华山松林凋落物养分释放及土壤生态化学计量特征对模拟氮沉降的短期响应

研究模拟氮(N)沉降下森林生态系统凋落物-土壤C/N/P化学计量特征,对探究在全球气候变化背景下森林生态系统物质循环内在机理具有重要科学意义.以滇中亚高山华山松林(Pinus armandii forest)为研究对象,采用尼龙网袋法于2018年2月—2019年1月在华山松林开展模拟N沉降下凋落叶、枝原位分解试验,分别...     

氮沉降下鼎湖山森林凋落物分解及与土壤动物的关系

研究了南亚热带3种森林生态系统凋落物在N沉降下的分解动态及其与土壤动物群落的关系。选取季风常绿阔叶林、针阔混交林和马尾松林建立野外模拟N沉降样地,实施四个处理组,对照(Control)、低氮(50kg·hm-2·a-1,LowN)、中氮(100kg·hm-2·a-1,MediujmN)和高氮处理(150kg·hm-2·a-1,HighN),利用凋落物网袋法,在18个月的时间内调查分析了凋落物分解过程及其中的土壤动物密度特征。研究结果表明,植被演替阶段对凋落物的分解速度存在影响,季风林凋落物降解速度显著性快于混交林和针叶林(P<0.05);18个月后,季风林各处理地凋落物残留率为0.05、0.14、0.13和0.17,混交林为0.64、0.56和0.62,针叶林为0.66、0.63和0.62。N沉降增加对凋落物分解存在一定影响。且这种影响与植被类型之间存在明显的交互作用。N沉降处理对季风林凋落物分解表现出了一定的抑制作用,而且这种差异随时间推移愈益明显,但在混交林和针叶林内,试验后期凋落物分解受到了N沉降处理的促进作用。在试验后期,尤其是12个月后,凋落物网袋土壤动物密度在不同林地和不同水平N处理下体现了差异化发展趋势。在季风林内,N处理地土壤动物密度受到了明显的抑制;在混交林和针叶林内,低N样地动物密度显示了相比对照样地的明显优势,但在较高强度的中N处理地无论在凋落物的降解速率还是在动物密度上都与对照样地没有明显差别。文章认为,N沉降处理所产生的影响可能受环境N饱和程度的调控。文章还提出,在凋落物分解进程中,土壤动物群落具有“后期进入”特征,这对于进一步准确分析森林凋落物分解进程及土壤动物的贡献有重要意义。     

中国森林凋落叶氮、磷化学计量特征及控制因素

建立中国森林凋落叶养分浓度及其化学计量比数据库,分析养分浓度及其化学计量比与主要环境因素之间的关系,对预测中国森林生态系统生物地球化学循环具有重要意义.通过收集已报道的中国森林凋落叶氮(N)、磷(P)浓度及其相关变量,探讨地理因素(纬度,LAT)、气候因素(年平均气温,MAT和年平均降水量,MAP)和叶特性(常绿与落叶、阔叶与针叶)对中国森林凋落叶N、P和N/P的影响.结果显示,N浓度和N/P随LAT的升高而降低,P浓度随LAT的升高而升高;N浓度和N/P随MAT和MAP的升高而升高,而P浓度随MAT和MAP的升高而降低;常绿树种和落叶树种N浓度差异不显著,落叶树种P浓度比常绿树种高53%,而N/P比常绿树种低57%;相反,阔叶树种N浓度比针叶树种的高37%,而P浓度和N/P在两者之间没有显著差异.综上,中国森林凋落叶N、P及N/P受环境因素和叶特性综合影响,特别是气候因素对凋落叶P浓度和N/P的影响尤为显著,这为预测全球气候变暖背景下森林物质循环提供了理论依据.     

晋西吕梁山区3种森林碳氮磷生态化学计量特征

以吕梁山区3种人工林(山杨林、落叶松林和油松林)为研究对象,采用标准样地的实测数据,探索植物叶片、枯落物及表层(0-20 cm)土壤的碳(C)、氮(N)、磷(P)生态化学计量特征,并进行相关性分析.结果显示,不同森林类型同一组分C、N、P含量差异显著,叶片、枯落物、土壤的C、N含量均为山杨林大于落叶松林和油松林,P含量为落叶松林大于山杨林和油松林.3种森林C、N、P含量均为叶片枯落物土壤,且叶片与枯落物C、N、P含量显著高于土壤;C:N、C:P均表现为枯落物叶片土壤,N:P则表现为叶片土壤枯落物.山杨林枯落物N:P与土壤N:P呈现显著正相关;落叶松林叶片C:N与枯落物N:P呈现显著负相关,叶片C:P与土壤N:P呈现显著正相关;油松林叶片N:P与土壤N:P呈现显著正相关.以3种森林类型总体来说,叶片与土壤N含量呈现显著正相关,而枯落物与土壤C、N、P之间均无显著相关.上述研究表明,环境因素对土壤C、N、P计量特征的影响较大,尤其是纬度和海拔对土壤C、N、P及C:N、C:P的影响最为显著,且均为显著正相关;结果可为进一步研究该地区不同树种的养分利用和循环特征提供科学依据.     

植被因子和土壤氮对南亚热带常绿阔叶次生林细根生物量的影响

森林细根生物量与乔木层胸高断面积关系,以及在高大气N沉降背景下细根对土壤氮的响应程度,目前仍不明确。本研究选择广州市受保护40 a左右的南亚热带常绿阔叶次生林为研究对象,在全市范围内设置了48个森林样地,开展乔木层、灌草层、细根(直径≤2 mm)、土壤C含量和N含量的调查,研究乔木胸高断面积、乔木密度、灌草层生物量、土壤N含量、土壤单位碳的N含量与细根生物量之间的关系,探讨地上植被因子和土壤N对细根生物量的影响。结果表明,(1)土壤表层(0~20 cm)和土壤下层(20~40 cm)细根生物量与乔木层总胸高断面积均不相关,但与胸径30 cm以上乔木的胸高断面积所占比例负相关。(2)表层细根生物量与灌草层生物量不相关,下层细根生物量与灌草层生物量负相关。(3)除了表层细根生物量与相应土层土壤N含量不相关外,下层细根生物量与相应土层土壤N含量,以及表层和下层细根生物量与相应土层单位碳的N含量均负相关。研究表明,对于林木胸径组成差异大的南亚热带常绿阔叶次生林,对细根生物量产生影响的是胸径30 cm以上乔木的胸高断面积所占全部乔木总胸高断面积比例,而不是乔木层总胸高断面积。细根生物量与土壤N含量负相关,表明即使在高大气N沉降背景下,南亚热带常绿阔叶次生林的森林植被仍对土壤N存在响应。     

亚热带4种林木幼苗的碳氮磷含量及其化学计量比特征

以亚热带区域4种主要森林植物杉木(Cunninghamia lanceolata)、马尾松(Pinus massoniana)、木荷(Schima superba)、米槠(Castanopsis carlesii)幼苗为研究对象,分别测定其不同器官碳(C)、氮(N)、磷(P)含量,并计算其化学计量比.结果表明:(1)针叶树种C、N含量以及C/P、N/P高于阔叶树种,P含量及C/N则低于阔叶树种.(2)杉木叶片N、P含量及茎P含量高于马尾松,但各器官的C/N与C/P低于马尾松;米槠根N含量、C/P、N/P高于木荷.(3)4种林木幼苗的C、N含量及C/P、N/P均以叶为最高,根的C、N含量为最低,茎的C/P和N/P为最低;P含量以茎为最高,叶为最低;C/N以根为最高,叶为最低.本研究结果可为我国湿润亚热带区域森林植物苗木培育、幼林养分管理及人工造林时立地选择等提供参考价值.     

降水变化、氮添加对鼎湖山主要森林土壤有机碳矿化和土壤微生物碳的影响

全球变化对土壤有机碳(SOC)存贮与分解的影响在全球碳(C)循环中具有重要地位.分别通过室内土壤培养法和氯仿熏蒸法,研究了降水变化和氮(N)添加处理对鼎湖山3种不同演替阶段的季风常绿阔叶林、针阔混交林和马尾松针叶林SOC矿化和土壤微生物量碳(SMBC)的影响.结果表明:1)降水量增加能够提高森林演替晚期SOC累积矿化量和矿化速率,而对森林演替早期SOC累积矿化量和矿化速率没有显著影响(P>0.05).2)干旱条件(降水量减少)降低森林SMBC含量,且在鼎湖山季风林表层土壤(0～10 cm)中SMBC的减少达到显著水平(P<0.05).3)N添加处理对鼎湖山3种森林类型SOC累积矿化量、矿化速率以及SMBC都没有显著影响(P>0.05).未来关于SOC矿化对全球变化响应的研究,要综合考虑土壤有机质质量、C/N比例、外源性氮输入等因素的作用.图4表2参37     

南亚热带森林优势树种氮、磷可利用性与菌根侵染率的关系

氮(N)和磷(P)是植物生长所必需的大量元素,其供应是否充足对植物生长发育具有重要影响。菌根真菌侵染有助于植物对N、P等营养元素的吸收利用,而以往研究大多集中于草地、农田生态系统中,并多以控制实验或者室内盆栽实验为主。选择南亚热带森林演替前期与演替中期的共有优势树种马尾松(Pinus massoniana)以及演替中期与演替顶级阶段的共有优势树种木荷(Schima superba)与锥栗(Castanopsis chinensis)为研究对象,测定其菌根侵染率、根际土酸性磷酸酶活性(acid phosphatase activity,APA)、根际土以及叶片元素含量,探讨不同演替阶段森林共有优势树种N、P可利用性与菌根真菌侵染率之间的关系。结果表明:(1)马尾松菌根侵染率由演替前期的48.18%显著增加到演替中期的65.7%;木荷以及锥栗菌根侵染率分别由演替中期的57.7%、50.79%增加到演替顶级的64.03%、53.18%;(2)共有优势树种菌根侵染率与叶片磷含量呈显著负相关关系,马尾松菌根侵染率与根际土铵态氮含量呈显著正相关关系;(3)共有优势树种根际土APA、叶片N:P均随着演替的进行而升高,木荷和锥栗根际土APA分别由演替中期的8.8、9.38μmol·g~(-1)·h~(-1)显著增加到演替顶级的16.96、15.55μmol·g~(-1).h~(-1);共有优势树种根际土APA与有效磷含量存在显著负相关关系,而与根际土C:P、N:P以及叶片N:P间存在极显著正相关关系。可见,随着南亚热带森林演替的进行,磷限制的增强将使得不同演替阶段森林共有优势树种菌根侵染率升高,菌根真菌的侵染能够缓解不同演替阶段森林共有优势树种所受的磷限制。     

不同施肥处理对棕壤N2O排放量的影响

采用田间试验研究不同施肥处理对棕壤N2O排放量的影响。结果表明,N2O释放量随着耕层土壤硝态氮含量增加而上升。不同施肥处理对N2O排放量影响不同,低氮处理(N1)排放量(整个玉米生育期按185d计算)为1.18kg·hm-2,高氮处理(N2)为2.39kg·hm-2。随着施氮量的增加,反硝化作用加强,N2O排放量上升,以N2O形式损失加剧。相同施氮水平条件下,随着有机肥施入量的增加土壤N2O排放量上升,其中以高氮高有机肥处理(M2N2)N2O排放量最高,达到了7.05kg·hm-2,占所投入氮肥的2.34%。相同氮素供应水平条件下增施磷、钾肥,也会增加N2O排放量。整个玉米生育时期通过N2O排放损失的肥料占投入氮肥比例为0.99%～2.46%。     

亚热带3种典型常绿森林土壤和植物叶片碳氮磷化学计量特征

为深入了解亚热带常绿森林生态系统养分循环和系统稳定机制,以四川省宜宾市老君山国家级自然保护区内3种典型森林(水杉和柳杉人工林、中华木荷和鸡爪槭次生林以及总状山矾和天全钓樟原始常绿阔叶林)作为研究对象,研究其表层土壤(0-10 cm)和乔木、灌木、草本生活型植物叶片的C、N、P化学计量特征.结果表明:(1)表层土壤C、N、P含量以原始林最高,人工林最低,人工林土壤C:N最高,次生林土壤C:P最高,原始林土壤N:P最高;(2)乔木叶片C、N、P含量最高,草本植物最低;人工林乔木叶片C:N、C:P比最高;乔木、灌木、草本N:P比分别为13.9、14.1、9.3;人工林乔木N:P(10.2)最低,次生林乔木N:P(14.5)与原始林(13.8)较高;(3)植被整体及乔木、灌木、草本整体叶片N、P含量间Ⅱ类线性回归斜率约等于1,表明叶片N与P含量呈等速投入关系.可见,乔、灌木整体更易受P限制,草本更易受N限制;在土壤N、P供应较为贫瘠的人工林受N限制更为明显,而物种较为丰富、土壤养分供应较为充足的天然林P限制更为明显.(图4表3参37)     

岷江干旱河谷灌木幼苗的土壤养分适应性研究

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

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

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

Nitrogen to phosphorus ratios of two understory plant species in response to nitrogen and phosphorus addition in tropical forest of southern China北大核心CSCD

The secondary tropical forests in southern China have suffered from frequent human disturbance and increasing high N deposition. In order to explore the nutrient limitation status in secondary tropical forests of South China, this 3-year field experiment of nitrogen (+N) and phosphorus (+P) addition investigated nitrogen (N) and phosphorus (P) concentrations of the aboveground tissue (leaf and branch) of two widely distributed understory native species Clerodendrum cyrtophyllum and Uvaria microcarpa in a secondary tropical forest of South China. The results showed that: 1) the N and P concentrations of the two species were significantly different (P < 0.001); N and P concentrations of different tissues in the same species were different; N&P addition greatly affected N and P concentrations in branch rather than new leaf and older leaf. 2) +N treatment had no significant effect on N or P concentrations of either species, but significantly decreased N:P ratios (P = 0.001), at the level of 9% for C. cyrtophyllum and 50% for U. microcarpa, respectively. 3) +P treatment had no significant effect on tissue N concentrations, but significantly increased plant P concentrations (P < 0.001), at 54% for C. cyrtophyllum and 88% for U. microcarpa, respectively; +P treatment significantly decreased plant N:P ratios (P < 0.001), at 28% and 60%, respectively. 4) The alterations of P concentrations of two species had significantly negative correlations with N:P alterations under +N/+P treatment (P < 0.001), suggesting that the alteration of P concentrations in plant tissue was the major driver for N:P alteration. Our results show that N and P addition would affect tissue N and P concentrations of the two species, with +P treatment having relatively greater effect on nutrient concentrations than +N treatment; the branch is more sensitive than new or older leaf in response to nutrient addition. Therefore, P availability may be the limiting factor for plant growth in the tropical forests.     

Nutrient utilization by Casuarina equisetifolia plantation of different ages in the tropical coastal area of South China北大核心CSCD

Casuarina equisetifolia plantation plays a key role in protecting coastal areas from hazardous climate. However, the plantations in the tropical coastal area of south China have degraded severely in recent years. This research aimed to investigate the nutrient status of the plantation ecosystem along a chronological sequence. The results showed that different parts of the Casuarina equisetifolia had very similar level of Carbon (C), 448-462 g kg-1 in the branch and trunk, 416-430 g kg-1 in the leaf and shed leaf, 320-391 g kg-1 in the fine root. Carbon content did not vary with the plantation age. High fine root biomass did not definitely lead to high soil carbon stock. Casuarina equisetifolia had Nitrogen (N) content of 9.9-11.9 g kg-1, with the highest N found in the leaf and fine root. The Phosphorus (P) content was in the order of leaf > fine root > trunk. The plantation in fast growth period of age 6 had the lowest N and P. The soil of 3-year plantation had the highest P content among the 4 age classes, which also resulted in the highest soil C and N content in plantation of 3 years among all. However, the C and N stock of the sandy soil was extremely low compared to normal soil of the region. Soil N was weakly correlated with leaf N, but soil P not correlated with leaf P. Except for the obvious dynamics of C/N and C/P ratios in the leaf, which showed a peak in 6-year plantations, the C/N and C/P ratios of different organs did not change with the plantation age. Casuarina equisetifolia retranslocated nutrients from aging leaf at a rate of 18-30% for N and 43-58% for P. The nutrient resorption efficiency was not correlated with nutrient level in either soil or plant. In summary, Casuarina equisetifolia has low level of nutrient status. The plantation growth is limited by N and P in young period, but by P in relatively older period.     

不同林龄木麻黄人工林生物多样性与土壤养分状况研究

木麻黄（Casuarina equisetifolia）是我国东南沿海海岸防护林骨干树种。本文对广东省茂名市木麻黄防护林开展了不同林龄的种群结构、植物多样性、以及土壤养分特征的研究。结果表明,滨海沙地木麻黄群落在生长过程中有明显的自疏现象,18年林个体数（975株·hm-2）不足3年林个体数一半（2350株·hm-2）,3年至6年龄木麻黄生长最快。调查林地内共有植物27种,其中灌木8种,草本植物18种。随林龄的增长,林下植物种数显著增加,多样性指数、均匀度指数逐步增加,优势度指数下降。林地土壤有机碳、全氮与速效氮供应水平极低;土壤磷供应相对较好,表层有效磷一般高于3.5mg·kg-1。土壤有效养分与植物多样性显著相关,显示养分是植物定居的主要限制因素。结果表明,木麻黄林结构简单,生物多样性低,土壤养分贫乏。     

Canopy interception of atmospheric chloride ion during precipitation processes in the subalpine Picea asperata plantation in the upper reaches of the Yangtze River北大核心CSCD

Increasing concentrations of chloride ions (Cl-) in the atmosphere is one of the most important global environmental factors influencing processes in terrestrial ecosystems. To understand the roles played by plantations in intercepting atmospheric Cl- in mountainous regions, a one-year field observation of precipitation and throughfall was conducted in a subalpine artificial Picea asperata plantation in western Sichuan and the dynamics of Cl- concentrations in rainfall, snowfall, and throughfall, and canopy interception on atmospheric Cl- were investigated from August 2015 to July 2016. During the observational year, 27 rainfalls and 6 snowfalls were observed and sampled. The average concentration of Cl- in the precipitation was 1.33 mg/L and the average concentrations of Cl- in rainfall and snowfall were 1.41 mg/L and 0.98 mg/L, respectively. Correspondingly, the total input of Cl- by the precipitation processes was 7.56 kg/hm2, which included 6.31 kg/ hm2 by rainfall and the remainder by snowfall. In one entire year, the total canopy interception of Cl- was 2.61 kg/hm2 and the average interception rate was 38.08%. During the rainy season, the canopy interception of Cl- was 2.20 kg/hm2 and the average rate was 38.90%, whereas during the snow season, the canopy interception of Cl- was 0.41 kg/hm2 and the average rate was 34.39%. The maximum and minimum interception rates occurred in August and September, respectively. Significantly negative correlation was found between the interception rate of Cl- and precipitation. In summary, the P. asperata plantation in the subalpine forest region of western Sichuan exerts an important role in intercepting atmospheric Cl-, which is of great significance for maintaining and improving the environment of the water conservation region in the upper reaches of the Yangtze River. Furthermore, changes in the precipitation pattern caused by global changes might affect the role of the artificial forest canopy interception on atmospheric Cl-. © 2018 Science Press. All rights reserved.     

Progressive N limitation in forests: review and implications for long-term responses to elevated CO2

Field studies have shown that elevated CO2 can cause increased forest growth over the short term (<6 years) even in the face of N limitation. This is facilitated to some degree by greater biomass production per unit N uptake (lower tissue N concentrations), but more often than not, N uptake is increased with elevated CO2 as well. Some studies also show that N sequestration in the forest floor is increased with elevated CO2. These findings raise the questions of where the "extra" N comes from and how long such growth increases can continue without being truncated by progressive N limitation (PNL). This paper reviews some of the early nutrient cycling literature that describes PNL during forest stand development and attempts to use this information, along with recent developments in soil N research, to put the issue of PNL with elevated CO2 into perspective. Some of the early studies indicated that trees can effectively "mine" N from soils over the long term, and more recent developments in soil N cycling research suggest mechanisms by which this might have occurred. However, both the early nutrient cycling literature and more recent simulation modeling suggest that PNL will at some point truncate the observed increases in growth and nutrient uptake with elevated CO2, unless external inputs of N are increased by either N fixation or atmospheric deposition.     

不同恢复年限刺槐林土壤碳、氮、磷含量及其生态化学计量特征

地处西秦岭山地的甘肃天水吕二沟小流域土壤侵蚀问题严重,上世纪五六十年代开始,当地政府在研究区实施了多年的生态恢复措施以保持水土,但目前尚未有植被恢复对土壤生态计量特征的系统调查.分析不同林龄人工林土壤碳、氮、磷含量及其生态化学计量特征可在一定程度上揭示土壤养分的限制情况.选取黄土丘陵沟壑区不同生长年限(5、20、40、...     

No evidence that chronic nitrogen additions increase photosynthesis in mature sugar maple forests

Atmospheric nitrogen (N) deposition can increase forest growth. Because N deposition commonly increases foliar N concentrations, it is thought that this increase in forest growth is a consequence of enhanced leaf-level photosynthesis. However, tests of this mechanism have been infrequent, and increases in photosynthesis have not been consistently observed in mature forests subject to chronic N deposition. In four mature northern hardwood forests in the north-central United States, chronic N additions (30 kg N ha(-1) yr(-1) as NaNO3 for 14 years) have increased aboveground growth but have not affected canopy leaf biomass or leaf area index. In order to understand the mechanism behind the increases in growth, we hypothesized that the NO3(-) additions increased foliar N concentrations and leaf-level photosynthesis in the dominant species in these forests (sugar maple, Acer saccharum). The NO3(-) additions significantly increased foliar N. However, there was no significant difference between the ambient and +NO3(-) treatments in two seasons (2006-2007) of instantaneous measurements of photosynthesis from either canopy towers or excised branches. In measurements on excised branches, photosynthetic nitrogen use efficiency (micromol CO2 s(-1) g(-1) N) was significantly decreased (-13%) by NO3(-) additions. Furthermore, we found no consistent NO3(-) effect across all sites in either current foliage or leaf litter collected annually throughout the study (1993-2007) and analyzed for delta 13C and delta 18O, isotopes that can be used together to integrate changes in photosynthesis over time. We observed a small but significant NO3(-) effect on the average area and mass of individual leaves from the excised branches, but these differences varied by site and were countered by changes in leaf number. These photosynthesis and leaf area data together suggest that NO3(-) additions have not stimulated photosynthesis. There is no evidence that nutrient deficiencies have developed at these sites, so unlike other studies of photosynthesis in N-saturated forests, we cannot attribute the lack of a stimulation of photosynthesis to nutrient limitations. Rather than increases in C assimilation, the observed increases in aboveground growth at our study sites are more likely due to shifts in C allocation.     

Potassium, phosphorus, or nitrogen limit root allocation, tree growth, or litter production in a lowland tropical forest

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 x 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.