Nitrogen Fixation in Bryophytes, Lichens, and Decaying Wood along a Soil-age Gradient in Hawaiian Montane Rain Forest

We determined rates of acetylene reduction and estimated total nitrogen fixation associated with bryophytes, lichens, and decaying wood in Hawaiian montane rain forest sites with underlying substrate ranging in age from 300 to 4.1 million years. Potential N fixation ranged from ca 0.2 kg/ha annually in the 300‐year‐old site to ca 1 kg/ha annually in the 150,000‐year‐old site. Rates of acetylene reduction were surprisingly uniform along the soil‐age gradient, except for high rates in symbiotic/associative fixers at the 150,000‐year‐old site and in heterotrophic fixers at the 2100‐year‐old site. Low fixation at the youngest site, where plant production is known to be N‐limited, suggests that demand for N alone does not govern N fixation. Total N fixation was highest in sites with low N:P ratios in leaves and stem wood, perhaps because epiphytic bryophytes and lichens depend on canopy leachate for mineral nutrients and because heterotrophic fixation is partly controlled by nutrient supply in the decomposing substrate; however, differences in substrate cover, rather than in fixation rates, had the largest effect on the total N input from fixation at these sites.     

Nutrient limitation and soil development: Experimental test of a biogeochemical theory

Walker & Syers (1976) proposed a conceptual model that describesthe pattern and regulation of soil nutrient pools and availability during long-term soil and ecosystem development. Their model implies that plantproduction generally should be limited by N on young soils and by P on oldsoils; N and P supply should more or less equilibrate onintermediate-aged soils. We tested the application of this model to nutrientlimitation, using a well characterized substrate age sequence in Hawaiianmontane rain forest. Earlier experiments had evaluated nutrient limitationin forests on a young (300 y) and an old (4,100,000 y) substrate on the samedevelopmental sequence; N alone limited tree growth on the youngsubstrate, while P alone did so on the old one. An additional fertilizerexperiment based on replicated treatments with N, P, and all othernutrients combined, applied in individually and in all factorialcombinations, was established in an intermediate-aged site in theLaupahoehoe Forest Reserve, Hawaii. Here, diameter increments of thedominant tree Metrosideros polymorpha increased slightly with Nadditions, and nearly doubled when N and P were added together.Additions of elements other than N and P had no significant effecton growth. These results show that N and P had equilibrated (relativeto plant requirements) in the intermediate aged site. Together withthe earlier experiments, these results suggest that the Walker and Syersmodel provides a useful starting point for explaining the nature anddistribution of nutrient limitation in forest ecosystems.     

Nutrient limitations to plant growth during primary succession in Hawaii Volcanoes National Park

We determined the effects of nutrient amendments on plant growth in three tropical montane rainforest sites representing a sequence of soil ages (< 30, 200, and 2000 y). Factorial fertilization with nitrogen, phosphorus, and all other essential nutrients (combined) was applied to the two younger sites; only nitrogen was applied to the oldest one. Nitrogen supply represented the most important limitation to plant growth in the two younger sites; additions of nitrogen caused significant increases in tree diameter increment, height growth, litterfall, and most other growth-related parameters. In contrast, nitrogen additions had no significant effect on plant growth in the oldest site. Phosphorus additions increased extractable soil phosphorus and plant tissue phosphorus, but did not increase plant growth at the young sites. The results are consistent with Walker & Syers' (1976) model for the control of nutrient limitation during soil development.     

Relationships among net primary productivity, nutrients and climate in tropical rain forest: a pan-tropical analysis

Tropical rain forests play a dominant role in global biosphere-atmosphere CO(2) exchange. Although climate and nutrient availability regulate net primary production (NPP) and decomposition in all terrestrial ecosystems, the nature and extent of such controls in tropical forests remain poorly resolved. We conducted a meta-analysis of carbon-nutrient-climate relationships in 113 sites across the tropical forest biome. Our analyses showed that mean annual temperature was the strongest predictor of aboveground NPP (ANPP) across all tropical forests, but this relationship was driven by distinct temperature differences between upland and lowland forests. Within lowland forests (相似文献   

氮添加诱导的磷限制改变了亚热带黄山松林土壤微生物群落结构

土壤微生物在陆地生态系统的生物地球化学循环中起着重要作用。然而目前尚不清楚氮(N)添加量及其持续时间如何影响土壤微生物群落结构,以及微生物群落结构变化与微生物相对养分限制状况是否存在关联。本研究在亚热带黄山松林开展了N添加试验以模拟N沉降,并设置3个处理:对照(CK, 0 kg N·hm^-2·a^-1)、低N(LN, 40 kg N·hm^-2·a^-1)和高N(HN, 80 kg N·hm^-2·a^-1)。在N添加满1年和3年时测定土壤基本理化性质、磷脂脂肪酸含量和碳(C)、N、磷(P)获取酶活性,并通过生态酶化学计量分析土壤微生物的相对养分限制状况。结果表明: 1年N添加对土壤微生物群落结构无显著影响,3年LN处理显著提高了革兰氏阳性菌(G⁺)、革兰氏阴性菌(G^-)、放线菌(ACT)和总磷脂脂肪酸(TPLFA)含量,而3年HN处理对微生物的影响不显著,表明细菌和ACT对N添加可能更为敏感。N添加加剧了微生物C和P限制,而P限制是土壤微生物群落结构变化的最佳解释因子。这表明,N添加诱导的P限制可能更有利于部分贫营养菌(如G⁺)和参与P循环的微生物(如ACT)的生长,从而改变亚热带黄山松林土壤微生物群落结构。     

海南岛热带山地雨林幼苗幼树光合与叶氮、叶磷及比叶面积的关系

以海南岛吊罗山热带山地雨林101个物种的幼苗幼树为试验材料,测定其光合、叶片氮、磷含量及比叶面积;检验其相关关系,并按乔木,乔灌木(小乔木至大灌木)和灌木3个生活型组进行分组检验。研究结果表明,单位叶面积(Aarea)和单位叶重量的光合速率(Amass)均表现出灌木>乔木>乔灌木,方差分析表明,灌木和乔灌木之间Aarea差异显著;灌木和乔木以及灌木和乔灌木之间Amass差异显著(p<0.05)。Aarea与叶氮含量之间的相关性在不同生态型组和所有物种之间均达到极显著水平(p<0.0001);与叶磷之间的相关关系在灌木(p=0.0038),乔灌木(p=0.0002)以及所有物种(p<0.0001)之间达到极显著水平,但是在乔木中未达到显著水平(p>0.05);与SLA之间在灌木(p=0.0006)、乔木(p<0.0001)和所有物种(p<0.0001)之间达到极显著水平,但是在乔灌木中未达到显著水平(p>0.05)。Amass与叶片氮含量、SLA的相关关系在不同生活型组和所有物种中都达到极显著水平(p<0.0001);与叶磷含量之间的相关性在灌木(p=0.0004),乔灌木(p=0.0018)及所有物种(p<0.0001)中极显著,在乔木生活型组中也达显著水平(p=0.0377)。逐步回归表明,与Aarea相比,Amass估计结果更接近于实际测值。由此可见,海南岛热带山地雨林林下幼苗幼树的光合和叶氮、磷含量及SLA之间相关关系与基于成树的研究非常相似,并且A比A更能稳定体现这种相关性。     

Functional traits determine tree growth and ecosystem productivity of a tropical montane forest: Insights from a long‐term nutrient manipulation experiment

Trait‐response effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long‐term tree growth in experimental nutrient addition plots (N, P, and N + P) varied as a function of morphological traits, tree size, and species identity. We also evaluated how trait‐based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N + P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N + P grew faster irrespective of their initial size relative to trees in control or to trees in other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long‐term experimental evidence that trait‐based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.     

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.     

Variation in Canopy Litterfall Along a Precipitation and Soil Fertility Gradient in a Panamanian Lower Montane Forest

Fertilization experiments in tropical forests have shown that litterfall increases in response to the addition of one or more soil nutrients. However, the relationship between soil nutrient availability and litterfall is poorly defined along natural soil fertility gradients, especially in tropical montane forests. Here, we measured litterfall for two years in five lower montane 1‐ha plots spanning a soil fertility and precipitation gradient in lower montane forest at Fortuna, Panama. Litterfall was also measured in a concurrent nitrogen fertilization experiment at one site. Repeated‐measures ANOVA was used to test for site (or treatment), year, and season effects on vegetative, reproductive and total litterfall. We predicted that total litterfall, and the ratio of reproductive to leaf litterfall, would increase with nutrient availability along the fertility gradient, and in response to nitrogen addition. We found that total annual litterfall varied substantially among 1‐ha plots (4.78 Mg/ha/yr to 7.96 Mg/ha/yr), and all but the most aseasonal plot showed significant seasonality in litterfall. However, litterfall accumulation did not track soil nutrient availability; instead forest growing on relatively infertile soil, but dominated by an ectomycorrhizal tree species, had the highest total litterfall accumulation. In the fertilization plots, significantly more total litter fell in nitrogen addition relative to control plots, but this increase in response to nitrogen (13%) was small compared to variation observed among 1‐ha plots. These results suggest that while litterfall at Fortuna is nutrient‐limited, compositional and functional turnover along the fertility gradient obscure any direct relationship between soil resource availability and canopy productivity.     

Prediction of forest aboveground net primary production from high‐resolution vertical leaf‐area profiles

Temperature and precipitation explain about half the variation in aboveground net primary production (ANPP) among tropical forest sites, but determinants of remaining variation are poorly understood. Here, we test the hypothesis that the amount of leaf area, and its vertical arrangement, predicts ANPP when other variables are held constant. Using measurements from airborne lidar in a lowland Neotropical rain forest, we quantify vertical leaf‐area profiles and develop models of ANPP driven by leaf area and other measurements of forest structure. Vertical leaf‐area profiles predict 38% of the variation among plots. This number is 4.5 times greater than models using total leaf area (disregarding vertical arrangement) and 2.1 times greater than models using canopy height alone. Furthermore, ANPP predictions from vertical leaf‐area profiles were less biased than alternate metrics. Variation in ANPP not attributable to temperature or precipitation can be predicted by the vertical distribution of leaf area in this system.     

Both nitrogen and phosphorus limit plant production on young Hawaiian lava flows

We applied fertilizers in a 2³complete factorial design to determine the effects of nutrient amendments on plant growth in Hawaiian montane forests growing on two different volcanic substrates: a and phoehoe lava. Both sites were about 140 years old and their overstories were nearly monospecific stands of Metrosideros polymorpha. Fertilizer applications included N, P, a mixture of essential macro- and micronutrients excepting P and N, and all combinations thereof in each of four blocks. Additions of nutrients other than N or P had no significant effects on measured plant-growth variables. In contrast, additions of either N or P significantly increased tree height growth, diameter increments, biomass growth, and height growth of the understory fern Dicranopteris linearis in both sites. The effect of N was greater than that of P. Greatest growth rates occurred in plots receiving both N and P, and signficant N^*P interactions occurred in several cases, suggesting a synergistic effect between these two elements. Plant growth on these young, poorly weathered, basaltic lavas is colimited by N and P availability. Growth in a similar-aged stand growing on a mixture of volcanic ash and cinders is N but not P limited, indicating that the texture of the parent material influences nutrient-availability patterns during early primary succession.     

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。     

Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems

The cycles of the key nutrient elements nitrogen (N) and phosphorus (P) have been massively altered by anthropogenic activities. Thus, it is essential to understand how photosynthetic production across diverse ecosystems is, or is not, limited by N and P. Via a large-scale meta-analysis of experimental enrichments, we show that P limitation is equally strong across these major habitats and that N and P limitation are equivalent within both terrestrial and freshwater systems. Furthermore, simultaneous N and P enrichment produces strongly positive synergistic responses in all three environments. Thus, contrary to some prevailing paradigms, freshwater, marine and terrestrial ecosystems are surprisingly similar in terms of N and P limitation.     

Nutrient limitation of phytoplankton in a central Arizona reservoir

The influence of added nutrients nitrogen, phosphorus and carbon on the phytoplankton of a small recreational reservoir in central Arizona was investigated during the summer, 1974. Polyethylene bags were used to isolate lake water and the natural populations for the addition of nitrogen, phosphorus and carbon individually and in combination. A large increase in phytoplankton numbers, extractable chlorophyll, pH and dissolved oxygen occurred only in bags to which both nitrogen and phosphorus were added, suggesting that both nitrogen and phosphorus levels were limiting to the primary producers. Little alteration in species composition resulted from the addition of the above nutrients.     

Phosphorus limitation on photosynthesis of two dominant understory species in a lowland tropical forest

Aims Elevated nitrogen (N) deposition in tropical regions may accelerate ecosystem phosphorus (P) limitation. However, it is not explicitly addressed that how changes in soil N and P availability affect foliar nutrients and photosynthesis of plants in tropical forests. In this study, we examined the effects of N and P additions on foliar nutrients and net photosynthesis of two dominant understory species, Randia canthioides (R. canthioides) and Cryptocarya concinna (C. concinna) in an N-saturated old-growth tropical forest (>400-year-old) in southern China.Methods A full factorial NP addition experiment (2×2) was established in 2007 and continued through August 2010. Four treatments, including control, N addition (150kg N ha^-1 year^-1), P addition (150kg P ha^-1 year^-1) and NP addition (150kg N ha^-1 year^-1 plus 150kg P ha^-1 year^-1) were set up in this experiment. Photosynthetic traits (maximum photosynthetic CO₂ assimilation (A max), stomatal conductance (g s), leaf transpiration (E), light saturating point, concentrations of chlorophyll a/b and foliar nutrients (N and P) of the two species were measured with standard methods.Important findings Three years of N addition had no significant effects on any measured photosynthetic parameter of either species. However, N addition significantly elevated foliar N and P concentrations of one species (R. canthioides), resulting in lower photosynthetic nitrogen use efficiency (PNUE). N treatments decreased foliar P concentration of the other (C. concinna), resulting in increased photosynthetic phosphorus use efficiency, which was potentially related to N-induced P shortage. In contrast, positive effects of P treatments on g s of R. canthioides, A max and chlorophyll a+b of C. concinna were observed. P treatments also elevated foliar P and PNUE of both species, implying P induced more efficient use of N. Our results suggested a more important role of P than N on influencing photosynthetic traits of these two understory species. Alleviation of P shortage through P addition may enhance photosynthetic performances of some understory species in N-rich tropical forests.     

Phosphorus Fertilization Increases the Abundance and Nitrogenase Activity of the Cyanolichen Pseudocyphellaria crocata in Hawaiian Montane Forests

Nitrogen-fixing epiphytes (especially lichens with a cyanobacterial symbiont—cyanolichens) have the potential to contribute significant amounts of nitrogen (N) to montane tropical forests, which are typically low in N—but the factors controlling the abundance and distribution of epiphytic cyanolichens are poorly understood. In long-term fertilization experiments in montane forests on a 4.l million-yr-old Oxisol on the island of Kauà'i and on a 152-yr-old lava flow on the island of Hawai'i, the epiphytic cyanolichen Pseudocyphellaria crocata increased significantly in abundance in canopies of host trees fertilized with phosphorus (P). There was no response to fertilization with N or other essential elements. Nitrogen-fixation rates were also elevated in lichens in P-fertilized plots at both sites. Phosphorus supply to host trees may be an important factor controlling N inputs to montane tropical forests by N-fixing epiphytes.     

亚热带次级森林演替过程中模拟氮磷沉降对土壤微生物生物量及土壤养分的影响

氮（N）沉降深刻影响着森林生态系统的生物多样性、生产力和稳定性。亚热带地区森林土壤磷（P）的有效性较低,N沉降将更突显P的限制作用。N、P输入对亚热带次级森林土壤的影响是否依赖于森林演替阶段知之甚少。选取两种不同演替年龄阶段（年轻林：<40 a;老年林：>85 a）的亚热带常绿阔叶林,设置模拟N和/或P沉降（10 g m^-2 a^-1）4个处理（Ctrl、N、P、NP）,连续处理4.5年后采集表层、次表层和下底层（0-15、15-30、30-60 cm）土壤样品,综合分析了土壤微生物生物量碳（MBC）氮（MBN）和多种土壤养分含量。结果表明,MBC、MBN及土壤养分含量均随土壤深度增加而降低。N添加对两种演替阶段森林土壤中MBC和MBN均无显著影响。施P相关处理（P和NP）对年轻林表层土壤MBC和MBN无显著影响,但显著增加了老年林表层土壤MBC和MBN（P<0.05）,表明老年林可能比年轻林更易受P限制。N添加显著增加了两种演替森林表层土壤可溶性有机氮（DON）、氨态氮（NH₄⁺-N）和硝态氮（NO₃^--N）的含量（P<0.05）;P相关处理（P和NP）显著增加两种演替阶段表层和次表层土壤速效磷（AP）以及表层土壤全磷（TP）的含量（P<0.05）。土壤MBC和MBN与土壤中各养分指标（可溶性有机碳DOC、DON、NH₄⁺-N、NO₃^--N、AP、全碳TC、全氮TN和TP）呈显著正相关关系,土壤TC、TN和DOC是影响土壤微生物生物量的主要因子。研究可为评估和揭示未来全球环境变化背景下不同演替林龄亚热带森林的土肥潜力及土壤质量的演变提供一定的科学理论依据。     

Long-term nitrogen deposition inhibits soil priming effects by enhancing phosphorus limitation in a subtropical forest

It is widely accepted that phosphorus (P) limits microbial metabolic processes and thus soil organic carbon (SOC) decomposition in tropical forests. Global change factors like elevated atmospheric nitrogen (N) deposition can enhance P limitation, raising concerns about the fate of SOC. However, how elevated N deposition affects the soil priming effect (PE) (i.e., fresh C inputs induced changes in SOC decomposition) in tropical forests remains unclear. We incubated soils exposed to 9 years of experimental N deposition in a subtropical evergreen broadleaved forest with two types of ¹³C-labeled substrates of contrasting bioavailability (glucose and cellulose) with and without P amendments. We found that N deposition decreased soil total P and microbial biomass P, suggesting enhanced P limitation. In P unamended soils, N deposition significantly inhibited the PE. In contrast, adding P significantly increased the PE under N deposition and by a larger extent for the PE of cellulose (PE_cellu) than the PE of glucose (PE_glu). Relative to adding glucose or cellulose solely, adding P with glucose alleviated the suppression of soil microbial biomass and C-acquiring enzymes induced by N deposition, whereas adding P with cellulose attenuated the stimulation of acid phosphatase (AP) induced by N deposition. Across treatments, the PE_glu increased as C-acquiring enzyme activity increased, whereas the PE_cellu increased as AP activity decreased. This suggests that P limitation, enhanced by N deposition, inhibits the soil PE through varying mechanisms depending on substrate bioavailability; that is, P limitation regulates the PE_glu by affecting soil microbial growth and investment in C acquisition, whereas regulates the PE_cellu by affecting microbial investment in P acquisition. These findings provide new insights for tropical forests impacted by N loading, suggesting that expected changes in C quality and P limitation can affect the long-term regulation of the soil PE.     

Soil nutrient dynamics in response to irrigation of a Panamanian tropical moist forest

We measured concentrations of soil nutrients (0–15 and 30–35 cm depths) before and after the dry season in control and dry-season irrigated plots of mature tropical moist forest on Barro Colorado Island (BCI) in central Panama to determine how soil moisture affects availability of plant nutrients. Dry-season irrigation (January through April in 1986, 1987, and 1988) enhanced gravimetric soil water contents to wet-season levels (ca. 400 g kg^–1 but did not cause leaching beyond 0.8 m depth in the soil. Irrigation increased concentrations of exchangeable base cations (Ca²⁺, Mg²⁺, K⁺, Na⁺), but it had little effect on concentrations of inorganic N (NH₄ ^+C, NO₃ ^– and S (SO₄ ^2–). These BCI soils had particularly low concentrations of extractable P especially at the end of the dry season in April, and concentrations increased in response to irrigation and the onset of the rainy season. We also measured the response of soil processes (nitrification and S mineralization) to irrigation and found that they responded positively to increased soil moisture in laboratory incubations, but irrigation had little effect on rates in the field. Other processes (plant uptake, soil organic matter dynamics) must compensate in the field and keep soil nutrient concentrations at relatively low levels.