Transferring soils from high- to low-elevation forests increases nitrogen cycling rates: climate change implications

It is well recognized that photosynthesis of C3 plants is highly responsive to CO₂ concentration. However, in natural ecosystems, plants are subject to a range of feed-back effects that can interact with increased photosynthetic carbon gain in different ways so that it is not clear to what extent increased photosynthesis will translate into increased growth. To assess the probable growth response of nutrient-limited forests to increasing CO₂ concentration, we use a previously developed modelling framework and apply it under conditions where the supply of nutrients is affected by a range of different factors. Our analysis indicates that forest growth is likely to be highly stimulated by increasing CO₂ concentration in forests with high fertility, in forests with nitrogen fixing plants, in those subject to fire or where nitrogen in wood is effectively removed from the biologically active cycle either through physical removal of stems in harvesting or through continued stem growth over long time periods. Forest growth is likely to be stimulated by CO₂ concentration in both phosphorus- and sulphur-limited forests provided nutrients in heartwood of trees are removed from the active nutrient cycle. Without this removal from the cycling system, however, sulphur-limited forests should show little response to increasing CO₂. In phosphorus-limited forests without phosphorus removal, the response to increasing CO₂ depends further on the equilibration state of the large pool of unavailable secondary phosphorus. Considered over periods of centuries during which the secondary pool has equilibrated, growth of phosphorus-limited forests is likely to be only weakly stimulated by increasing CO₂ concentration. However, over shorter periods, increasing CO₂ concentration should lead to a substantial increase in productivity. In general, it can be concluded that systems that are more open with respect to nutrient gains and losses are likely to be more responsive to increasing CO₂ concentration than systems where the amount of available nutrients is less variable. In more open systems, operation at a lower internal nutrient concentration as a result of increasing atmospheric CO₂ concentration can lead to reduced nutrient losses per unit carbon gain. Our analysis shows that the effect of increasing CO₂ on forest growth can differ substantially between forests due to interactions with a range of factors that affect nutrient supply. The response of a particular forest to increasing CO₂ concentration can only be predicted if the main factors controlling nutrient supply and growth in that forest are understood and incorporated into an assessment.     

Potential impacts of climate change on nitrogen transformations and greenhouse gas fluxes in forests: a soil transfer study

Relatively little research has been conducted on how climate change may affect the structure and function of arid to semiarid ecosystems of the American Southwest. Along the slopes of the San Francisco Peaks of Arizona, USA, I transferred intact soil cores from a spruce‐fir to a ponderosa pine forest 730 m lower in elevation to assess the potential impacts of climate change on soil N cycling and trace gas fluxes. The low‐elevation site has a mean annual soil temperature about 2.5°C higher than the high‐elevation site. Net rates of N transformations and trace gas fluxes were measured in high‐elevation soil cores incubated in situ and soil cores transferred to the low‐elevation site. Over a 13‐month period, volumetric soil water content was similar in transferred soil cores relative to soil cores incubated in situ. Net N mineralization and nitrification increased over 80% in transferred soil cores compared with in situ soil cores. Soil transfer significantly increased net CO₂ efflux (120%) and net CH₄ consumption (90%) relative to fluxes of these gases from soil cores incubated in situ. Soil net N₂O fluxes were relatively low and were not generally altered by soil transfer. Although the soil microbial biomass as a whole decreased in transferred soil cores compared with in situ soil cores after the incubation period, active bacterial biomass increased. Transferring soil cores from the low‐elevation to the high‐elevation site (i.e. simulated global cooling) commonly, but not consistently, resulted in the opposite effects on soil pools and processes. In general, soil containment (root trenching) did not significantly affect soil measurements. My results suggest that small increases in mean annual temperature can have large impacts on soil N cycling, soil–atmosphere trace gas exchanges, and soil microbial communities even in ecosystems where water availability is a major limiting resource.     

森林土壤氮素转换及其对氮沉降的响应

近几十年人类活动向大气中排放的含氮化合物激增 ,并引起大气氮沉降也成比例增加。目前 ,氮沉降的增加使一些森林生态系统结构和功能发生改变 ,甚至衰退。近 2 0 a欧洲和北美有关氮沉降及其对森林生态系统的影响方面的研究较多 ,而我国少有涉及。森林土壤氮素转换是森林生态系统氮素循环的一个重要的组成部分 ,而矿化、硝化和反硝化作用是其核心过程 ,氮沉降作为驱动因子势必改变森林土壤氮素转换速度、方向和通量。根据国外近 2 0 a有关研究 ,首先介绍了森林土壤氮素转换过程和强度 ,论述森林土壤氮素在生态系统氮素循环中的作用 ,然后在此基础上 ,介绍了氮沉降对森林土壤氮素循环的研究途径 ,探讨了氮沉降对森林土壤氮素矿化、硝化和反硝化作用的影响及其机理     

四种温带森林土壤氮矿化与硝化时空格局

利用PVC管原位培养连续取样法测定了东北地区4种具有代表性的森林生态系统(硬阔叶林、蒙古栎林、红松林、落叶松林)土壤氮素矿化、硝化的时间动态及氮矿化的空间分布格局.结果表明:4种森林土壤氮素矿化存在明显的时空变异.蒙古栎和红松林土壤在6月份表现出强烈的氮矿化和硝化作用,而硬阔叶林及落叶松林7月份氮素矿化强烈.4种森林生态系统上层土壤的氮净矿(硝)化率显著高于下层土壤.4种林型土壤的硝化过程在氮矿化过程中占有重要地位,其NO-3-N在无机氮中的比例分别为:79.9%～91.1%(硬阔叶林)、50.7%～80.5%(蒙古栎林)、54.1%～92.0%(红松林)、63.7%～86.5%(落叶松林).生态系统构成决定了土壤氮素的矿化能力.阔叶林和针阔混交林生态系统矿化率大于纯针叶林生态系统.硬阔叶林、红松林、蒙古栎林、落叶松林的平均净矿化率分别为:(0.58±0.01) mg · kg-1 · d-1、(0.47±0.19) mg · kg-1 · d-1、(0.39±0.11) mg · kg-1 · d-1和(0.23±0.06) mg · kg-1 · d-1.4种林型氮素矿化作用与地下5 cm温度呈正相关,并受土壤表层 (0～10 cm)水分显著影响.土壤微生物量氮与土壤氮矿化呈显著正相关.     

Winter climate change affects growing‐season soil microbial biomass and activity in northern hardwood forests

Understanding the responses of terrestrial ecosystems to global change remains a major challenge of ecological research. We exploited a natural elevation gradient in a northern hardwood forest to determine how reductions in snow accumulation, expected with climate change, directly affect dynamics of soil winter frost, and indirectly soil microbial biomass and activity during the growing season. Soils from lower elevation plots, which accumulated less snow and experienced more soil temperature variability during the winter (and likely more freeze/thaw events), had less extractable inorganic nitrogen (N), lower rates of microbial N production via potential net N mineralization and nitrification, and higher potential microbial respiration during the growing season. Potential nitrate production rates during the growing season were particularly sensitive to changes in winter snow pack accumulation and winter soil temperature variability, especially in spring. Effects of elevation and winter conditions on N transformation rates differed from those on potential microbial respiration, suggesting that N‐related processes might respond differently to winter climate change in northern hardwood forests than C‐related processes.     

Response of nitrogen cycling to simulated climate change: differential responses along a subalpine ecotone

In situ nitrogen (N) transformations and N availability were examined over a four‐year period in two soil microclimates (xeric and mesic) under a climate‐warming treatment in a subalpine meadow/sagebrush scrub ecotone. Experimental plots that spanned the two soil microclimates were exposed to an in situ infrared (IR) climate change manipulation at the Rocky Mountain Biological Laboratory, near Crested Butte, Colorado. Although the two microclimates did not differ significantly in their rates of N transformations in the absence of heating, they differed significantly in their response to increased IR. Under a simulated warming in the sagebrush‐dominated xeric microclimate, gross N mineralization rates doubled and immobilization rates increased by up to 60% over the first 2 years of the study but declined to predisturbance rates by the fourth year. This temporal pattern of gross mineralization rates correlated with a decline in SOM. Concurrently, rates of net mineralization rates in the heated plots were 60% higher than the controls after the first year. There were no differences in gross or net nitrification rates with heating in the xeric soils. In contrast to the xeric microclimate, there were no significant effects of heating on any N transformation rates in the mesic microclimate. The differing responses in N cycling rates of the two microclimate to the increased IR is most certainly the result of differences in initial soil moisture conditions and vegetation type and cover.     

森林生态系统土壤氮矿化影响因素研究进展

森林生态系统土壤氮矿化是生态系统中最重要的功能之一,综述了近10余年来森林生态系统土壤氮矿化影响因素的研究,在前人的基础上将其影响因素归成3类;（1）环境因子,（2）凋落物质量,（3）土壤动物和微生物,其中环境因子中的土壤温、湿度是影响土壤氮矿化的最重要因子,氮素可利用性、氮转化与群落演替、植物多样性间相互关系的研究正受到愈来愈多的重视,研究CO2倍增及其引起的全球变暖对土壤氮素转化的潜在影响也已成为当前全球变化问题研究的热点之一。     

Effect of temperature on humus respiration rate and nitrogen mineralization: Implications for global climate change

Respiration and nitrogen mineralization rates of humus samples from 7 Scots pine stands located along a climatic transect across the European continent from the Pyrenees (42°40) to northern Sweden (66°08) were measured for 14 weeks under laboratory conditions at temperatures from 5 °C to 25 °C. The average Q₁₀ values for the respiration rate ranged from about 1.0 at the highest temperature to more than 5 at 10 °C to 15 °C in the northernmost samples. In samples from more northern sites, respiration rates remained approximately constant during the whole incubation period; in the southern end of the transect, rates decreased over time. Respiration rate was positively correlated with incubation temperature, soil pH and CN ratio, and negatively with soil total N. Regressions using all these variables explained approximately 71% of the total variability in the respiration rate. There was no clear relation between the nitrogen mineralization rate and incubation temperature. Below 15 °C the N-mineralization rate did not respond to increasing temperature; at higher temperatures, significant increases were found for samples from some sites. A regression model including incubation temperature, pH, N_tot and CN explained 73% of the total variability in N mineralization. The estimated increase in annual soil respiration rates due to predicted global warming at the high latitudes of the Northern Hemisphere ranged from approximately 0.07×10¹⁵ to 0.13×10¹⁵ g CO₂ at 2 °C and 4 °C temperature increase scenarios, respectively. Both values are greater than the current annual net carbon storage in northern forests, suggesting a switch of these ecosystems from net sinks to net sources of carbon with global warming.     

生物炭配施硝化/脲酶抑制剂对亚热带水稻土活性氮气体排放的影响

探究施用生物炭和脲酶抑制剂/硝化抑制剂对亚热带水稻土氮素硝化过程的调控作用、氨挥发和N₂O排放的温室效应潜能的影响,确定生物炭与硝化和脲酶抑制剂的最佳组合,可为削减施用氮肥带来的活性氮气体排放对环境的负面风险提供理论依据。本研究采用室内好气培养试验方式,以单施尿素(N)为对照,设置7个试验处理[尿素+生物炭(NB),尿素+硝化抑制剂(N+NI),尿素+脲酶抑制剂(N+UI),尿素+硝化抑制剂+脲酶抑制剂(N+NIUI),尿素+硝化抑制剂+生物炭(NB+NI),尿素+脲酶抑制剂+生物炭(NB+UI),尿素+硝化抑制剂+脲酶抑制剂+生物炭(NB+NIUI)],观测生物炭与脲酶抑制剂(NBPT)/硝化抑制剂(DMPP)配施下土壤无机氮含量、N₂O排放及氨挥发的变化动态。结果表明: 1)培养期间,与N处理(5.11 mg N·kg^-1·d^-1)相比,NB处理的土壤硝化速率常数显著增加33.9%,N+NI处理显著降低22.9%;NB处理显著提高了氨氧化细菌(AOB)丰度,增幅达56.0%。2)与N处理相比,N+NI和NB+NI处理的NH₃累积排放量均显著增加约49%;N+UI处理降低了NH₃累积损失量,NB+UI处理抑制效果更明显。3)各处理的N₂O排放速率高峰均出现在施肥后前10 d;NB处理的N₂O排放高峰出现最早,N处理排放速率最高(5.87 μg·kg^-1·h^-1);硝化抑制剂与脲酶抑制剂配施减少土壤N₂O排放的效果最佳。综合计算各处理直接N₂O和间接N₂O(NH₃)排放产生的温室效应潜能(GWP)发现,N+NI和NB+NI处理较N处理分别增加了34.8%和40.9%,而NB和NB+UI处理的GWP显著降低了45.9%和60.5%。因此,生物炭与脲酶抑制剂配施对降低土壤活性氮气体排放所产生的温室效应潜能效果最佳。     

雪被斑块对川西亚高山两个森林群落冬季土壤氮转化的影响

为了解气候变暖情景下雪况变化对高寒森林冬季土壤氮转化的影响,测定了川西亚高山冷杉(Abies faxoniana)+红桦(Betula albo-sinensis)混交林(MF)和冷杉次生林(SF)三类雪被斑块(浅雪被、中厚度雪被和厚雪被)内冬季土壤氮矿化特征。结果表明:经过一个冬季(2011-2012),两个森林群落土壤净氮氨化量都为负值,净氮硝化量都为正值,且净氮硝化量显著高于净氮氨化量;冬季土壤氮氨化、硝化、矿化和固持量都是中度雪被厚度最高,但各雪被斑块之间都未达到显著水平。各雪被斑块下,冷杉次生林土壤氮矿化参数都显著高于针阔混交林,但雪被斑块和林型交互作用对冬季土壤氮矿化无显著影响。这表明,该区冬季土壤氮矿化以硝化过程为主,硝化和氨化过程可能受不同微生物群落调控;短时期内,未来气候变化所导致的雪被减少对该区森林冬季土壤氮转化影响可能不明显。     

氮沉降对热带亚热带森林土壤氮循环微生物过程的影响研究进展

近年来,高速的城市化和工业化建设导致全球大气氮沉降量逐年递增,其中热带亚热带地区氮沉降量显著高于全球平均水平,而大部分热带亚热带森林土壤趋近氮饱和状态,氮沉降增加将持续向土壤输入外源活性氮,极易导致土壤氮过剩,进而破环整个森林生态系统氮循环的平衡。我国热带亚热带地区经济发展快速,氮沉降增加导致的土壤养分失衡和林地退化等生态问题日益凸显,森林土壤氮循环对大气氮沉降的响应及适应机制已引起了学术界的广泛关注。研究表明氮循环各环节均由特定的功能微生物驱动完成,明确氮沉降增加对热带亚热带森林土壤氮循环功能微生物及其介导的关键过程的影响,对评价未来氮沉降增加背景下全球森林土壤氮循环的响应及驱动机制有重要作用,可为促进我国热带亚热带地区森林修复、生态环境的改善与提升提供科学支撑。鉴于此,本文综述了热带亚热带森林土壤氮循环主要过程（如固氮、硝化、反硝化、厌氧氨氧化等）及其功能微生物群落丰度、活性、组成等对氮沉降增加的响应,同时分析了这些功能微生物的群落特征与主要环境因子（如NH₄⁺、NO₃^-、有机碳、pH、含水量等）的关联性。在此基础上探讨了氮沉降增加下功能微生物对热带亚热带森林土壤氮循环的调控作用,重点探讨了功能微生物如何通过改变丰度与群落组成而影响氮循环过程,并对目前研究中存在的主要问题与未来研究重点进行了简要剖析。     

季节性冻融期间川西亚高山/高山森林土壤净氮矿化特征

气候变暖情景下季节性冻融格局的改变可能显著影响高寒森林土壤氮素矿化过程.本文采用原状土壤移位培养的方法,以海拔梯度形成的温度差异模拟气候变暖,研究了川西亚高山/高山森林在生长季节和季节性冻融期间土壤的净氮矿化量和净氮矿化速率.结果表明： 在川西亚高山/高山森林,土壤铵态氮和硝态氮含量均表现为从生长季节至冻结初期明显下降,完全冻结期明显增加,而在融化初期明显降低的变化过程.季节性冻融期土壤的净氮矿化量和净氮矿化速率显著低于生长季节,并且出现明显的氮素固持现象.与低海拔相比,中海拔森林土壤的氮素固持作用相对较大,高海拔相对较小,可能与不同海拔梯度土壤温度变化及引起的冻融循环密切相关.在生长季节,土壤净氮矿化量和矿化速率均随海拔的降低呈明显增加趋势,尤其在低海拔处土壤的氮素矿化作用最为强烈.在气候变暖背景下,温度的增加明显促进了生长季节土壤氮素矿化,并且通过提高冻融循环频次、缩短冻结时间来影响土壤氮素矿化速率.这一过程可能受到微环境的影响.     

Nitrogen transformations in two nitrogen saturated forest ecosystems subjected to an experimental decrease in nitrogen deposition

Nitrogen transformations were studied in the forest floor and mineral soil (0–5 cm) of a Douglas fir forest (Pseudotsuga menziesii (Mirb.) Franco.) and a Scots pine forest (Pinus sylvestris L.) in the Netherlands. Curren nitrogen depositions (40 and 56 kg N ha^-1 yr^-1, respectively) were reduced to natural background levels (1–2 kg N ha^-1 yr^-1) by a roof construction. The study concentrated on rates and dynamic properties of nitrogen transformations and their link with the leaching pattern and nitrogen uptake of the vegetation under high and reduced nitrogen deposition levels. Results of an in situ field incubation experiment and laboratory incubations were compared. No effect of the reduced N deposition on nitrogen transformations was found in the Douglas fir forest. In the Scots pine forest, however, during some periods of the year nitrogen transformations were significantly decreased under the low nitrogen deposition level. At low nitrogen inputs a net immobilization occurred during most of the year leading to a very small net mineralization for the whole year. In laboratory and in individual field plots nitrogen transformations were negatively correlated with initial inorganic nitrogen concentrations. Nitrogen budget estimates showed that nitrogen transformations were probably underestimated by the in situ incubation technique. Nevertheless less nitrogen was available for plant uptake and leaching at the low deposition plots.     

Laboratory incubations reveal potential responses of soil nitrogen cycling to changes in soil C and N availability in Mojave Desert soils exposed to elevated atmospheric CO2

Elevated atmospheric carbon dioxide (CO₂) has the potential to alter soil carbon (C) and nitrogen (N) cycling in arid ecosystems through changes in net primary productivity. However, an associated feedback exists because any sustained increases in plant productivity will depend upon the continued availability of soil N. We took soils from under the canopies of major shrubs, grasses, and plant interspaces in a Mojave Desert ecosystem exposed to elevated atmospheric CO₂ and incubated them in the laboratory with amendments of labile C and N to determine if elevated CO₂ altered the mechanistic controls of soil C and N on microbial N cycling. Net ammonification increased under shrubs exposed to elevated CO₂, while net nitrification decreased. Elevated CO₂ treatments exhibited greater fluxes of N₂O–N under Lycium spp., but not other microsites. The proportion of microbial/extractable organic N increased under shrubs exposed to elevated CO₂. Heterotrophic N₂‐fixation and C mineralization increased with C addition, while denitrification enzyme activity and N₂O–N fluxes increased when C and N were added in combination. Laboratory results demonstrated the potential for elevated CO₂ to affect soil N cycling under shrubs and supports the hypothesis that energy limited microbes may increase net inorganic N cycling rates as the amount of soil‐available C increases under elevated CO₂. The effect of CO₂ enrichment on N‐cycling processes is mediated by its effect on the plants, particularly shrubs. The potential for elevated atmospheric CO₂ to lead to accumulation of NH₄⁺ under shrubs and the subsequent volatilization of NH₃ may result in greater losses of N from this system, leading to changes in the form and amount of plant‐available inorganic N. This introduces the potential for a negative feedback mechanism that could act to constrain the degree to which plants can increase productivity in the face of elevated atmospheric CO₂.     

Changes in nitrogen cycling and retention processes in soils under spruce forests along a nitrogen enrichment gradient in Germany

A network of long-term monitoring sites on nitrogen (N) input and output of forests across Germany showed that a number of Germany's forests are subject to or are experiencing N saturation and that spruce (Picea abies) stands have high risk. Our study was aimed at (1) quantifying the changes in gross rates of microbial N cycling and retention processes in forest soils along an N enrichment gradient and (2) relating the changes in soil N dynamics to N losses. We selected spruce sites representing an N enrichment gradient (indicated by leaching : throughfall N ratios) ranging from 0.04–0.13 (low N),≤0.26 (intermediate N enrichment) to≥0.42 (highly N enriched). To our knowledge, our study is the first to report on mechanistic changes in gross rates of soil N cycling and abiotic NO₃⁻ retention under ambient N enrichment gradient. Gross N mineralization, NH₄⁺ immobilization, gross nitrification, and NO₃⁻ immobilization rates increased up to intermediate N enrichment level and somewhat decreased at highly N-enriched condition. The turnover rates of NH₄⁺ and microbial N pools increased while the turnover rates of the NO₃⁻ pool decreased across the N enrichment gradient. Abiotic immobilization of NH₄⁺ did not differ across sites and was lower than that of NO₃⁻. Abiotic NO₃⁻ immobilization decreased across the N enrichment gradient. Microbial assimilation and turnover appeared to contribute largely to the retention of NH₄⁺. The increasing NO₃⁻ deposition and decreasing turnover rates of the NO₃⁻ pool, combined with decreasing abiotic NO₃⁻ retention, possibly contributed to increasing NO₃⁻ leaching and gaseous emissions across the N enrichment gradient. The empirical relationships of changes in microbial N cycling across the N enrichment gradient may be integrated in models used to predict responses of forest ecosystems (e.g. spruce) to increasing N deposition.     

Microbial biomass and nitrogen cycling responses to fertilization and litter removal in young northern hardwood forests

The influence of site fertility on soil microbial biomass and activity is not well understood but is likely to be complex because of interactions with plant responses to nutrient availability. We examined the effects of long-term (8 yr) fertilization and litter removal on forest floor microbial biomass and N and C transformations to test the hypothesis that higher soil resource availability stimulates microbial activity. Microbial biomass and respiration decreased by 20–30 % in response to fertilization. Microbial C averaged 3.8 mg C/g soil in fertilized, 5.8 mg C/g in control, and 5.5 mg C/g in litter removal plots. Microbial respiration was 200 µg CO₂-C g^–1 d^–1 in fertilized plots, compared to 270 µg CO₂-C g^–1 d^–1 in controls. Gross N mineralization and N immobilization did not differ among treatments, despite higher litter nutrient concentrations in fertilized plots and the removal of substantial quantities of C and N in litter removal plots. Net N mineralization was significantly reduced by fertilization. Gross nitrification and NO₃ ^– immobilization both were increased by fertilization. Nitrate thus became a more important part of microbial N cycling in fertilized plots even though NH₄ ⁺ availability was not stimulated by fertilization.Soil microorganisms did not mineralize more C or N in response to fertilization and higher litter quality; instead, results suggest a difference in the physiological status of microbial biomass in fertilized plots that influenced N transformations. Respiration quotients (qCO₂, respiration per unit biomass) were higher in fertilized plots (56 µg CO₂-C mg C^–1 d^–1) than control (48 µg CO₂-C mg C^–1 d ^–1) or litter removal (45 µg CO₂-C mg C^–1 d^–1), corresponding to higher microbial growth efficiency, higher proportions of gross mineralization immobilized, and lower net N mineralization in fertilized plots. While microbial biomass is an important labile nutrient pool, patterns of microbial growth and turnover were distinct from this pool and were more important to microbial function in nitrogen cycling.     

The neotropical shrub Lupinus elegans,from temperate forests,may not adapt to climate change

Considering that their distribution is limited to altitudinal gradients along mountains that are likely to become warmer and drier, climate change poses an increased threat to temperate forest species from tropical regions. We studied whether the understorey shrub Lupinus elegans, endemic to temperate forests of west‐central Mexico, will be able to withstand the projected temperature increase under seven climate change scenarios. Seeds were collected along an altitudinal gradient and grown in a shade‐house over 7 months before determining their temperature tolerance as electrolyte leakage. The plants from colder sites tolerated lower temperatures, i.e. the temperature at which half of the maximum electrolyte leakage occurred (LT₅₀), ranged from −6.4 ± 0.7 to −2.4 ± 0.3 °C. In contrast, no pattern was found for tolerance to high temperature (LT₅₀ average 42.8 ± 0.3 °C). The climate change scenarios considered here consistently estimated an increase in air temperature during the present century that was higher for the maximum air temperature than for the mean or minimum. In particular, the anomaly from the normal maximum air temperature at the study region ranged from 2.8 °C by 2030 to 5.8 °C by 2090. In this respect, the inability of L. elegans to adapt to increasingly higher temperatures found here, in addition to a possible inhibition of reproduction caused by warmer winters, may limit its future distribution.     

Global soil nitrogen cycle pattern and nitrogen enrichment effects: Tropical versus subtropical forests

Tropical and subtropical forest biomes are a main hotspot for the global nitrogen (N) cycle. Yet, our understanding of global soil N cycle patterns and drivers and their response to N deposition in these biomes remains elusive. By a meta-analysis of 2426-single and 161-paired observations from 89 published ¹⁵ N pool dilution and tracing studies, we found that gross N mineralization (GNM), immobilization of ammonium ($I_{{\mathrm{NH}}_4}$) and nitrate ($I_{{\mathrm{NO}}_3}$), and dissimilatory nitrate reduction to ammonium (DNRA) were significantly higher in tropical forests than in subtropical forests. Soil N cycle was conservative in tropical forests with ratios of gross nitrification (GN) to $I_{{\mathrm{NH}}_4}$ (GN/$I_{{\mathrm{NH}}_4}$) and of soil nitrate to ammonium (NO₃⁻/NH₄⁺) less than one, but was leaky in subtropical forests with GN/$I_{{\mathrm{NH}}_4}$ and NO₃⁻/NH₄⁺ higher than one. Soil NH₄⁺ dynamics were mainly controlled by soil substrate (e.g., total N), but climatic factors (e.g., precipitation and/or temperature) were more important in controlling soil NO₃⁻ dynamics. Soil texture played a role, as GNM and $I_{{\mathrm{NH}}_4}$ were positively correlated with silt and clay contents, while $I_{{\mathrm{NO}}_3}$ and DNRA were positively correlated with sand and clay contents, respectively. The soil N cycle was more sensitive to N deposition in tropical forests than in subtropical forests. Nitrogen deposition leads to a leaky N cycle in tropical forests, as evidenced by the increase in GN/$I_{{\mathrm{NH}}_4}$, NO₃⁻/NH₄⁺, and nitrous oxide emissions and the decrease in $I_{{\mathrm{NO}}_3}$ and DNRA, mainly due to the decrease in soil microbial biomass and pH. Dominant tree species can also influence soil N cycle pattern, which has changed from conservative in deciduous forests to leaky in coniferous forests. We provide global evidence that tropical, but not subtropical, forests are characterized by soil N dynamics sustaining N availability and that N deposition inhibits soil N retention and stimulates N losses in these biomes.     

Non-climatic thermal adaptation: implications for species' responses to climate warming

There is considerable interest in understanding how ectothermic animals may physiologically and behaviourally buffer the effects of climate warming. Much less consideration is being given to how organisms might adapt to non-climatic heat sources in ways that could confound predictions for responses of species and communities to climate warming. Although adaptation to non-climatic heat sources (solar and geothermal) seems likely in some marine species, climate warming predictions for marine ectotherms are largely based on adaptation to climatically relevant heat sources (air or surface sea water temperature). Here, we show that non-climatic solar heating underlies thermal resistance adaptation in a rocky–eulittoral-fringe snail. Comparisons of the maximum temperatures of the air, the snail''s body and the rock substratum with solar irradiance and physiological performance show that the highest body temperature is primarily controlled by solar heating and re-radiation, and that the snail''s upper lethal temperature exceeds the highest climatically relevant regional air temperature by approximately 22°C. Non-climatic thermal adaptation probably features widely among marine and terrestrial ectotherms and because it could enable species to tolerate climatic rises in air temperature, it deserves more consideration in general and for inclusion into climate warming models.     

Nitrogen cycling in an acid forest ecosystem in the Netherlands under increased atmospheric nitrogen input

Within a long-term research project studying the biogeochemical budget of an oak-beech forest ecosystem in the eastern part of the Netherlands, the nitrogen transformations and solute fluxes were determined in order to trace the fate of atmospherically deposited NH₄ ⁺ and to determine the contribution of nitrogen transformations to soil acidification.The oak-beech forest studied received an annual input of nitrogen via throughfall and stemflow of 45 kg N ha^–1 yr^–1, mainly as NH₄ ⁺, whereas 8 kg N ha^–1 yr^–1 was taken up by the canopy. Due to the specific hydrological regime resulting in periodically occurring high groundwater levels, denitrification was found to be the dominant output flux (35 kg N ha^–1 yr^–1). N₂0 emmission rate measurements indicated that 57% of this gaseous nitrogen loss (20 kg N ha^–1 yr^–1) was as N₂O. The forest lost an annual amount of 11 kg N ha^–1 yr^–1 via streamwater output, mainly as N0₃ ^–.Despite the acid conditions, high nitrification rates were measured. Nitrification occurred mainly in the litter layer and in the organic rich part of the mineral soil and was found to be closely correlated with soil temperature. The large amount of NH₄ ⁺ deposited on the forest floor via atmospheric deposition and produced by mineralization was to a large extent nitrified in the litter layer. Almost no NH₄ ⁺ reached the subsurface soil horizons. The N0₃ ^– was retained, taken up or transformed mainly in the mineral soil. A small amount of N0₃ ^– (9 kg N ha^–1 yr^–1) was removed from the system in streamwater output. A relatively small amount of nitrogen was measured in the soil water as Dissolved Organic Nitrogen.On the basis of these data the proton budget of the system was calculated using two different approaches. In both cases net proton production rates were high in the vegetation and in the litter layer of the forest ecosystem. Nitrogen transformations induced a net proton production rate of 2.4 kmol ha^–1 yr^–1 in the soil compartment.