Species‐specific activation time‐lags can explain habitat restrictions in hydrophilic lichens

Photosystem II (PSII) activation after hydration with water or humid air was measured in four hydrophilic and a generalist lichen to test the hypothesis that slow activation might explain habitat restriction in the former group. For the hydrophilic species, activation was after 4 h nearly completed in Lobaria amplissima and Platismatia norvegica, while only c. 50% for Bryoria bicolor and Usnea longissima. The generalist Platismatia glauca was activated instantaneously. The effect of this on lichen field performance was investigated using a dynamic model separating the two water sources rain and humid air. Model simulations were made using the species‐specific characteristics and climate data from 12 stream microhabitats. For U. longissima, slow PSII activation could reduce realized photosynthesis by a factor of five. Bryoria bicolor was almost as severely affected, while P. norvegica displayed moderate reductions. Lobaria amplissima displayed longer realized activity periods even in unfavourable microclimates, possibly because of a higher water loss resistance. Both close proximity to streams and presence of turbulent water had a positive impact on realized activity among the slowly activated species, coinciding with observed distribution patterns of hydrophilic species. The results presented here may thus partly explain observed habitat restrictions of rare hydrophilic lichens.     

土壤微生物数量对模拟氮沉降增加的早期响应

通过以南亚热带森林三种主要树种即荷木、锥栗和黄果厚壳桂为主的苗圃试验地的苗圃控制实验,初步探讨土壤微生物数量对模拟氮沉降增加的响应。结果表明,施氮增加对土壤微生物数量的影响根据类群和氮处理水平不同而异。总的来说,施氮增加对土壤微生物数量具有促进作用,这种促进作用对放线菌数量仅在一定(中氮)处理水平以下,超过此水平施氮增加则表现为抑制作用,而施氮对真菌数量则始终表现为抑制作用,尤其以中N处理水平的抑制作用最强。但目前细菌仍占微生物总量的绝对优势,放线菌次之,真菌则占微生物总量的比例最小。分析结果还表明,有效氮与放线菌、有效氮与细菌数量变化呈显著相关关系。     

Nitrogen deposition drives lichen community changes through differential species responses

Nitrogen (N) deposition has increased globally over the last 150 years and further increases are predicted. Epiphytic lichens decline in abundance and diversity in areas with high N loads, and the abundance of lichens decreases along gradients of increased deposition. Thus, although N is an essential nutrient for lichens, excessive loads may be detrimental for them. However, these gradients include many correlated pollutants and the mechanisms behind the decline are thus poorly known. The aim of this study was to assess effects of N deposition, alone, on the epiphytic lichen community composition in a naturally N‐poor boreal forest. For this purpose, whole spruce trees were fertilized daily with N at five levels, equivalent to 0.6, 6, 12.5, 25, and 50 kg N ha^?1 yr^?1, during four consecutive growing seasons (2006–2009), and changes in the abundance of lichens were monitored each autumn from the preceding year (2005). The studied lichen communities were highly dynamic and responded strongly to the environmental perturbation. N deposition detectably altered the direction of succession and reduced the species richness of the epiphytic lichen communities, even at the lowest fertilization application (6 kg N ha^?1 yr^?1). The simulated N deposition caused significant changes in the abundance of Alectoria sarmentosa, Bryoria spp., and Hypogymnia physodes, which all increased at low N loads and decreased at high loads, but with species‐specific optima. The rapid decline of A. sarmentosa may have been caused by the added nitrogen reducing the stability of the lichen thalli, possibly due to increases in the photobiont: mycobiont ratio or parasitic fungal attacks. We conclude that increases in nitrogen availability, per se, could be responsible for the reductions in lichen abundance and diversity observed along deposition gradients, and those community responses may be due to physiological responses of the individual species rather than changes in competitive interactions.     

Modelling hydration and photosystem II activation in relation to in situ rain and humidity patterns: a tool to compare performance of rare and generalist epiphytic lichens

A dynamic water and activity model was developed to assess how efficiently lichens can exploit in situ rain and humid air. The capacity to rehydrate and activate photosynthesis [i.e. photosystem II (PSII)] by these water sources was compared among four hydrophilic and one generalist epiphytic lichen. Hydration status, potential (instant activation) and realized (delayed activation) day‐light activity were simulated using a model based on species‐specific hydration, PSII activation characteristics and in situ water content for Platismatia norvegica in three microclimatic scenarios. The results showed that delayed PSII activation could have profound effects on lichens' ability to exploit environmental water sources. During rain, realized activity was reduced by 19, 34 and 56% compared to simulations assuming instant activation for three hydrophilic lichens in the driest microclimate. During humid air, the reduction was 81% for the most extreme species and scenario, because of slow hydration and low equilibrium water content. Many and brief hydration events may thus hamper species with slow activation and fast desiccation kinetics. No evidence of compensation by a ‘water‐holding’ morphology was observed among studied species. The developed model may provide a tool for identifying suitable habitats for long‐term persistence of lichens with physiological constraints.     

Impacts of short-term nitrogen addition on the thallus nitrogen and phosphorus balance of the dominant epiphytic lichens in the Shennongjia mountains,China

Aims To better understand whether and how nitrogen addition impacts the epiphytic lichens in the Shennongjia Nature Reserve (China).     

川南天然常绿阔叶林土壤酶活性特征及其对模拟N沉降的响应

通过原位进行低氮(LN,50 kg N·hm-2·a-1)、中氮(MN,100 kg N·hm-2·a-1)和高氮(HN,150 kg N·hm-2·a-1)处理,研究了川南天然常绿阔叶林土壤酶活性特征及其对模拟N沉降的响应.结果表明,森林土壤过氧化氢酶、脲酶、纤维素酶和蔗糖酶的活性在垂直分布上均表现为0～10 cm土层高于10～20 cm土层;各种土壤酶活性的季节变化明显,但土壤酶活性还因取样时间和土层不同而异.其中,0～10 cm和10～20 cm土层中过氧化氢酶活性高峰均出现在秋季(92 d);0～10 cm土层中脲酶活性高峰出现在秋季,而10～20 cm土层中脲酶活性高峰出现在冬季(183 d);0～10 cm和10～20 cm土层中土壤纤维素酶活性高峰则出现在翌年春季(274 d);0～10 cm土层中蔗糖酶活性有两个高峰,分别出现在秋季和翌年春季;而10～20 cm土层中蔗糖酶活性只有一个高峰,出现在翌年春季.总体上,常绿阔叶林中不同土壤酶活性对N沉降的响应有所差异.其中,N沉降降低了常绿阔叶林土壤纤维素酶和过氧化氢酶活性,增加了土壤脲酶和蔗糖酶活性.不同浓度N处理间酶活性表现出了不同程度的差异性,但其变化的规律还不明显.     

Anthropogenic nitrogen deposition enhances carbon sequestration in boreal soils

It is proposed that carbon (C) sequestration in response to reactive nitrogen (N_r) deposition in boreal forests accounts for a large portion of the terrestrial sink for anthropogenic CO₂ emissions. While studies have helped clarify the magnitude by which N_r deposition enhances C sequestration by forest vegetation, there remains a paucity of long‐term experimental studies evaluating how soil C pools respond. We conducted a long‐term experiment, maintained since 1996, consisting of three N addition levels (0, 12.5, and 50 kg N ha^?1 yr^?1) in the boreal zone of northern Sweden to understand how atmospheric N_r deposition affects soil C accumulation, soil microbial communities, and soil respiration. We hypothesized that soil C sequestration will increase, and soil microbial biomass and soil respiration will decrease, with disproportionately large changes expected compared to low levels of N addition. Our data showed that the low N addition treatment caused a non‐significant increase in the organic horizon C pool of ~15% and a significant increase of ~30% in response to the high N treatment relative to the control. The relationship between C sequestration and N addition in the organic horizon was linear, with a slope of 10 kg C kg^?1 N. We also found a concomitant decrease in total microbial and fungal biomasses and a ~11% reduction in soil respiration in response to the high N treatment. Our data complement previous data from the same study system describing aboveground C sequestration, indicating a total ecosystem sequestration rate of 26 kg C kg^?1 N. These estimates are far lower than suggested by some previous modeling studies, and thus will help improve and validate current modeling efforts aimed at separating the effect of multiple global change factors on the C balance of the boreal region.     

低温胁迫后苔藓植物对模拟氮沉降条件的生理响应

2008年初受强寒潮的影响, 中国华南大部分地区出现持续的异常低温。该文研究了华南地区常见的3种苔藓植物 ——刺边小金发藓拟刺亚种(Pogonatum cirratum subsp. fuscatum)、大灰藓(Hypnum plumaeforme)和石地钱(Reboulia hemisphaerica), 在人工模拟氮(N)沉降两年并经历2008年初异常低温气候后的生理响应变化, 并与2007年正常气候情况下人工加N一年后的结果进行比较, 分析苔藓植物的生长与N沉降之间的关系, 并探讨N沉降对低温胁迫后苔藓植物的补偿生长的影响。结果显示: 2008年3种苔藓植物的净光合速率和淀粉含量在加N量为0-60 kg N·hm ^-2·a^-1的范围内均随着N浓度的上升而下降, 总N含量在加N量处于0-40 kg N·hm^-2·a^-1的范围内随着N浓度的上升而上升, 至60 kg N·hm^-2·a^-1时不再上升, 甚至有所下降。2008年, 3种苔藓植物大多数碳氮代谢指标在对照及低N条件下与2007年加N 1年且在正常气候时同种N处理时相比均有不同程度的上升, 但上升幅度与加N浓度成反比, 至中高N条件时两者常较接近, 显示苔藓植物在经历低温胁迫后会出现超补偿效应, 但是在N沉降升高的条件下, 补偿生长能力下降。     

几种海南岛热带雨林优势种植物挥发性有机物排放对模拟氮沉降的短期响应

BVOCs(Biogenic Volatile Organic Compounds)是植物向大气释放的一类重要气态化合物,能参与大气化学过程和陆地生态系统碳素循环。分析环境因子对BVOCs排放的影响,对科学认识未来气候变化具有重要意义。氮素作为植物生长、发育所需的大量营养元素之一,其沉降增加是当前全球气候变化的主要驱动因素之一,但学者对BVOCs如何应对氮沉降增加知之甚少。因此以海南岛热带雨林树种：木荷(Schima superba)、厚壳桂(Cryptocarya chinensis)和线枝蒲桃(Syzygium araiocladum)为研究对象,通过温室盆栽实验模拟氮沉降对3个树种BVOCs释放的短期效应。主要结论如下：(1)自然状态下,从木荷、厚壳桂和线枝蒲桃的枝叶中鉴定出14、34和24种挥发性有机化合物,包括异戊二烯、单萜烯、倍半萜烯和其他挥发性有机化合物(烷烃、羰基、醛、醇、酯、醚和酸),此外三个阔叶树种释放BVOCs的速率呈厚壳桂>木荷>线枝蒲桃;(2)外源施氮均促进了三种植物幼苗VOCs释放,其中总VOCs释放速率和成分数量均随施氮浓度的升高而增加,且叶面...     

模拟氮沉降和降雨对华西雨屏区常绿阔叶林凋落物分解的影响

从2013年11月至2015年5月,采用凋落物分解袋法,设置了对照(CK)、氮沉降(N)、减雨(R)、增雨(A)、氮沉降+减雨(NR)、氮沉降+增雨(NA)6个处理水平,研究了模拟氮沉降和降雨对华西雨屏区常绿阔叶林凋落物分解的影响。结果表明:华西雨屏区常绿阔叶林凋落叶分解较快,凋落枝分解较慢;凋落物夏季分解较快,其他季节分解较慢。经过18个月的分解后,凋落叶和枝的质量残留率分别为45.86%和86.67%,凋落叶分解50%需要的时间为1.42 a,比枝短6.19 a。各处理凋落物叶分解系数表现为:k(A)k(CK)k(NA)k(N)k(R)k(NR),凋落枝质量残留率表现为:NNRRNACKA。模拟氮沉降、减雨和增雨处理凋落叶分解50%分别需要1.79、1.94a和1.36a,凋落枝分解50%分别需要8.84、8.63 a和6.47 a。各处理凋落叶分解95%需要5.37—11.33 a,凋落枝分解95%需要27.41—33.84 a。同一氮沉降条件下,增雨处理促进凋落叶分解,减雨处理抑制凋落叶分解;同一降雨条件下,氮沉降抑制凋落叶分解。氮沉降或降雨对凋落物的分解产生显著影响(P0.05),其交互作用影响不显著(P0.05)。可见,在氮沉降持续增加和降雨格局改变的背景下,增雨促进了华西雨屏区天然常绿阔叶林凋落物的分解,氮沉降和减雨抑制了凋落物的分解,模拟氮沉降和降雨对凋落物的分解交互作用表现不明显。     

模拟氮沉降对温带阔叶红松林地土壤氮素净矿化的影响

以长白山阔叶红松混交林为研究对象,于2006—2008年原位模拟不同形态氮((NH4)2SO4、NH4Cl和KNO3)沉降水平(22.5和45kgN·hm-2·a-1),利用树脂芯法技术(resin-core incubation technique)测定了表层(有机层0～7cm)和土层(0～15cm)土壤氮素净矿化、净氨化和净硝化通量的季节和年际变化规律。同时,结合前人报道的有关林地碳、氮过程及其环境变化影响的结果,力求有效预估森林生态系统中氮素年矿化通量对大气氮沉降量和水热条件等因子变化的响应。结果表明,长白山阔叶红松林地土壤氮素年净矿化通量为1.2～19.8kgN·hm-2·a-1,2008年不同深度的土壤氮素年净矿化通量均显著高于2006和2007年(P<0.05)。随着模拟氮沉降量增加,土壤氮素净矿化通量也随之增加,尤其外源NH4+-N输入对净矿化通量的促进作用更为明显(P<0.05),但随着施肥年限的延长,这种促进作用逐渐减弱。与林地0～15cm土壤相比,氮沉降增加对0～7cm有机层氮素净氨化和净矿化通量的促进作用更为明显,尤其NH4Cl处理的促进作用更大。结合前人报道的野外原位观测结果,土壤氮素年净矿化通量随氮素沉降量的增加而增大,氮沉降量对不同区域森林土壤氮素净矿化通量的贡献率约为52%;氮沉降量(x1)和pH值(x2)可以解释区域森林土壤氮素年净矿化通量(y)变化的70%(y=0.54x1-18.38x2-109.55,R2=0.70,P<0.0001)。前人研究结果仅提供区域年均温度,未考虑积温的影响,这可能是造成年净矿化通量与温度无关的原因。今后的研究工作应该加强区域森林土壤积温观测,进而更加准确地预估森林土壤氮素的年净矿化通量。     

樟树人工林凋落物养分含量及归还量对氮沉降的响应

氮沉降的持续增加对陆地生态系统的健康发展构成严重威胁,森林是陆地生态系统中重要的组成部分,大量的氮沉降对其结构和功能造成严重影响。凋落物是森林生态系统养分循环的重要组成部分,它对土壤肥力、森林生态系统养分循环等方面具有重要作用。为了探讨亚热带常绿阔叶森林凋落物对氮沉降增加的响应,在湖南省森林植物园以樟树人工林为研究对象进行模拟氮沉降的实验,实验设置4种氮添加水平CK(0g N m~(-2)a~(-1),对照)、LN(5g N m~(-2)a~(-1)),MN(15g N m~(-2)a~(-1)),HN(30g N m~(-2)a~(-1)),研究氮沉降对樟树林年凋落物量、凋落物养分含量以及归还量的影响。结果表明:不同施氮水平下(CK、LN、MN、HN),樟树林凋落物的年凋落量分别为(4.53±0.32)t hm~(-2)a~(-1)、(3.95±0.28)t hm~(-2)a~(-1)、(3.56±0.41)t hm~(-2)a~(-1)、(4.46±0.48)t hm~(-2)a~(-1),施氮抑制了樟树林的凋落量,且低、中氮处理下差异显著(P0.05);施氮处理后凋落物的养分含量大小顺序为:CNCaKMg,凋落物的碳含量没有显著变化,但氮含量都有所增加,因此,施氮降低了樟树凋落物各组分的C/N比;凋落物中元素的年归还量大小顺序表现为:CNCaKMg,施氮处理对凋落物C、K、Ca、Mg归还量有抑制作用,但对凋落物N归还量表现为促进作用。     

Nonlinear responses of ecosystem carbon fluxes to nitrogen deposition in an old-growth boreal forest

Nitrogen (N) deposition is known to increase carbon (C) sequestration in N-limited boreal forests. However, the long-term effects of N deposition on ecosystem carbon fluxes have been rarely investigated in old-growth boreal forests. Here we show that decade-long experimental N additions significantly stimulated net primary production (NPP) but the effect decreased with increasing N loads. The effect on soil heterotrophic respiration (Rh) shifted from a stimulation at low-level N additions to an inhibition at higher levels of N additions. Consequently, low-level N additions resulted in a neutral effect on net ecosystem productivity (NEP), due to a comparable stimulating effect on NPP and Rh, while NEP was increased by high-level N additions. Moreover, we found nonlinear temporal responses of NPP, Rh and NEP to low-level N additions. Our findings imply that actual N deposition in boreal forests likely exerts a minor contribution to their soil C storage.     

降水和氮沉降增加对草地土壤酶活性的影响

为探究降水和氮沉降增加对草地生态系统土壤酶活性的影响,于2014年生长季在内蒙古温带典型羊草草原开展了野外原位控制实验。试验共设置降水(对照,W0,自然降水;W15,增加15%的年均降水量)、施氮(对照,CK,0 kg N hm~(-2)a~(-1);低氮,LN,25 kg N hm~(-2)a~(-1);中氮,MN,50 kg N hm~(-2)a~(-1);高氮,HN,100 kg N hm~(-2)a~(-1))及其交互作用等8个不同的处理水平来模拟降水和氮沉降增加的全球变化情景,分别定量探讨了不同水、氮添加条件下草地表层土壤中与氮循环相关的蛋白酶,脲酶,硝酸还原酶,亚硝酸还原酶活性的月动态变化及其与土壤理化性质之间的相关性。研究结果表明:在自然降水条件下,不同施氮水平蛋白酶、脲酶和硝酸还原酶活性无显著差异,亚硝酸还原酶活性相比于对照显著降低;在增加降水条件下,不同施氮水平对蛋白酶和硝酸还原酶活性未产生显著性影响,高氮水平显著降低脲酶和亚硝酸还原酶活性。不同施氮水平是否添加降水对亚硝酸还原酶活性无影响,而增添降水使低氮处理的蛋白酶活性和中、高氮处理水平的硝酸还原酶活性增加、高氮处理的脲酶活性降低。降水在影响蛋白酶和硝酸还原酶活性方面具有主效应,氮沉降在影响亚硝酸还原酶活性方面具有主效应,而降水和施氮处理未表现出明显地交互作用。土壤亚硝酸还原酶活性与土壤碳氮比和NH~+_4-N含量极显著正相关,与NO-3-N含量显著正相关。     

华北平原大气氮素沉降的时空变异

利用量雨器和湿沉降自动收集仪在华北平原9个监测点通过2a的试验,研究了农田生态系统中大气氮素沉降的时空变异。结果表明:华北平原大气氮素混合沉降的平均值为28.0 kg/(hm2.a),降水中铵态氮和硝态氮量平均分别为3.76 mg/L和1.85 mg/L。不同地区比较,北京大气氮素沉降为32.5 kg/(hm2.a),明显高于山东和河北两省的23.6 kg/(hm2.a)。北京各监测点的大气氮素沉降也存在明显空间变异,东北旺、房山的氮素沉降水平较高,延庆、顺义的氮素沉降水平较低。大气氮素沉降的年内分布不均,60%的沉降集中在降水较丰沛的6~9月份。氮素的输入与降雨量呈乘幂型正相关(r2=0.67),在农田生态系统中以铵态氮的沉降为主,铵态氮的沉降量是硝态氮的2.0倍;城市生态系统中以硝态氮的沉降为主,铵态氮的沉降量是硝态氮的0.79倍。在东北旺试验点近两年的监测结果表明,在等量降雨量条件下湿沉降输入的氮素(18~20.6 kg/hm2)明显低于混合沉降(26.3 kg/hm2),湿沉降的氮素输入仅占后者的73%,而混合沉降中的超量部分主要来自铵态氮,表明干沉降尤其是降尘带入的铵态氮也是华北平原大气氮素沉降的重要来源。     

Growth and nitrogen relations in the mat-forming lichens Stereocaulon paschale and Cladonia stellaris

BACKGROUND AND AIMS: Mat-forming lichens in the genera Stereocaulon and Cladonia have ecosystem-level effects in northern boreal forests. Yet the factors affecting the productivity of mat-forming lichens are not known. The aim of the presented work was to investigate whether mat-forming lichens adapted to low N availability employ N-conserving mechanisms similar to those of vascular plants in nutrient-poor ecosystems. Specifically, the following questions were asked: (a) Do lichens translocate N from basal areas to apical growth areas? (b) Are the quantities of N translocated of ecological significance. (c) Is lichen growth dependent on tissue N concentration [N]. METHODS: Two different, but complementary, field experiments were conducted using the mat-forming N2-fixing Stereocaulon paschale and non-fixing Cladonia stellaris as model species. First, N translocation was investigated by feeding lichens with Na(15)NO3 either directly to the apex (theoretical sink) or to the basal part (theoretical source) and observing the redistribution of (15)N after a growth period. Secondly, growth and variation in [N] in thalli of different lengths was measured after a growth period. KEY RESULTS: (15)N fed to lower parts of lichen was translocated towards the growing top, but not vice versa, indicating physiologically dependent translocation that follows a sink-source relationship. In the growth experiment where thalli were cut to different lengths, the significant decrease in [N] in apices of short vs. longer thalli after a growth period is consistent with internal relocation as an ecologically important source of N. CONCLUSIONS: The presented results demonstrate that internal recycling of N occurs in both species investigated and may be ecologically important in these mat-forming lichens under field conditions. The higher nitrogen use efficiency and relative growth rate in C. stellaris in comparison with S. paschale probably enable C. stellaris to dominate the ground cover vegetation in dry boreal coniferous forests under undisturbed conditions.     

模拟氮沉降及经营方式对毛竹林土壤酶活性的影响

研究粗放经营和集约经营条件下毛竹林蔗糖酶、纤维素酶、硝酸还原酶、脲酶和过氧化氢酶5种土壤酶活性对4种氮沉降水平(0、30、60和90 kg·hm^-2·a^-1)的响应.结果表明:与粗放经营相比,集约经营分别显著提高土壤蔗糖酶、纤维素酶和脲酶活性55.5%、112.9%和28.6%,显著抑制硝酸还原酶活性31.5%,对过氧化氢酶活性的影响不显著.氮沉降显著抑制粗放和集约经营方式下毛竹林蔗糖酶活性20.0%～49.4%和36.2%～45.1%、纤维素酶活性20.5%～46.3%和18.3%～49.0%、硝酸还原酶活性67.9%～85.2%和15.2%～34.2%,以及集约经营毛竹林脲酶活性23.1%～47.6%,显著增加了粗放经营毛竹林土壤脲酶活性8.1%～50.6%,对过氧化氢酶活性的影响不显著.氮沉降与经营方式的复合作用除对过氧化氢酶活性的影响不显著外,对其他4种土壤酶活性的影响均达到显著水平.     

氮沉降下杉木(Cunninghamia lanceolata)人工林凋落叶分解过程中C、N元素动态变化

在12年生的杉木(Cunninghamia lanceolata)人工林中开展模拟氮沉降试验,分N0(对照)、N1、N2、N3等4种处理,N沉降量依次为0、60、120、240 kg N hm-2 a-1,每处理重复3次.通过2 a的监测和分析发现,经N0、N1、N2、N3处理后,凋落物分解过程中C平均含量分别为46.47%、46.35%、46.79%、46.6%,各处理之间无显著差异,但均随着分解时间的增加呈下降趋势.氮沉降明显增加了凋落物中的N含量,且随着沉降水平的增加而增加.各处理凋落物C的分解系数依次为0.739、0.744、0.936、0.708,周转期为4.26 a、4.26 a、3.46 a、4.41 a;而N的分解系数分别为0.458、0.543、0.776、0.565,周转期为6.26 a、5.44 a、3.91 a、5.20 a.N1处理表现出促进N释放的作用,但对C释放影响不明显;N2处理促进了凋落物C、N元素的释放,而N3处理则表现出一定的抑制作用.氮沉降处理也明显降低了凋落物的C/N比,N1、N2、N3处理使C/N比分别比N0下降8.59%、14.20%和17.54%.