岷江干旱河谷灌木幼苗的土壤养分适应性研究

岷江上游干旱河谷地区的植被恢复是目前急需解决的问题,小马鞍羊蹄甲（Bauhinia faberi var. microphylla）是干旱河谷地区的优势乡土灌木,并作为当地植被恢复过程中的重要备选物种,因而进行小马鞍羊蹄甲种群在土壤贫瘠的干旱河谷的养分限制研究十分必要。针对性地进行施肥将影响到小马鞍羊蹄甲幼苗存活、生长和定殖,而如何有效提高该物种的存活和生长速率对于干旱河谷植被恢复将具有重要的意义。采用野外调查与模拟实验相结合的方法,分别研究了干旱河谷地区3个典型群落（干旱河谷的核心区——飞虹,干旱河谷灌丛与亚高山森林的过渡区——北部的石大关、干旱河谷灌丛与温性森林的过渡区——南部的蓝新镇）中小马鞍羊蹄甲幼苗叶片的化学计量特征,和施肥试验[w（有效氮）分别为100、280、460 mg·kg–1、w（有效磷）分别为12、24、48 mg·kg–1、w（氮）分别为40、70、100 mg·kg–1、w（磷）分别为12和24 mg·kg–1]中叶片化学计量特征及幼苗生长参数（叶片数、基径和株高）和各器官（根茎叶）生物量。结果表明：在野外各演替阶段的小马鞍羊蹄甲幼苗生物量和营养元素质量分数都随着磷肥的增加而增加,表明磷素是植物生长的限制因子,同时N∶P比均大于16,也暗示其受到P养分的限制;在室内施肥试验中,施加N肥没有促进小马鞍羊蹄甲生物量的积累,反而抑制了幼苗生长;施加P肥促进了幼苗生物量的积累,表明幼苗缺乏P元素,养分限制类型为P限制。     

Spectral analysis of forest model time series

The use of spectral analysis to elucidate the cyclic behavior in time series generated by a forest stand growth simulation model is discussed. A stand-level simulator, FORET, for an Appalachian deciduous forest is described. An estimate of the power spectral density of the total biomass time series is calculated. The power spectral density estimate indicates a dominant cyclic behavior with a period of about 200 years. In addition the spectral density is approximately bandlimited. This characteristic makes possible the application of the sampling theorem for analysis of sampling rates.     

氮沉降下西南亚高山针叶林生长的养分限制特征

长期氮(N)沉降及其诱导的N、磷(P)养分平衡性对森林生长与生产力的生态反馈效应已成为当前森林生态学研究的前沿与热点,但目前大多研究主要基于已有文献数据整合分析,而缺乏野外原位系统性研究与试验证据.以西南亚高山两种典型人工针叶林——云杉(Picea asperata)林和华山松(Pinus armandii)林为对象,...     

Extending the leaf economics spectrum to decomposition: evidence from a tropical forest

I investigated the relationship between leaf physiological traits and decomposition of leaf litter for 35 plant species of contrasting growth forms from a lowland tropical forest in Panama to determine whether leaf traits could be used to predict decomposition. Decomposition rate (k) was correlated with specific leaf area (SLA), leaf nitrogen (N), phosphorus (P), and potassium (K) across all species. Photosynthetic rate per unit mass (Amass) was not correlated with k, but structural equation modeling showed support for a causal model with significant indirect effects of Amass on k through SLA, N, and P, but not K. The results indicate that the decomposability of leaf tissue in this tropical forest is related to a global spectrum of leaf economics that varies from thin, easily decomposable leaves with high nutrient concentrations and high photosynthetic rates to thick, relatively recalcitrant leaves with greater physical toughness and defenses and low photosynthetic rates. If this pattern is robust across biomes, then selection for suites of traits that maximize photosynthetic carbon gain over the lifetime of the leaf may be used to predict the effects of plant species on leaf litter decomposition, thus placing the ecosystem process of decomposition in an evolutionary context.     

Resource availability and the abundance of an N-based defense in Australian tropical rain forests

Plant defense theories predict that relatively resource-rich environments (those with more fertile soil) will support a greater abundance of plants with nitrogen-based chemical defense, but this has yet to be adequately tested. We tested this prediction by measuring the diversity and contribution to total biomass of cyanogenic plants (those that release hydrogen cyanide from endogenous cyanide-containing compounds) in the Australian tropical rain forest. We examined 401 species in thirty 200-m2 plots, six at each of five sites, for cyanogenesis. In upland/highland rain forest, two pairs of sites similar in rainfall and altitude, but differing in soil nutrients, were selected, as well as one site in lowland rain forest. Sites differed markedly in species composition and foliar N was positively related to soil fertility. Holding altitude constant, we did not detect significant differences in the proportion of cyanogenic species with soil fertility, nor did we consistently detect significant increases in the contribution of cyanogenic species to total biomass on higher nutrient sites. Thus we found no clear evidence that soil fertility affects the distribution and prevalence of species investing in a constitutive N-based defense at the community level.     

Patterns in soil fertility and root herbivory interact to influence fine-root dynamics

Fine-scale soil nutrient enrichment typically stimulates root growth, but it may also increase root herbivory, resulting in trade-offs for plant species and potentially influencing carbon cycling patterns. We used root ingrowth cores to investigate the effects of microsite fertility and root herbivory on root biomass in an aggrading upland forest in the coastal plain of South Carolina, USA. Treatments were randomly assigned to cores from a factorial combination of fertilizer and insecticide. Soil, soil fauna, and roots were removed from the cores at the end of the experiment (8-9 mo), and roots were separated at harvest into three diameter classes. Each diameter class responded differently to fertilizer and insecticide treatments. The finest roots (< 1.0 mm diameter), which comprised well over half of all root biomass, were the only ones to respond significantly to both treatments, increasing when fertilizer and when insecticide were added (each P < 0.0001), with maximum biomass found where the treatments were combined (interaction term significant, P < 0.001). These results suggest that root-feeding insects have a strong influence on root standing crop with stronger herbivore impacts on finer roots and within more fertile microsites. Thus, increased vulnerability to root herbivory is a potentially significant cost of root foraging in nutrient-rich patches.     

Multigenerational genetic analysis of tropical secondary regeneration in a canopy palm

Iriartea deltoidea (Arecaceae) is an abundant canopy palm with a wide geographic distribution in Neotropical wet forests. We analyzed the genetic profile across three generations of Iriartea within a 43-ha area encompassing two areas of second-growth and adjoining old-growth forest at La Selva Biological Field Station in northeastern Costa Rica. A total of 311 reproductively mature trees, 99 large saplings, 207 small saplings, and 601 seedlings were genotyped using 141 AFLP loci. Parentage analysis revealed high dispersal distances, both for seed (over 2.3 km) and pollen (over 3.8 km), indicating a large genetic neighborhood within La Selva Biological Station. In a 20-ha area of second growth, the founding palm population was dominated by a small number of parental trees located in the adjacent old-growth forest; two old-growth trees contributed 48% of the second-growth genes. The genetic diversity of reproductively mature trees in this second-growth forest was significantly reduced compared to adjacent old-growth forest. Within 400 m of the border with old-growth forest, we observed a similar reduction of genetic diversity in saplings, and an even greater loss of genetic diversity in the second generation of seedlings. Nearly half of these seedlings were offspring of local parents. In contrast, in the distant portion of second-growth forest (400-800 m from the old-growth border), parentage analysis showed that 40% of seedlings originated from outside the study area and only 10% were offspring of local parents. These high levels of gene flow maintained genetic diversity in saplings and seedlings similar to levels observed in old-growth forest. Our findings highlight the importance of gene flow from diverse seed and pollen sources for sustaining levels of genetic diversity of tree populations in second-growth forests.     

不同林龄马尾松（Pinus massoniana）人工林碳氮磷分配格局及化学计量特征

为了解长江上游低山丘陵区马尾松（Pinus massoniana）人工林生态系统的C、N、P分配格局及化学计量特征,本文采用时空互代的方法,在宜宾高县来复林区选取三种不同林龄（5年生幼龄林、14年生中龄林、39年生成熟林）,但立地条件相近、样地情况基本一致的马尾松（Pinus massoniana）人工林作为研究对象,对马尾松针叶、凋落物及土壤中的C、N、P含量及 w（C）？w（N）？w（P）化学计量特征进行测定和分析。结果表明,（1）C、N、P 含量均表现为针叶〉凋落物〉土壤,且在三个库之间差异显著;（2）林龄对针叶、凋落物、土壤的 C、N、P 及 w（C）？w（N）、w（C）？w（P）计量比均有显著影响。（3）土壤 C、N、P含量在成熟林中最高;针叶和凋落物的C含量在成熟林中最低,N、P含量则在中龄林中最高。（4）随林龄增加马尾松对N、P的利用效率降低,针叶、凋落物及土壤的w（C）？w（N）与 w（C）？w（P）均表现为下降。（5）马尾松针叶w（N）？w（P）比值在14.37~15.53之间,说明该地区马尾松人工林受N和P的共同限制,但林龄对N、P养分限制的影响不显著。为提高该区马尾松人工林的生产力,建议在人工林的抚育管理中要适当增加N肥和P肥,同时也可在马尾松人工林引入豆科固氮植物以提高地力。该研究将马尾松针叶、凋落物及土壤结合起来探究随林龄增长C、N、P养分元素的分配格局及化学计量特征的变化,有助于全面、系统地揭示马尾松人工林生态系统的养分循环,对指导马尾松人工林生产,调节和改善林木生长环境,提高系统的养分利用效率及林地生产力具有重要意义。     

Atmospheric deposition may affect northern hardwood forest composition by altering soil nutrient supply.

In the northeastern United States, the input of reactive nitrogen (N) via atmospheric deposition has increased rapidly since the onset of the industrial revolution. During the same period of time, acid precipitation and forest harvest have removed substantial quantities of base cations from soil. Because of the dominance of base-poor soils and the low rates of atmospheric base cation deposition, soils throughout the northeastern United States may be increasingly rich in N but poor in calcium (Ca). We studied the consequences of a change in soil N and Ca availability on forest composition by transplanting seedlings of four tree species into replicate plots in the understory and in canopy gaps amended with N and Ca in factorial combination. In this paper, we report on the growth and survivorship of seedlings over a four-year period. Relative to control plots, fertilization with N increased red maple growth by an average of 39% whereas fertilization with Ca decreased survivorship in the understory by 41%. In sugar maple, fertilization with Ca increased growth by 232% and 46% in the forest understory and in canopy gaps, respectively, and significantly increased high light survivorship. Fertilization with N decreased white pine survivorship by 69% in the understory whereas high Ca availability significantly increased survivorship. Fertilization with N or Ca alone reduced red oak growth but had no effect on survivorship. The results of this study suggest that historical losses of soil Ca and the continuing effects of atmospheric-N deposition on N availability are likely to alter the composition of northeastern North American forests because of the positive effects of N enrichment on the growth of red maple and the negative effects of Ca loss on the growth and survivorship of sugar maple and white pine.     

The cladoceran trophic status in the nitrogen limited ecosystem of Lake Kinneret (Israel)

The impactof cladocerans metabolic activities on the carbon (C), nitrogen (N) and phosphorus (P) dynamics in Lake Kinneret (Israel) is presented. The study, is based on the incorporation of field data and experimental measurements. Grazing, respiration and production rates of Diaphanosoma spp., Ceriodaphnia spp. and Bosmina spp. were experimentally measured at three temperatures, and the results were extrapolated to the field biomass distribution atthese respective temperatures, and the total lake capacity was calculated using the following equation: consumption = respiration + production + excretion. The field capacity of consumption, respiration and production were found to be mostly correlated with biomass density, but the temporal fluctuation of the percent of excretion from consumed energy differed. The increase in P, decline in N and decrease in the TN/TP mass ratio in the epilimnion of Lake Kinneret during 1969-2004 created N limitation. An increase in C and dedine in TN, with a consequential increase in the C/TN ratio were documented. TP was augmented but the C/TP ratio was only slightly increased. During 1975-2004, P was probably, a minor limitation for cladoceran growth. The positive impact of recycled P by cladocerans underthe N limitation in Lake Kinneret is discussed. There is a current threat on the water quality, derived from N limitation (mostly in summer-fall) and consequent Cyanophyta blooms. Thus, the role of recycled P bygrazers may be significant.     

Long-term CO2 enrichment of a forest ecosystem: implications for forest regeneration and succession.

The composition and successional status of a forest affect carbon storage and net ecosystem productivity, yet it remains unclear whether elevated atmospheric carbon dioxide (CO2) will impact rates and trajectories of forest succession. We examined how CO2 enrichment (+200 microL CO2/L air differential) affects forest succession through growth and survivorship of tree seedlings, as part of the Duke Forest free-air CO2 enrichment (FACE) experiment in North Carolina, USA. We planted 2352 seedlings of 14 species in the low light forest understory and determined effects of elevated CO2 on individual plant growth, survival, and total sample biomass accumulation, an integrator of plant growth and survivorship over time, for six years. We used a hierarchical Bayes framework to accommodate the uncertainty associated with the availability of light and the variability in growth among individual plants. We found that most species did not exhibit strong responses to CO2. Ulmus alata (+21%), Quercus alba (+9.5%), and nitrogen-fixing Robinia pseudoacacia (+230%) exhibited greater mean annual relative growth rates under elevated CO2 than under ambient conditions. The effects of CO2 were small relative to variability within populations; however, some species grew better under low light conditions when exposed to elevated CO2 than they did under ambient conditions. These species include shade-intolerant Liriodendron tulipifera and Liquidambar styraciflua, intermediate-tolerant Quercus velutina, and shade-tolerant Acer barbatum, A. rubrum, Prunus serotina, Ulmus alata, and Cercis canadensis. Contrary to our expectation, shade-intolerant trees did not survive better with CO2 enrichment, and population-scale responses to CO2 were influenced by survival probabilities in low light. CO2 enrichment did not increase rates of sample biomass accumulation for most species, but it did stimulate biomass growth of shade-tolerant taxa, particularly Acer barbatum and Ulmus alata. Our data suggest a small CO2 fertilization effect on tree productivity, and the possibility of reduced carbon accumulation rates relative to today's forests due to changes in species composition.     

Tree species control rates of free-living nitrogen fixation in a tropical rain forest

Tropical rain forests represent some of the most diverse ecosystems on earth, yet mechanistic links between tree species identity and ecosystem function in these forests remains poorly understood. Here, using free-living nitrogen (N) fixation as a model, we explore the idea that interspecies variation in canopy nutrient concentrations may drive significant local-scale variation in biogeochemical processes. Biological N fixation is the largest "natural" source of newly available N to terrestrial ecosystems, and estimates suggest the highest such inputs occur in tropical ecosystems. While patterns of and controls over N fixation in these systems remain poorly known, the data we do have suggest that chemical differences among tree species canopies could affect free-living N fixation rates. In a diverse lowland rain forest in Costa Rica, we established a series of vertical, canopy-to-soil profiles for six common canopy tree species, and we measured free-living N fixation rates and multiple aspects of chemistry of live canopy leaves, senesced canopy leaves, bulk leaf litter, and soil for eight individuals of each tree species. Free-living N fixation rates varied significantly among tree species for all four components, and independent of species identity, rates of N fixation ranged by orders of magnitude along the vertical profile. Our data suggest that variations in phosphorus (P) concentration drove a significant fraction of the observed species-specific variation in free-living N fixation rates within each layer of the vertical profile. Furthermore, our data suggest significant links between canopy and forest floor nutrient concentrations; canopy P was correlated with bulk leaf litter P below individual tree crowns. Thus, canopy chemistry may affect a suite of ecosystem processes not only within the canopy itself, but at and beneath the forest floor as well.     

计划烧除攀枝花苏铁林区地面覆盖物对苏铁生长和土壤理化性质的影响

选择攀枝花苏铁林下计划烧除试验后苏铁及其根系周围土壤(0~15 cm)为对象,研究火烧对攀枝花苏铁植株生长、叶片生理和苏铁根系周围土壤理化性质的影响,以期为区域攀枝花苏铁种群恢复和生态环境保护提供理论依据.结果表明:计划烧除区域苏铁成年树的株数及株高没有显著变化,新生叶片数目和幼苗株数相对未烧除区域显著增加,增幅分别达201.66%和317.7%;烧除区域苏铁植株新生叶片叶绿素a、叶绿素b、叶绿素a+b,可溶性糖、蛋白质含量,硝酸还原酶(NR)、谷氨酰胺合成酶(GS)活性,叶片碳(C)、氮(N)、磷(P)、钾(K)含量,C/N、N/P比与对照区域新生叶片相比无显著性差异,只有类胡萝卜素含量显著降低.计划烧除改变了苏铁根围土壤(0~15 cm)的理化性质,表现在烧除后的土壤含水量、pH值及总N、P、K含量显著降低;但是,火烧显著提高了土壤有机碳(TOC)、硝态氮(NO3--N)、铵态氮(NH4+-N)含量,微生物量C、N含量也略有升高;这说明火烧有利于增加苏铁林下土壤养分(N)的有效性,为烧除后苏铁快速恢复生长提供有利条件.     

Nitrogen turnover in the leaf litter and fine roots of sugar maple

In order to better understand the nitrogen (N) cycle, a pulse of 15NO3- was applied in 1998 to a sugar maple (Acer saccharum) dominated northern hardwood forest receiving long-term (1994-2008) simulated atmospheric N deposition. Sugar maple leaf litter and live fine-root 15N were quantified for four years prior to labeling and for 11 subsequent years. Continuous sampling of 15N following addition of the tracer enabled calculation of leaf litter and fine-root N pool turnover utilizing an exponential decay function. Fine-root 15N recovery peaked at 3.7% +/- 1.7% the year the tracer was applied, while leaf litter 15N recovery peaked in the two years following tracer application at approximately 8%. These results suggest shoots are primarily constructed from N taken up in previous years, while fine roots are constructed from new N. The residence time of N was 6.5 years in leaf litter and 3.1 years in fine roots. The longer residence time and higher recovery rate are evidence that leaves were a stronger sink for labeled N than fine roots, but the relatively short residence time of tracer N in both pools suggests that there is not tight intra-ecosystem cycling of N in this mature forest.     

Atmospheric CO2 and O3 alter the flow of 15N in developing forest ecosystems

Anthropogenic O3 and CO2-induced declines in soil N availability could counteract greater plant growth in a CO2-enriched atmosphere, thereby reducing net primary productivity (NPP) and the potential of terrestrial ecosystems to sequester anthropogenic CO2. Presently, it is uncertain how increasing atmospheric CO2 and O3 will alter plant N demand and the acquisition of soil N by plants as well as the microbial supply of N from soil organic matter. To address this uncertainty, we initiated an ecosystem-level 15N tracer experiment at the Rhinelander (Wisconsin, USA) free air CO2-O3 enrichment (FACE) facility to understand how projected increases in atmospheric CO2 and 03 alter the distribution and flow of N in developing northern temperate forests. Tracer amounts of 15NH4+ were applied to the forest floor of developing Populus tremuloides and P. tremuloides-Betula papyrifera communities that have been exposed to factorial CO2 and O3 treatments for seven years. One year after isotope addition, both forest communities exposed to elevated CO2 obtained greater amounts of 15N (29%) and N (40%) from soil, despite no change in soil N availability or plant N-use efficiency. As such, elevated CO2 increased the ability of plants to exploit soil for N, through the development of a larger root system. Conversely, elevated O3 decreased the amount of 15N (-15%) and N (-29%) in both communities, a response resulting from lower rates of photosynthesis, decreases in growth, and smaller root systems that acquired less soil N. Neither CO2 nor 03 altered the amount of N or 15N recovery in the forest floor, microbial biomass, or soil organic matter. Moreover, we observed no interaction between CO2 and 03 on the amount of N or 15N in any ecosystem pool, suggesting that 03 could exert a negative effect regardless of CO2 concentration. In a CO2-enriched atmosphere, greater belowground growth and a more thorough exploitation of soil for growth-limiting N is an important mechanism sustaining the enhancement of NPP in developing forests (0-8 years following establishment). However, as CO2 accumulates in the Earth's atmosphere, future O3 concentrations threaten to diminish the enhancement of plant growth, decrease plant N acquisition, and lessen the storage of anthropogenic C in temperate forests.     

Life history trade-offs in tropical trees and lianas

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10 mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species.     

Invasion through quantitative effects: intense shade drives native decline and invasive success.

The effects of invasive nonnative species on community composition are well documented. However, few studies have determined the mechanisms by which invaders drive these changes. The literature indicates that many nonnative plant species alter light availability differently than natives in a given community, suggesting that shading may be such a mechanism. We compared light quantity (photosynthetically active radiation, PAR) and quality (red: far-red ratio, R:Fr) in riparian reaches heavily invaded by a nonnative tree (Acer platanoides) to that in an uninvaded forest and experimentally tested the effects of our measured differences in PAR and R:Fr on the survival, growth, and biomass allocation of seedlings of the dominant native species and Acer platanoides. Light conditions representative of the understory of Acer platanoides-invaded forest decreased survival of the native maple Acer glabrum by 28%; Amelanchier alnifolia by 32%; Betula occidentalis by 55%; Elymus glaucus by 46%; and Sorbus aucuparia by 52%, relative to seedlings growing in PAR similar to that of native understories. In contrast, Acer platanoides and the native shrub Symphoricarpos albus were not affected by reductions in PAR. Acer platanoides seedlings and saplings are uniquely adapted to shade relative to native species. Acer platanoides was the only species tested that decreased allocation to roots relative to shoots in the invaded forest vs. the native forest light conditions. Therefore it was the only species to demonstrate an adaptive response to the particular light environment associated with Acer platanoides invasion as predicted by optimal partitioning theory. The profound change in light quantity associated with Acer platanoides canopies appears to act as an important driver of native suppression and conspecific success in invaded riparian communities. Further research is necessary to determine whether the effect of nonnative plant-driven changes on light quantity and quality is a widespread mechanism negatively affecting resident species and facilitating invasion by nonnatives.     

磷胁迫条件下北美红杉幼苗生长的适应性反应

采用温室盆栽的方法探讨了不同的P质量浓度(0、0.018、0.036、0.054、0.071、0.108、0.142和0.213g.L-1,以0.071g.L-1作为对照)处理下北美红杉一年生幼苗生物量及根系生长的反应,结果表明,P供应不足时,幼苗将更多的生物量分配到地下以扩大根系的生长,地下/地上生物量比率增加,缺P时为0.47。高水平供P条件下,增加幅度较大,供P水平增加2倍时,达0.66。幼苗细根/叶生物量比率与地下/地上生物量比率变化规律相似。当供P水平较低时,幼苗的一级侧根数增多,根系的分枝密度增加,二级侧根节点之间距离减小,细根的特定根长增加,这些根系结构特征的变化有利于幼苗吸收更多的养分和水分。P养分供应适宜时,幼苗增加了地上部分的分枝数,以争取更多的地上资源空间。     

Nitrogen limitation of net primary productivity in terrestrial ecosystems is globally distributed

Our meta-analysis of 126 nitrogen addition experiments evaluated nitrogen (N) limitation of net primary production (NPP) in terrestrial ecosystems. We tested the hypothesis that N limitation is widespread among biomes and influenced by geography and climate. We used the response ratio (R approximately equal ANPP(N)/ANPP(ctrl)) of aboveground plant growth in fertilized to control plots and found that most ecosystems are nitrogen limited with an average 29% growth response to nitrogen (i.e., R = 1.29). The response ratio was significant within temperate forests (R = 1.19), tropical forests (R = 1.60), temperate grasslands (R = 1.53), tropical grasslands (R = 1.26), wetlands (R = 1.16), and tundra (R = 1.35), but not deserts. Eight tropical forest studies had been conducted on very young volcanic soils in Hawaii, and this subgroup was strongly N limited (R = 2.13), which resulted in a negative correlation between forest R and latitude. The degree of N limitation in the remainder of the tropical forest studies (R = 1.20) was comparable to that of temperate forests, and when the young Hawaiian subgroup was excluded, forest R did not vary with latitude. Grassland response increased with latitude, but was independent of temperature and precipitation. These results suggest that the global N and C cycles interact strongly and that geography can mediate ecosystem response to N within certain biome types.     

Nutrient limitation in soils exhibiting differing nitrogen availabilities: what lies beyond nitrogen saturation?

The nature of nutrient limitation in large areas of temperate forest may be changing due to human activities. As N availability in these forests increases, other nutrients could increasingly constrain productivity and other ecosystem processes. To determine the nature of nutrient limitation (N, P, and Ca) in forest soils exhibiting differing N availability, we conducted three field studies in the Fernow Experimental Forest, West Virginia, USA. The first used a ubiquitous herbaceous species, Viola rotundifolia, to compare indices of N availability to the activity of root-associated phosphomonoesterase (PME) activity at two spatial scales. The second study used fertilized, root in-growth cores to assess the extent of N, P, and Ca limitation. Finally, we measured the root-associated PME activity of V. rotundifolia growing in experimental plots that have received various combinations of nutrient additions and harvest treatments. For entire watersheds, stream water nitrate concentrations were positively related to PME activities (R2 = 0.986). For small plots, PME activities were positively associated with soil nitrate availability (R2 = 0.425), and to a lesser extent with the leaf N concentrations (R2 = 0.291). Root growth into microsites fertilized with P was greater than growth into microsites fertilized with either N or Ca, especially in watersheds with high N availability. Experimental additions of N increased the root-associated PME activity of V. rotundifolia, supporting the causality of the relationship between N availability and PME activity. Collectively, our results indicate that, as N availability increases, P becomes increasingly limiting at the sites examined. Understanding how nutrient limitations change during N saturation should improve ecosystem models and better inform our attempts to mitigate any undesired effects.