色土壤Ca2-P、Ca8-P、Al-P、Fe-P呈显著正相关,与OC-P、Ca10-P无相关性。紫色土各形态无机磷与两种蔬菜产量的相关系数r大小依次为Ca2-P>Fe-P>AL-P>Ca8-P>Ca10-P>OC-P,这与各形态无机磷与有效磷简单相关分析的结果一致。     

Plant phenols contents and their changes with nitrogen availability in peatlands of northeastern China

气候变暖和大气氮沉降增加会改变北方泥炭地的养分状态,从而影响其植被的物种组成和固碳功能。酚类物质是植物用于 防御植食性动物和适应环境的次生代谢产物,由于其具有抗分解的特性,在调节泥炭地碳动态方面也起着重要的作用。然而,北方 泥炭地不同功能型植物的酚含量及其如何随氮有效性变化尚不清楚。本论文通过测定中国东北大兴安岭地区18个泥炭地共11 种植物的叶片总酚含量(Total Phenols Contents, TPC),研究了它们随叶片氮、磷含量的变化关系,并探讨了其潜在机制。结果表明,灌木叶片TPC高于草本植物,说明生长较快且无菌根的草本植物比生长较慢且具有菌根的灌木对防御的碳投入较少。灌木叶片TPC随叶片氮含量增 加而降低,表明其防御碳投入随氮有效性增加而减少。相反,草本植物叶片TPC随氮含量增加而增加,随磷含量增加而减少,由于草本植物相对于灌木具有较强的氮限制和较弱的磷限制,草本植物防御碳投入可能随养分有效性增加而增加。我们的结果表明,泥炭地在氮有效性随气候变暖和氮沉降增加而增加的背景下,灌木将投入相对较多的碳用于生长而非防御,而草本植物则与之相反。这些发现将有助于对北方泥炭地灌木入侵及其资源竞争机制的理解。     

长期施肥和增水对半干旱草地土壤性质和植物性状的影响

本文对内蒙古多伦退化草地2005年建立的长期野外控制试验中施肥和增水对土壤性质和植物群落特征的影响进行了总结和综合评述.结果表明:加氮导致了表土酸化并降低酸缓冲容量,提高了表土中碳氮磷硫有效性及DTPA-浸提态铁锰铜含量,导致盐基离子钙镁钾钠总量的消耗,降低了土壤微生物群落多样性,促进了优势植物物种叶片对氮磷硫钾及锰铜锌的吸收,抑制植物叶片对铁的吸收,而对钙镁吸收无显著影响,增加了植物地上净初级生产力(ANPP),降低了植物物种多样性和群落稳定性.单独加磷增加了表土全磷和Olsen-P含量及真菌丰度,促进了植物叶片对氮、磷、硫的吸收,但对其他土壤基本化学性质及ANPP、物种多样性无显著影响.增水提高了植物群落对干旱的抵抗力,但对ANPP增长的贡献受到土壤氮有效性的限制.增水对于加氮导致的土壤酸化、植物和微生物多样性降低等具有一定的缓冲作用;加氮增水和加磷增水下,土壤微生物多样性及功能受地上植物群落结构及功能变化的影响.长期野外控制试验对于深入理解草地生态系统结构和功能对环境变化的响应具有重要意义,但单点的研究结果仍需与不同区域多点控制试验的联网研究相结合,深入开展地上与地下生态过程的关联研究,才能深入理解草地生态系统生态学的相关机制.     

Bringing function to structure: Root–soil interactions shaping phosphatase activity throughout a soil profile in Puerto Rico

1. Large areas of highly productive tropical forests occur on weathered soils with low concentrations of available phosphorus (P). In such forests, root and microbial production of acid phosphatase enzymes capable of mineralizing organic phosphorus is considered vital to increasing available P for plant uptake.
2. We measured both root and soil phosphatase throughout depth and alongside a variety of root and soil factors to better understand the potential of roots and soil biota to increase P availability and to constrain estimates of the biochemical mineralization within ecosystem models.
3. We measured soil phosphatase down to 1 m, root phosphatase to 30 cm, and collected data on fine‐root mass density, specific root length, soil P, bulk density, and soil texture using soil cores in four tropical forests within the Luquillo Experimental Forest in Puerto Rico.
4. We found that soil phosphatase decreased with soil depth, but not root phosphatase. Furthermore, when both soil and root phosphatase were expressed per soil volume, soil phosphatase was 100‐fold higher that root phosphatase.
5. Both root and soil factors influenced soil and root phosphatase. Soil phosphatase increased with fine‐root mass density and organic P, which together explained over 50% of the variation in soil phosphatase. Over 80% of the variation in root phosphatase per unit root mass was attributed to specific root length (positive correlation) and available (resin) P (negative correlation).
6. Synthesis: Fine‐root traits and soil P data are necessary to understand and represent soil and root phosphatase activity throughout the soil column and across sites with different soil conditions and tree species. These findings can be used to parameterize or benchmark estimates of biochemical mineralization in ecosystem models that contain fine‐root biomass and soil P distributions throughout depth.

     

Plant phenological responses to a long‐term experimental extension of growing season and soil warming in the tussock tundra of Alaska

Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis‐idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species‐specific and growth form‐specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.     

Plant community responses to nitrogen addition and increased precipitation: the importance of water availability and species traits

Global nitrogen (N) enrichment and changing precipitation regimes are likely to alter plant community structure and composition, with consequent influences on biodiversity and ecosystem functioning. Responses of plant community structure and composition to N addition and increased precipitation were examined in a temperate steppe in northern China. Increased precipitation and N addition stimulated and suppressed community species richness, respectively, across 6 years (2005–2010) of the manipulative experiment. N addition and increased precipitation significantly altered plant community structure and composition at functional groups levels. The significant relationship between species richness and soil moisture (SM) suggests that plant community structure is mediated by water under changing environmental conditions. In addition, plant height played an important role in affecting the responses of plant communities to N addition, and the effects of increased precipitation on plant community were dependent on species rooting depth. Our results highlight the importance and complexity of both abiotic (SM) and biotic factors (species traits) in structuring plant community under changing environmental scenarios. These findings indicate that knowledge of species traits can contribute to mechanistic understanding and projection of vegetation dynamics in response to future environmental change.     

Effects of nitrogen fertilization of grapefruit trees on soil acidification and nutrient availability in a Riviera fine sand

Nitrogen (N) fertilizer applied in the NH4+ form results in some degree of soil acidification, which could influence nutrient availability to plants and nutrient losses through leaching. Effects of various N rates (0 – 168 kg N ha-1 yr-1) on soil acidification and nutrient availability were investigated in a Riviera fine sand with 26-year-old white Marsh grapefruit (Citrus paradisi MacFadyen) trees. Soil pH significantly decreased with increasing NH4–N rates. Application of 112 kg N ha-1 yr-1 for four years decreased the pH by 0.7 to 1.7 unit. Soil acidification was greater when the NH4+ form of N fertilizer was applied as dry soluble granular material compared to fertigation or controlled release forms. The marked effect of NH4–N fertilization on the pH of the Riviera fine sand was due to its low buffering capacity. Soil acidification increased the concentration of extractable Fe and P but decreased that of K, Zn and Mn. Soil pH was positively correlated with concentration of Ca, but negatively with concentrations of Fe, Mn and Zn in six-month-old spring flush leaves of the grapefruit trees. Leaf P concentrations, however, were poorly correlated with soil pH. This study also demonstrated an increase in leaching of P and K below the grapefruit trees rootzone with a decrease in soil pH.     

Available soil nitrogen in relation to fractions of soil nitrogen and other soil properties

The available N of 27 soils from England and Wales was assessed from the amounts of N taken up over a 6-month period by perennial ryegrass grown in pots under uniform environmental conditions. Relationships between availability and the distribution of soil N amongst various fractions were then examined using multiple regression. The relationship: available soil N (mg kg^–1 dry soil)=(N_min×0.672)+(N_inc×0.840)+(N_mom×0.227)–5.12 was found to account for 91% of the variance in available soil N, where N_min=mineral N, N_inc=N mineralized on incubation and N_mom=N in macro-organic matter. The N mineralized on incubation appeared to be derived largely from sources other than the macro-organic matter because these two fractions were poorly correlated. When availability was expressed in terms of available organic N as % of soil organic N (N_ao) the closest relationship with other soil characteristics was: N_ao=[N_inc×(1.395–0.0347×CN_mom]+[N_mom×0.1416], where CN_mom=CN ratio of the macro-organic matter. This relationship accounted for 81% of the variance in the availability of the soil organic N.The conclusion that the macro-organic matter may contribute substantially to the available N was confirmed by a subsidiary experiment in which the macro-organic fraction was separated from about 20 kg of a grassland soil. The uptake of N by ryegrass was then assessed on two subsamples of this soil, one without the macro-organic matter and the other with this fraction returned: uptake was appreciably increased by the macro-organic matter.     

Links between plant community composition,soil organic matter quality and microbial communities in contrasting tundra habitats

Plant communities, soil organic matter and microbial communities are predicted to be interlinked and to exhibit concordant patterns along major environmental gradients. We investigated the relationships between plant functional type composition, soil organic matter quality and decomposer community composition, and how these are related to major environmental variation in non-acid and acid soils derived from calcareous versus siliceous bedrocks, respectively. We analysed vegetation, organic matter and microbial community compositions from five non-acidic and five acidic heath sites in alpine tundra in northern Europe. Sequential organic matter fractionation was used to characterize organic matter quality and phospholipid fatty acid analysis to detect major variation in decomposer communities. Non-acidic and acidic heaths differed substantially in vegetation composition, and these disparities were associated with congruent shifts in soil organic matter and microbial communities. A high proportion of forbs in the vegetation was positively associated with low C:N and high soluble N:phenolics ratios in soil organic matter, and a high proportion of bacteria in the microbial community. On the contrary, dwarf shrub-rich vegetation was associated with high C:N and low soluble N:phenolics ratios, and a high proportion of fungi in the microbial community. Our study demonstrates a strong link between the plant community composition, soil organic matter quality, and microbial community composition, and that differences in one compartment are paralleled by changes in others. Variation in the forb-shrub gradient of vegetation may largely dictate variations in the chemical quality of organic matter and decomposer communities in tundra ecosystems. Soil pH, through its direct and indirect effects on plant and microbial communities, seems to function as an ultimate environmental driver that gives rise to and amplifies the interactions between above- and belowground systems. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Nutrient limitation of phytoplankton production in Alaskan Arctic foothill lakes

We used 54 enrichment bioassays to assess nutrient limitation (N, P) of ¹⁴C uptake by natural phytoplankton assemblages in 39 lakes and ponds in the Arctic Foothills region of Alaska. Our purpose was to categorize phytoplankton nutrient status in this under-represented region of North America and to improve our ability to predict the response of primary production to anticipated anthropogenically mediated increases in nutrient loading. Experiments were performed across several watersheds and included assays on terminal lakes and lakes occupying various positions in chains (lakes in series within a watershed and connected by streams). In total, 89% (48 of 54) of the bioassays showed significant stimulation of ¹⁴C primary production by some form of nutrient addition relative to unamended controls. A significant response was observed following enrichment with N and P, N alone and P alone in 83, 35 and 22% of the bioassays, respectively. In experiments where N and P proved stimulatory, the influence of N alone was significantly greater than the influence of P alone. Overall, the data point to a greater importance for N than P in regulating phytoplankton production in this region. The degree of response to N and P enrichment declined as the summer progressed and showed no relationship to irradiance or water temperature, suggesting secondary limitation by some micronutrient such as iron as the summer advanced. Phytoplankton nutrient status was often consistent across lakes within a watershed, suggesting that watershed characteristics influence nutrient availability. Lakes in this region will clearly show increased phytoplankton production in response to anthropogenic activities and anticipated changes in climate that will increase nutrient loading.     

Complex vegetation responses to soil disturbances in mountain grassland

We studied vegetation responses to disturbances originated by ants and voles in subalpine grasslands in the Eastern Pyrenees. We compared the effects of these small-scale disturbances with those of a large-scale disturbance caused by ploughing. We wanted to know if these soil disturbances promoted species richness through the existence of a specific guild of plants colonizing these areas, and if this guild was the same for all soil disturbances, independently of their extent. In general, grassland vegetation seemed to recover relatively quickly from soil-displacement disturbances, and the effects could be scaled up in time and space in terms of species richness and composition. Vole mound composition was similar to that in the surrounding grassland, suggesting that mounds were rapidly colonized by the neighbouring vegetation. Vegetation composition differed between the grassland and the ant mounds. Grasses and erect dicots coped well with repeated disturbance, while rosette-forming species and sedges were very sensitive to it. Landscape processes could be important to understanding recolonization. Species from xeric grasslands were found in mesic grasslands when disturbed by ploughing and on the tops of active ant mounds. Furrows in mesic grasslands recovered well, but decades after disturbance showed long persistence of some xeric species and increased species richness compared to terraces, while xeric grasslands showed decreased richness. This suggests that, because of those disturbances, within-habitat diversity was increased, although landscape diversity was not. However, specific disturbances showed idiosyncratic effects, which could enhance the species richness globally. In ant-affected areas, the grassland itself showed the highest plant species richness, partially associated to the presence of some species with elaiosomes not, or only rarely, found in adjacent grasslands without ant mounds. Therefore, soil disturbances occurring at different spatial scales contributed to complexity in vegetation patterns in addition to abiotic factors and grazing. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Nomenclature of the species follows Tutin et al. (1964–1980) and Bolòs et al. (1993).     

Plant responses to fertilization and exclusion of grazers on an arctic tundra heath

This study investigated the impacts of fertilization and grazing by Norwegian lemmings (Lemmus lemmus), grey‐sided voles (Clethrionomys rufocanus), and reindeer (Rangifer tarandus) on a diverse tundra plant community dominated by deciduous shrubs. Four out of eight study areas, having a size of 2500 m² each, were fertilized with a N‐P‐K fertilizer and four areas served as unfertilized controls. Two types of exclosures were used within each study area, one to exclude solely reindeer, and one to exclude both rodents and reindeer. Open, grazed plots served as controls. During 5 years following the fertilization event the changes in vegetation inside and outside the exclosures were monitored using a point frequency method. The densities of rodents on the fertilized and unfertilized areas were investigated by live trapping and by counting nests of overwintering individuals. Reindeer do not graze on the study area during the growing season but migrate through this area in autumn and spring. Fertilization increased the abundance of vascular plants while grazing by reindeer and rodents decreased the abundance of vascular plants significantly on both fertilized and unfertilized areas. Rodents preferred clearly the fertilized areas during winter, decreasing the abundance of Vaccinium myrtillus and Vaccinium vitis‐idaea, while very little grazing occurred during summer. Graminoids showed the strongest positive response to fertilization and dominated the plant community on ungrazed plots, while winter grazing by both reindeer and rodents significantly decreased the abundance of graminoids. Deciduous shrubs (Betula nana, Vaccinium myrtillus) increased slightly but significantly due to fertilization and evergreen dwarf shrubs showed no response to fertilization. However, the use of functional growth forms for predicting the responses of nutrient enrichment and grazing must be questioned, as responses to fertilization as well as preferences by herbivores were shown to be species‐specific rather than uniform within functional groups based on plant growth forms.