增温施氮对高寒草甸生产力及生物量分配的影响

在青藏高原高寒草甸区设置模拟增温和氮添加处理,研究长期增温与外源氮输入对高寒草甸群落生产及其分配的影响.结果表明:开顶箱增温装置造成小环境暖干化,即显著提高地表空气温度1.6℃,提高表层土壤温度1.4℃,降低土壤含水量4.7%.2012、2013和2014年不施氮处理下增温分别降低地上生物量61.5%、108.8%和77.1%,在高氮(40和80kg N·hm^-2·a^-1)处理下增温对群落地上生物量无显著影响,这说明增温的影响依赖于氮添加水平,且施氮补偿了增温导致的土壤氮损失.增温导致根冠比增加,2012、2013和2014年不施氮处理下增温分别增加根冠比98.6%、60.7%和97.8%.在不增温处理下,植物群落地上、地下生物量的变化率均表现出低氮(10、20 kg N·hm^-2·a^-1)促进、高氮抑制的趋势,达到饱和阈值时的氮添加剂量分别为56.0和55.5 kg N·hm^-2·a^-1;而在增温处理下,地上、地下生物量随施氮量增加呈线性增加趋势.这说明增温改变了高寒草甸生物量分配对外源氮输入的响应模式,增温导致的土壤无机氮含量变化是生物量分配模式改变的主要原因.由氮添加试验估算的高寒草甸氮饱和阈值表明,高寒草甸对氮输入的敏感性高于其他类型草地.     

Shifting grassland plant community structure drives positive interactive effects of warming and diversity on aboveground net primary productivity

Ecosystems worldwide are increasingly impacted by multiple drivers of environmental change, including climate warming and loss of biodiversity. We show, using a long‐term factorial experiment, that plant diversity loss alters the effects of warming on productivity. Aboveground primary productivity was increased by both high plant diversity and warming, and, in concert, warming (≈1.5 °C average above and belowground warming over the growing season) and diversity caused a greater than additive increase in aboveground productivity. The aboveground warming effects increased over time, particularly at higher levels of diversity, perhaps because of warming‐induced increases in legume and C4 bunch grass abundances, and facilitative feedbacks of these species on productivity. Moreover, higher plant diversity was associated with the amelioration of warming‐induced environmental conditions. This led to cooler temperatures, decreased vapor pressure deficit, and increased surface soil moisture in higher diversity communities. Root biomass (0–30 cm) was likewise consistently greater at higher plant diversity and was greater with warming in monocultures and at intermediate diversity, but at high diversity warming had no detectable effect. This may be because warming increased the abundance of legumes, which have lower root : shoot ratios than the other types of plants. In addition, legumes increase soil nitrogen (N) supply, which could make N less limiting to other species and potentially decrease their investment in roots. The negative warming × diversity interaction on root mass led to an overall negative interactive effect of these two global change factors on the sum of above and belowground biomass, and thus likely on total plant carbon stores. In total, plant diversity increased the effect of warming on aboveground net productivity and moderated the effect on root mass. These divergent effects suggest that warming and changes in plant diversity are likely to have both interactive and divergent impacts on various aspects of ecosystem functioning.     

青藏高原矮嵩草草甸地下和地上生物量分配格局及其与气象因子的关系

基于2006—2015年青海海北站10年生物量及气候因子监测数据,分析了青藏高原高寒矮嵩草草甸生物量的季节及年际动态,并探讨了气候因子对其影响。结果表明:(1)季节尺度上,高寒矮嵩草草甸地上生物量表现为单峰变化曲线,8月为其峰值点,为(345.72±27.01) g/m~2,代表了高寒草甸的地上净初级生产力。而地下根系的现存量变化较为复杂,其中5—7月呈现持续上升趋势,8月快速下降,之后9月份急剧,且各月份之间未达到显著水平(P0.05);年际尺度上,10年间高寒矮嵩草草甸地上生物量整体呈现波动增加趋势,2014年为其峰值点,达(437.12±32.01) g/m~2。地下生物量呈现波动性变化,变异较大,10年间平均值为(2566.99±138.11) g/m~2;(2)高寒草甸光合产物分配主要分布在地下,80%地下根系生物量分布于地表0—10 cm土层,且不同土层根系生物量占总地下生物量的比值在不同月份较为稳定。(3)气候因子中,大气相对湿度是影响高寒草甸地上生物量大小的主要因素;而气候因子对地下根系生物量的影响极为微弱。研究表明,高寒嵩草草甸对环境变化具有较高的自我调节能力,且高寒草甸的演化受制于人类干扰,而非气候变化。     

Precipitation and nitrogen addition enhance biomass allocation to aboveground in an alpine steppe

There are two important allocation hypotheses in plant biomass allocation: allometric and isometric. We tested these two hypotheses in an alpine steppe using plant biomass allocation under nitrogen (N) addition and precipitation (Precip) changes at a community level. An in situ field manipulation experiment was conducted to examine the two hypotheses and the responses of the biomass to N addition (10 g N m^?2 y^?1) and altered Precip (±50% precipitation) in an alpine steppe on the Qinghai–Tibetan Plateau from 2013 to 2016. We found that the plant community biomass differed in its response to N addition and reduced Precip such that N addition significantly increased aboveground biomass (AGB), while reduced Precip significantly decreased AGB from 2014 to 2016. Moreover, reduced Precip enhanced deep soil belowground biomass (BGB). In the natural alpine steppe, the allocation between AGB and BGB was consistent with the isometric hypotheses. In contrast, N addition or altered Precip enhanced biomass allocation to aboveground, thus leading to allometric growth. More importantly, reduced Precip enhanced biomass allocation into deep soil. Our study provides insight into the responses of alpine steppes to global climate change by linking AGB and BGB allocation.     

Permafrost response to temperature rise in carbon and nutrient cycling: Effects from habitat‐specific conditions and factors of warming

Permafrost is experiencing climate warming at a rate that is two times faster than the rest of the Earth''s surface. However, it is still lack of a quantitative basis for predicting the functional stability of permafrost ecosystems in carbon (C) and nutrient cycling. We compiled the data of 708 observations from 89 air‐warming experiments in the Northern Hemisphere and characterized the general effects of temperature increase on permafrost C exchange and balance, biomass production, microbial biomass, soil nutrients, and vegetation N dynamics through a meta‐analysis. Also, an investigation was made on how responses might change with habitat‐specific (e.g., plant functional groups and soil moisture status) conditions and warming variables (e.g., warming phases, levels, and timing). The net ecosystem C exchange (NEE) was found to be downregulated by warming as a result of a stronger sensitivity to warming in respiration (15.6%) than in photosynthesis (6.2%). Vegetation usually responded to warming by investing more C to the belowground, as belowground biomass increased much more (30.1%) than aboveground biomass (2.9%). Warming had a minor effect on microbial biomass. Warming increased soil ammonium and nitrate concentrations. What''s more, a synthesis of 70 observations from 11 herbs and 9 shrubs revealed a 2.5% decline of N in green leaves. Compared with herbs, shrubs had a stronger response to respiration and had a decline in green leaf N to a greater extent. Not only in dry condition did green leaf N decline with warming but also in wet conditions. Warming in nongrowing seasons would negatively affect soil water, C uptake, and biomass production during growing seasons. Permafrost C loss and vegetation N decline may increase with warming levels and timing. Overall, these findings suggest that besides a positive C cycling–climate feedback, there will be a negative feedback between permafrost nutrient cycling and climate warming.     

短期氮素添加和模拟放牧对青藏高原高寒草甸生态系统呼吸的影响

氮沉降和放牧是影响草地碳循环过程的重要环境因子,但很少有研究探讨这些因子交互作用对生态系统呼吸的影响。在西藏高原高寒草甸地区开展了外源氮素添加与刈割模拟放牧实验,测定了其对植物生物量分配、土壤微生物碳氮和生态系统呼吸的影响。结果表明:氮素添加显著促进生态系统呼吸,而模拟放牧对其无显著影响,且降低了氮素添加的刺激作用。氮素添加通过提高微生物氮含量和土壤微生物代谢活性,促进植物地上生产,从而增加生态系统的碳排放;而模拟放牧降低了微生物碳含量,且降低了氮素添加的作用,促进根系的补偿性生长,降低了氮素添加对生态系统碳排放的刺激作用。这表明,放牧压力的存在会抑制氮沉降对高寒草甸生态系统碳排放的促进作用,同时外源氮输入也会缓解放牧压力对高寒草甸生态系统生产的负面影响。     

青藏高原多年冻土区不同水分条件的高寒草甸根系功能性状对增温的响应

在青藏高原多年冻土广泛分布的风火山地区,选择小嵩草（Kobresia pygmea）草甸和藏嵩草（Kobresia tibetica）沼泽化草甸为研究对象,采用开顶增温室（Open top chambers, OTCs）模拟气候变暖,探讨模拟增温对土壤水分差异的两种草甸地下生物量及根系功能性状的影响。结果显示,（1）增温显著增加小嵩草草甸0—20 cm根系生物量,主要是由于表层（0—10 cm）根系生物量显著增加,而对藏嵩草沼泽化草甸根系生物量无影响。（2）增温显著增加了小嵩草草甸根组织密度,同时提高了藏嵩草沼泽化草甸10—20 cm的比根长和比根面积（3）增温降低了小嵩草草甸的根系碳含量及10—20 cm根系氮含量,增加了藏嵩草沼泽化草甸的碳含量及10—20 cm根系氮含量,显著提高了小嵩草草甸和藏嵩草沼泽化草甸深层（10—20 cm）根系碳氮比。这些结果预示着增温使得土壤水分较低的小嵩草草甸朝着资源保守的慢速生长型发展,以适应暖干化的环境;土壤水分较高的藏嵩草沼泽化草甸朝着资源获取的快速生长型发展,加速利用土壤中的养分满足植物生长需要。可见,土壤水分可以调节高寒草甸对气候变暖的演变趋势,强调了水分的重要性。     

Nitrogen availability mediates soil carbon cycling response to climate warming: A meta-analysis

Global climate warming may induce a positive feedback through increasing soil carbon (C) release to the atmosphere. Although warming can affect both C input to and output from soil, direct and convincing evidence illustrating that warming induces a net change in soil C is still lacking. We synthesized the results from field warming experiments at 165 sites across the globe and found that climate warming had no significant effect on soil C stock. On average, warming significantly increased root biomass and soil respiration, but warming effects on root biomass and soil respiration strongly depended on soil nitrogen (N) availability. Under high N availability (soil C:N ratio < 15), warming had no significant effect on root biomass, but promoted the coupling between effect sizes of root biomass and soil C stock. Under relative N limitation (soil C:N ratio > 15), warming significantly enhanced root biomass. However, the enhancement of root biomass did not induce a corresponding C accumulation in soil, possibly because warming promoted microbial CO₂ release that offset the increased root C input. Also, reactive N input alleviated warming-induced C loss from soil, but elevated atmospheric CO₂ or precipitation increase/reduction did not. Together, our findings indicate that the relative availability of soil C to N (i.e., soil C:N ratio) critically mediates warming effects on soil C dynamics, suggesting that its incorporation into C-climate models may improve the prediction of soil C cycling under future global warming scenarios.     

基于OTCs模拟增温方式探讨气候变暖对青藏高原草地生态系统的影响

开顶式生长室(OTCs)增温实验是研究全球气候变化与陆地生态系统关系的主要方法之一,已广泛应用于青藏高原地区。该文通过对近些年国内外研究文献的回顾,分别从植物物候、群落结构、生物量和土壤方面综合分析青藏高原草地生态系统对OTCs模拟增温实验的响应。研究发现:增温使群落返青期提前、枯黄期延迟,生长季延长;有利于禾本科植物的生长;高寒草甸地下生物量分配格局向深层转移;高寒草地生态系统对模拟增温的响应存在不确定性,受到地域、群落类型和实验时间的影响;在增温条件下,降雨和冻土融化引起的土壤水分变化通过调控生态系统的物候、生产力、土壤等途径控制着生态系统对气候变暖的响应。并在此基础上,提出了将来应着重研究的几个方面。     

短期增温对西藏念青唐古拉山典型植物群落结构和稳定性的影响

青藏高原气候变暖幅度显著高于全球其他区域,深刻影响着该地区植物群落的结构和稳定性。选择西藏念青唐古拉山的三种典型植物群落(高寒草原、高寒草甸和流石滩)作为研究对象,采用开顶式增温箱(OTC)模拟增温,研究了短期增温对植物群落结构和稳定性的影响。结果表明:(1)增温改变了群落的优势物种,影响其结构组成,而对物种多样性无显著影响;(2)增温显著降低了高寒草甸的地上生物量(P < 0.05),增加地下生物量(P < 0.01),从而导致了群落地下地上生物量分配策略的改变;(3)增温降低群落中部分物种的生态位宽度,进而影响群落稳定性,其中高寒草甸变化最大,达到-66.8%。研究结果可为青藏高原高寒草地生态系统应对和适应未来气候变化提供一定科学依据。     

Grazing intensity significantly affects belowground carbon and nitrogen cycling in grassland ecosystems: a meta‐analysis

Livestock grazing activities potentially alter ecosystem carbon (C) and nitrogen (N) cycles in grassland ecosystems. Despite the fact that numerous individual studies and a few meta‐analyses had been conducted, how grazing, especially its intensity, affects belowground C and N cycling in grasslands remains unclear. In this study, we performed a comprehensive meta‐analysis of 115 published studies to examine the responses of 19 variables associated with belowground C and N cycling to livestock grazing in global grasslands. Our results showed that, on average, grazing significantly decreased belowground C and N pools in grassland ecosystems, with the largest decreases in microbial biomass C and N (21.62% and 24.40%, respectively). In contrast, belowground fluxes, including soil respiration, soil net N mineralization and soil N nitrification increased by 4.25%, 34.67% and 25.87%, respectively, in grazed grasslands compared to ungrazed ones. More importantly, grazing intensity significantly affected the magnitude (even direction) of changes in the majority of the assessed belowground C and N pools and fluxes, and C : N ratio as well as soil moisture. Specifically,light grazing contributed to soil C and N sequestration whereas moderate and heavy grazing significantly increased C and N losses. In addition, soil depth, livestock type and climatic conditions influenced the responses of selected variables to livestock grazing to some degree. Our findings highlight the importance of the effects of grazing intensity on belowground C and N cycling, which may need to be incorporated into regional and global models for predicting effects of human disturbance on global grasslands and assessing the climate‐biosphere feedbacks.     

Plant community structure regulates responses of prairie soil respiration to decadal experimental warming

Soil respiration is recognized to be influenced by temperature, moisture, and ecosystem production. However, little is known about how plant community structure regulates responses of soil respiration to climate change. Here, we used a 13‐year field warming experiment to explore the mechanisms underlying plant community regulation on feedbacks of soil respiration to climate change in a tallgrass prairie in Oklahoma, USA. Infrared heaters were used to elevate temperature about 2 °C since November 1999. Annual clipping was used to mimic hay harvest. Our results showed that experimental warming significantly increased soil respiration approximately from 10% in the first 7 years (2000–2006) to 30% in the next 6 years (2007–2012). The two‐stage warming stimulation of soil respiration was closely related to warming‐induced increases in ecosystem production over the years. Moreover, we found that across the 13 years, warming‐induced increases in soil respiration were positively affected by the proportion of aboveground net primary production (ANPP) contributed by C₃ forbs. Functional composition of the plant community regulated warming‐induced increases in soil respiration through the quantity and quality of organic matter inputs to soil and the amount of photosynthetic carbon (C) allocated belowground. Clipping, the interaction of clipping with warming, and warming‐induced changes in soil temperature and moisture all had little effect on soil respiration over the years (all P > 0.05). Our results suggest that climate warming may drive an increase in soil respiration through altering composition of plant communities in grassland ecosystems.     

青藏高原高寒草甸植物群落生物量对氮、磷添加的响应

青藏高原正经历着明显的温暖化过程, 由此引起的土壤温度的升高促进了土壤中微生物的活性, 同时青藏高原东缘地区大气氮沉降十分明显, 并呈逐年增加的趋势, 这些环境变化均促使土壤中可利用营养元素增加, 因此深入了解青藏高原高寒草甸植物生物量对可利用营养元素增加的响应, 是准确预测未来全球变化背景下青藏高原高寒草甸碳循环过程的重要基础。该研究基于在青藏高原高寒草甸连续4年(2009-2012年)氮、磷添加后对不同功能群植物地上生物量、群落地上和地下生物量的测定, 探讨高寒草甸生态系统碳输入对氮、磷添加的响应。结果表明: (1)氮、磷添加均极显著增加了禾草的地上绝对生物量及其在群落总生物量中所占的比例, 同时均显著降低了杂类草在群落总生物量中的比例, 此外磷添加极显著降低了莎草地上绝对生物量及其在群落总生物量中所占的比例。(2)氮、磷添加均显著促进了青藏高原高寒草甸的地上生物量增加, 分别增加了24%和52%。(3)氮添加对高寒草甸地下生物量无显著影响, 而磷添加后地下生物量有增加的趋势。(4)氮添加对高寒草甸植物总生物量无显著影响, 而磷添加后植物总生物量显著增加。研究表明, 氮、磷添加可缓解青藏高原高寒草甸植物生长的营养限制, 促进植物地上部分的生长, 然而高寒草甸植物的生长极有可能更受土壤中可利用磷含量的限制。     

Effects of field experimental warming on wheat root distribution under conventional tillage and no‐tillage systems

Despite the obvious importance of roots to agro‐ecosystem functioning, few studies have attempted to examine the effects of warming on root biomass and distribution, especially under different tillage systems. In this study, we performed a field warming experiment using infrared heaters on winter wheat, in long‐term conventional tillage and no‐tillage plots, to determine the responses of root biomass and distribution to warming. Soil monoliths were collected from three soil depths (0–10, 10–20, and 20–30 cm). Results showed that root biomass was noticeably increased under both till and no‐till tillage systems (12.1% and 12.9% in 2011, and 9.9% and 14.5% in 2013, in the two tillage systems, respectively) in the 0–30 cm depth, associated with a similar increase in shoot biomass. However, warming‐induced root biomass increases occurred in the deeper soil layers (i.e., 10–20 and 20–30 cm) in till, while the increase in no‐till was focused in the surface layer (0–10 cm). Differences in the warming‐induced increases in root biomass between till and no‐till were positively correlated with the differences in soil total nitrogen (R² = .863, p < .001) and soil bulk density (R² = .853, p < .001). Knowledge of the distribution of wheat root in response to warming should help manage nutrient application and cycling of soil C‐N pools under anticipated climate change conditions.     

Plant diversity and dry-matter dynamics of peri-urban plant communities in an Indian dry tropical region

The emergence of variety of land-use changes due to continuous anthropogenic pressure in peri-urban areas may concomitantly result in modification of the structure of associated plant communities for their sustainable growth. In the present study, plant diversity, and above- and belowground biomass distribution among species were investigated to understand their dynamics across different season, soil, and site conditions in a dry tropical peri-urban region in India. From four study sites that covered contrasting land uses: abandoned brick kiln (ABK), grazing land (GL), Kali river bank (KRB), and agricultural land (AL), a total of 360 monoliths were randomly extracted in three seasons, and dry weights were estimated for aboveground and belowground parts of species individuals. Seasonal soil samples of the sites were analyzed for physico-chemical characteristics. Of the total 87 recorded species that were mainly annual weeds and ruderals, 77% were forbs and 23% grasses. The ranges of plant biomass recorded across all sites and seasons were: aboveground 228–738 g m⁻², belowground 83–288 g m⁻², and a total of 344–1,026 g m⁻². The dominance of species differed between above- and belowground; some species dominated only above- or belowground, and others dominated in both layers. Above- and belowground biomass of the sites, differential community-biomass allocation to above- and belowground parts and species dominants varied significantly with site and season. ABK and AL sites showed lower species diversity and soil nutrients compared to GL and KRB sites. Belowground biomass significantly declined with increasing soil organic C and total N, indicating altered dry matter allocation under resource-scarce habitat conditions. Higher diversity occurred at both low- and high-biomass sites, reflective of enhanced ability of these plant communities to exploit resources maximally in spatio-temporal pattern.     

青藏高原高寒草甸初级生产力及其主要影响因素

青藏高原有各类天然草地14×108hm2,其中高寒草甸和高寒灌丛约占青藏高原天然草地面积的50%,占全国草地总面积的16.2%。嵩草草甸是高寒草甸的主体,包括矮嵩草草甸、金露梅灌丛草甸、藏嵩草草甸、小嵩草草甸和高山嵩草草甸等,这5类高寒草甸平均地上生物量分别为354.2、422.4、445.1、227.3和368.5g/m2,地下生物量分别为3389.6、3548.3、11922.7、4439.3、5604.8g/m2,地下与地上生物量的比例分别为10.55、10.15、27.82、14.82和15.21,远大于IPCC(2006)报告中地下/地上生物量比例的默认值(2.8±95%)。地下生物量对气候变化和放牧的反应比地上生物量更敏感,干旱和重度放牧均降低了地下/地上生物量的比例。在极度退化状态下地下/地上生物量的比例2。对于轻度和中度退化的高寒草甸应以围封禁牧为主要恢复措施,但如果结合补播和施肥,则恢复速率会加快;对于重度和极度退化的高寒草甸,由于草地植物群落中优良牧草的比例极低,仅靠自然恢复很难进行恢复或需要的年限很长,所以必须采用人工重建的措施,并结合毒杂草防除和施肥等措施进行恢复,通过建立人工或半人工草地的措施予以重建。     

Plant biomass and species composition along an environmental gradient in montane riparian meadows

In riparian meadows, narrow zonation of the dominant vegetation frequently occurs along the elevational gradient from the stream edge to the floodplain terrace. We measured plant species composition and above- and belowground biomass in three riparian plant communities—a priori defined as wet, moist, and dry meadow—along short streamside topographic gradients in two montane meadows in northeast Oregon. The objectives were to: (1) compare above- and belowground biomass in the three meadow communities; (2) examine relations among plant species richness, biomass distribution, water table depth, and soil redox potential along the streamside elevational gradients. We installed wells and platinum electrodes along transects (perpendicular to the stream; n=5 per site) through the three plant communities, and monitored water table depth and soil redox potential (10 and 25 cm depth) from July 1997 to August 1999. Mean water table depth and soil redox potential differed significantly along the transects, and characterized a strong environmental gradient. Community differences in plant species composition were reflected in biomass distribution. Highest total biomass (live+dead) occurred in the sedge-dominated wet meadows (4,311±289 g/m²), intermediate biomass (2,236±221 g/m²) was seen in the moist meadow communities, dominated by grasses and sedges, and lowest biomass (1,403±113 g/m²) was observed in the more diverse dry meadows, dominated by grasses and forbs. In the wet and moist communities, belowground biomass (live+dead) comprised 68–81% of the totals. Rhizome-to-root ratios and distinctive vertical profiles of belowground biomass reflected characteristics of the dominant graminoid species within each community. Total biomass was positively correlated with mean water table depth, and negatively correlated with mean redox potential (10 cm and 25 cm depths; P <0.01) and species richness (P <0.05), indicating that the distribution of biomass coincided with the streamside edaphic gradient in these riparian meadows.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Nitrogen addition promotes terrestrial plants to allocate more biomass to aboveground organs: A global meta-analysis

A significant increase in reactive nitrogen (N) added to terrestrial ecosystems through agricultural fertilization or atmospheric deposition is considered to be one of the most widespread drivers of global change. Modifying biomass allocation is one primary strategy for maximizing plant growth rate, survival, and adaptability to various biotic and abiotic stresses. However, there is much uncertainty as to whether and how plant biomass allocation strategies change in response to increased N inputs in terrestrial ecosystems. Here, we synthesized 3516 paired observations of plant biomass and their components related to N additions across terrestrial ecosystems worldwide. Our meta-analysis reveals that N addition (ranging from 1.08 to 113.81 g m⁻² year⁻¹) increased terrestrial plant biomass by 55.6% on average. N addition has increased plant stem mass fraction, shoot mass fraction, and leaf mass fraction by 13.8%, 12.9%, and 13.4%, respectively, but with an associated decrease in plant reproductive mass (including flower and fruit biomass) fraction by 3.4%. We further documented a reduction in plant root-shoot ratio and root mass fraction by 27% (21.8%–32.1%) and 14.7% (11.6%–17.8%), respectively, in response to N addition. Meta-regression results showed that N addition effects on plant biomass were positively correlated with mean annual temperature, soil available phosphorus, soil total potassium, specific leaf area, and leaf area per plant. Nevertheless, they were negatively correlated with soil total N, leaf carbon/N ratio, leaf carbon and N content per leaf area, as well as the amount and duration of N addition. In summary, our meta-analysis suggests that N addition may alter terrestrial plant biomass allocation strategies, leading to more biomass being allocated to aboveground organs than belowground organs and growth versus reproductive trade-offs. At the global scale, leaf functional traits may dictate how plant species change their biomass allocation pattern in response to N addition.     

Nitrogen availability increases in a tundra ecosystem during five years of experimental permafrost thaw

Perennially frozen soil in high latitude ecosystems (permafrost) currently stores 1330–1580 Pg of carbon (C). As these ecosystems warm, the thaw and decomposition of permafrost is expected to release large amounts of C to the atmosphere. Fortunately, losses from the permafrost C pool will be partially offset by increased plant productivity. The degree to which plants are able to sequester C, however, will be determined by changing nitrogen (N) availability in these thawing soil profiles. N availability currently limits plant productivity in tundra ecosystems but plant access to N is expected improve as decomposition increases in speed and extends to deeper soil horizons. To evaluate the relationship between permafrost thaw and N availability, we monitored N cycling during 5 years of experimentally induced permafrost thaw at the Carbon in Permafrost Experimental Heating Research (CiPEHR) project. Inorganic N availability increased significantly in response to deeper thaw and greater soil moisture induced by Soil warming. This treatment also prompted a 23% increase in aboveground biomass and a 49% increase in foliar N pools. The sedge Eriophorum vaginatum responded most strongly to warming: this species explained 91% of the change in aboveground biomass during the 5 year period. Air warming had little impact when applied alone, but when applied in combination with Soil warming, growing season soil inorganic N availability was significantly reduced. These results demonstrate that there is a strong positive relationship between the depth of permafrost thaw and N availability in tundra ecosystems but that this relationship can be diminished by interactions between increased thaw, warmer air temperatures, and higher levels of soil moisture. Within 5 years of permafrost thaw, plants actively incorporate newly available N into biomass but C storage in live vascular plant biomass is unlikely to be greater than losses from deep soil C pools.     

Individual and interactive effects of warming and nitrogen supply on CO2 fluxes and carbon allocation in subarctic grassland

Climate warming has been suggested to impact high latitude grasslands severely, potentially causing considerable carbon (C) losses from soil. Warming can also stimulate nitrogen (N) turnover, but it is largely unclear whether and how altered N availability impacts belowground C dynamics. Even less is known about the individual and interactive effects of warming and N availability on the fate of recently photosynthesized C in soil. On a 10-year geothermal warming gradient in Iceland, we studied the effects of soil warming and N addition on CO₂ fluxes and the fate of recently photosynthesized C through CO₂ flux measurements and a ¹³CO₂ pulse-labeling experiment. Under warming, ecosystem respiration exceeded maximum gross primary productivity, causing increased net CO₂ emissions. N addition treatments revealed that, surprisingly, the plants in the warmed soil were N limited, which constrained primary productivity and decreased recently assimilated C in shoots and roots. In soil, microbes were increasingly C limited under warming and increased microbial uptake of recent C. Soil respiration was increased by warming and was fueled by increased belowground inputs and turnover of recently photosynthesized C. Our findings suggest that a decade of warming seemed to have induced a N limitation in plants and a C limitation by soil microbes. This caused a decrease in net ecosystem CO₂ uptake and accelerated the respiratory release of photosynthesized C, which decreased the C sequestration potential of the grassland. Our study highlights the importance of belowground C allocation and C-N interactions in the C dynamics of subarctic ecosystems in a warmer world.