Canopy N and P dynamics of a southeastern US pine forest under elevated CO2

Forest production is strongly nutrient limited throughout the southeastern US. If nutrient limitations constrain plant acquisition of essential resources under elevated CO₂, reductions in the mass or nutrient content of forest canopies could constrain C assimilation from the atmosphere. We tested this idea by quantifying canopy biomass, foliar concentrations of N and P, and the total quantity of N and P in a loblolly pine (Pinus taeda) canopy subject to 4 years of free-air CO₂ enrichment. We also used N:P ratios to detect N versus P limitation to primary production under elevated CO₂. Canopy biomass was significantly higher under elevated CO₂ during the first 4 years of this experiment. Elevated CO₂ significantly reduced the concentration of N in loblolly pine foliage (5% relative to ambient CO₂) but not P. Despite the slight reduction foliage N concentrations, there were significant increases in canopy N and P contents under elevated CO₂. Foliar N:P ratios were not altered by elevated CO₂ and were within a range suggesting forest production is N limited not P limited. Despite the clear limitation of NPP by N under ambient and elevated CO₂ at this site, there is no evidence that the mass of N or P in the canopy is declining through the first 4 years of CO₂ fumigation. As a consequence, whole-canopy C assimilation is strongly stimulated by elevated CO₂ making this forest a larger net C sink under elevated CO₂ than under ambient CO₂. We discuss the potential for future decreases in canopy nutrient content as a result of limited changes in the size of the plant-available pools of N under elevated CO₂.     

大兴安岭次生林区优势种落叶松分布格局及竞争作用

竞争是形成特定群落结构、构成分布格局的基本驱动力之一,树种的空间分布和大小并不是相互独立的,而是广泛受到竞争过程影响。在大兴安岭设置一块具有代表性的天然次生林恢复样地,采用点格局分析中的最近邻体距离G（r）函数、双关联函数g（r）和基于个体胸径标记的点格局分析方法同时结合4种点格局零模型（完全随机模型、均值托马斯模型、随机标签模型、先决条件零模型）研究分析大兴安岭次生林区优势种落叶松不同生活史阶段的分布格局、关联性及竞争作用。研究结果显示：落叶松不同发育阶段分布格局不同、同一发育阶段不同研究尺度分布格局也不同;落叶松种群内存在潜在的竞争、且竞争作用与龄级和研究尺度有关;落叶松幼龄个体在小尺度（r<5m）下的聚集分布是由扩散限制作用和竞争作用所导致;落叶松种群内成龄与幼龄个体间有较强的正关联性、成龄个体对幼龄个体在小尺度下有明显的庇护作用;标记点格局分析方法在检测植物群落中的竞争作用时敏感性较高。     

松林中焦艺夜蛾幼虫的空间格局

应用聚集度指标和回归分析等方法,测定了焦艺夜蛾幼虫的空间格局。结果表明,焦艺夜蛾幼虫在马尾松林内呈聚集型分布,分布的基本成分为个体群,个体之间相互吸引;用马占山重新解释的Taylor幂法则分析表明,焦艺夜蛾幼虫的空间格局归属于“聚集度逆密度制约型”。     

暖温带森林优势种群的地形生境变异性

地形是影响植物种群分布格局、维持群落多样性的重要因子。以东灵山20hm2暖温带落叶阔叶林动态监测样地内的2个优势种群——辽东栎(Quercus wutaishanica)与胡桃楸(Juglans mandshurica)为研究对象,对比分析2种群同物种不同生长阶段、不同物种相同生长阶段群体的地形生境差异性。结果表明,样地内辽东栎种群主要分布在坡度大、南坡、西坡、凸出及高海拔的位置上,成年树比幼树更偏好凹地形;胡桃楸种群集中分布在坡度大、北坡、东坡、凹陷及低海拔的地形上,成年树相比幼树偏好分布在南坡、东坡及凹陷的位置。每个种群的幼树与成年树群体的地形因子之间差异都不显著(P>0.05),地形特征的分布范围一致;2物种成年树群体的海拔因子差异显著(P<0.05),幼树群体除了海拔外,南北坡向也表现了明显的差异(P<0.05)。同种群不同生长阶段地形因子的选择显示了物种地形生境的限制及保守性,不同种群同生长阶段地形因子的选择显示了物种间空间分布格局及多样性的维持。这对于理解暖温带森林的物种共存及生物多样性维持具有重要的意义。     

Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest

Summary The spatial distribution of seedlings of the dominant perennial plant species (Pinus halepensis, Cistus salviifolius, Rhus coriaria) and may annual species was studied after a wild fire in an eastern Mediterranean pine forest. The spatial distribution of all seedlings is affected by the location of the old burned pine trees. Seedling density of Pinus and Cistus is higher at a distance from the burned pine canopy and lower near the burned pine trunk. It is also higher beneath small burned pine trees than under big ones. Rhus seedling density is higher under big burned pine trees and also near the burned trunks. Seedlings of Pinus, Cistus and Rhus growing under the burned canopy of big pine trees tend to be taller than seedlings under small ones or outside the burned canopy. Most annual species germinate and establish themselves outside the burned canopies, and only a few annual species are found beneath them. It is suggested that variation in the heat of the fire, in the amount of ash between burned pine trees of different sizes, and in the distance from the burned canopy are responsible for the observed pattern of seedling distribution. The possible ecological significance of the spatial pattern of seedlings distribution and their differential growth rate are discussed.     

Spatial variability of soil properties under Pinus canariensis canopy in two contrasting soil textures

The aim of this study was to investigate how the biomass of extramatrical mycorrhizal mycelia (EMM) is influenced by the addition of different phosphorus (P), potassium (K) and PK fertilizers in peatland forests with variable P and K availability. Four fertilizers were used: apatite, biotite, Rauta-PK? (apatite and ferrosulphate) and a test fertilizer (50% apatite and 50% recycled iron phosphate). Forest plots with four different types of nutrient balance (deficient P and deficient K, deficient P and sufficient K, sufficient P and deficient K, and sufficient P and sufficient K) were studied. The effects on EMM biomass and ectomycorrhizal (EM) biomass in roots were estimated by ergosterol and phospholipid fatty acid (PLFA) analysis using in-growth mesh bags. Nutrients and rare-earth elements in EM roots surrounding the mesh bags were quantified and used as indicators of nutrient transport by the EMM in the mesh bags. The biomass of EMM was enhanced by P, K and PK deficiency of the trees, and EM fungal biomass in the roots was increased by P deficiency. The test fertilizer enhanced EMM biomass in all the plots studied, whereas the other fertilizers did not have any significant effect. No significant interactions between the P and K availability of host and mycelial fertilizer response could be detected. Deficiency of P or K or both in needles did not affect the concentrations of rare-earth elements in the tree roots. Earlier results from laboratory experiments have shown reduced carbon allocation to EM fungi under K deficiency, but this was not the case in these mature forests. Instead, we observed increased EMM biomass in response to both P and K deficiency.     

Changes in N and P availability and P fractions in Iwo soil from Nigeria under intensive cultivation

Summary Two N-P factorial experiments, one on a site previously in bush fallow and the other previously in arable crops, were used to follow changes in the P status of Iwo soils under continuous cultivation. Marked yield response of maize to N was found at both sites, with the response being relatively small in the early crop at the fallowed site. Yield response to P was found only at the cropped site and then only if N was applied. About 25% of the organic P in the surface soil was mineralized during the two cropping periods of the rainy season which resulted in release of about 3 times as much P as was taken up by the maize. Most of the mineralized P as well as any added P was converted to Fe and Al-P. Apparently the rate of organic P mineralization in this soil is rapid enough for a few years following clearing to maintain adequate P activity for plant growth whereas N deficiency shows up very early. Lecturer, Department of Soil Science, University of Ife, Nigeria, and Professor of Soil Science, University of Wisconsin, Madison, Wisconsin, formerly Professor of Soil Science, University of Ife, respectively. Lecturer, Department of Soil Science, University of Ife, Nigeria, and Professor of Soil Science, University of Wisconsin, Madison, Wisconsin, formerly Professor of Soil Science, University of Ife, respectively.     

Intraspecific functional trait variability across different spatial scales: a case study of two dominant trees in Korean pine broadleaved forest

Intraspecific functional trait variability plays an important role in the response of plants to environmental changes. However, it is still unclear how the variability differs across three nested spatial scales (individual, plot, and site) and which determinants (climatic, soil, and ontogenetic variables) shape the trait variability. Along a latitudinal gradient in Korean pine broadleaved forest of northeast China, we quantified the extent of intraspecific variability of four functional traits in two dominant trees Pinus koraiensis and Fraxinus mandshurica at eight sites, including specific leaf area, leaf dry matter content (morphological traits) and leaf nitrogen content, leaf phosphorus content (physiological traits). Results showed a large trait variation within and between species (coefficient variation: 6.07–23.3%). The leaf physiological traits of F. mandshurica and morphological traits of P. koraiensis were more responsive at site scale, while the morphological traits of F. mandshurica and physiological traits of P. koraiensis were more responsive at individual scale. In addition, abiotic and biotic factors explaining functional trait variation differ markedly between the two tree species, with physiological trait of F. mandshurica being more associated with climate and soil, while traits variability in P. koraiensis was not affected by climate, soil, and ontogeny, except for leaf phosphorus content. Overall, we can predict that the physiological traits of broadleaved species tend to be more sensitive to environmental changes, while pines are more sensitive to competition. It is critical to determine which spatial scale and trait type should be taken into account in predictive models of vegetation dynamics.     

生境梯度影响下的天然红松种群空间格局与种内关联

为理清生境梯度下天然红松种群空间分布规律,以黑龙江省凉水国家级自然保护区内自坡底至坡顶的四种生境(生境Ⅰ,谷地平坡潮湿生境;生境Ⅱ,坡下缓坡潮湿生境;生境Ⅲ,坡上斜坡半湿润生境;生境Ⅳ,坡顶陡坡半干旱生境)类型中建群种红松种群为研究对象。分析不同林层(主林层、亚林层、演替层、更新层)红松种群的数量特征,并基于O-ring函数,在重复采样条件下评价了红松种群空间分布格局。结果表明:(1)下坡位生境(生境Ⅰ、Ⅱ)红松更新层种群数量显著大于上坡位(生境Ⅲ、Ⅳ)(P<0.05),演替层中差异不显著;自坡底至坡顶的生境梯度上,主林层和亚林层中红松株数比例增加,红松种群年龄结构结构由倒J型向J型转变。(2)下坡位原始红松林更新层和演替层中红松种群聚集规模与聚集强度均高于上坡位,坡顶(生境Ⅳ)原始红松林更新层中红松种群聚集规模与聚集强度高于坡上(生境Ⅲ)。(3)四类生境的原始红松林中主林层与更新层中红松种群总体表现为空间独立,在个别尺度上表现出微弱的空间正关联或负关联;在0—5 m尺度上,下坡位红松林原始林内演替层与主林层红松种群表现为空间负关联,而上坡位生境林分对应林层间空间关系则为空间正关...     

Responses of fine roots and soil N availability to short-term nitrogen fertilization in a broad-leaved Korean pine mixed forest in northeastern China

Knowledge of the responses of soil nitrogen (N) availability, fine root mass, production and turnover rates to atmospheric N deposition is crucial for understanding fine root dynamics and functioning in forest ecosystems. Fine root biomass and necromass, production and turnover rates, and soil nitrate-N and ammonium-N in relation to N fertilization (50 kg N ha(-1) year(-1)) were investigated in a temperate forest over the growing season of 2010, using sequential soil cores and ingrowth cores methods. N fertilization increased soil nitrate-N by 16% (P<0.001) and ammonium-N by 6% (P<0.01) compared to control plots. Fine root biomass and necromass in 0-20 cm soil were 13% (4.61 vs. 5.23 Mg ha(-1), P<0.001) and 34% (1.39 vs. 1.86 Mg ha(-1), P<0.001) less in N fertilization plots than those in control plots. The fine root mass was significantly negatively correlated with soil N availability and nitrate-N contents, especially in 0-10 cm soil layer. Both fine root production and turnover rates increased with N fertilization, indicating a rapid underground carbon cycling in environment with high nitrogen levels. Although high N supply has been widely recognized to promote aboveground growth rates, the present study suggests that high levels of nitrogen supply may reduce the pool size of the underground carbon. Hence, we conclude that high levels of atmospheric N deposition will stimulate the belowground carbon cycling, leading to changes in the carbon balance between aboveground and underground storage. The implications of the present study suggest that carbon model and prediction need to take the effects of nitrogen deposition on underground system into account.     

中国四种森林类型主要优势植物的C∶N∶P化学计量学特征

为了评价不同森林类型的生态化学计量特征的差异,以吉林长白山温带针阔混交林、广东鼎湖山亚热带常绿阔叶林、云南西双版纳热带季雨林和江西千烟洲亚热带人工针叶林为研究对象,通过对2007年4月-2008年5月4种典型区域森林植物叶片和凋落物的碳(C)氮(N)磷(P)元素质量比与N、P再吸收率的分析,探讨了4种森林类型N、P养分限制和N、P养分再吸收的内在联系.结果表明:1)从森林类型上看,温带针阔混交林叶片的C∶N∶P为321∶13∶1,亚热带常绿阔叶林叶片的C∶N∶P为561∶22∶1,热带季雨林叶片的C∶N∶P为442∶19∶1,亚热带人工针叶林叶片的C∶N∶P为728∶18∶1;凋落物的C∶N∶P也是亚热带人工林最高,达1950∶27∶1,温带针阔混交林的最低,为552∶14∶1,热带季雨林的为723∶24∶1,亚热带常绿阔叶林的为1305∶35∶1,不同森林类型凋落物的C∶N∶P的计量大小关系与叶片的结果一致;2)从植物生活型上看,常绿针叶林叶片的C∶N均显著高于常绿阔叶林及落叶阔叶林;叶片C∶P与森林类型的关系并不十分密切;常绿阔叶林叶片的N∶P最高,常绿针叶林次之,落叶阔叶林最低;3)植物叶片的N∶P与月平均气温有显著的负相关关系,但叶片的C∶P基本不受月平均气温影响,叶片的C、N、P计量比与降水的线性关系不显著;4)高纬度地区的植物更易受N元素限制,而低纬度地区植物更易受P元素的限制;但受N或P限制的植物并不一定具有高的N或P再吸收率.研究结果表明,不同类型森林的叶片与凋落物的化学计量特征具有一致性,但是环境因子对不同类型植物化学计量比的影响并不相同.     

森林不同土壤层全氮空间变异特征

应用经典统计学和地统计学方法,分析了八达岭地区土壤全氮(TN)在不同层次(A,B,C)的空间变异特征。同时结合地理信息系统(GIS),分析了该地区植被类型和土壤TN之间的关系。应用分类回归树模型(classification and regression trees,CART)分析了土壤TN和海拔与植被分布格局的关系。得到以下结论:(1)TN在A、B、C层平均值分别为2.94、1.30,0.63 g/kg,变异系数(CV)分别为33%、33%、45%,都表现为中等变异。(2)TN在不同土层的变异函数理论模型符合球状模型,TN在A层为弱空间相关,在B、C层为中等空间相关。(3)泛可里格插值表明,TN在不同层次都表现出了明显的空间分布趋势。不同植被类型所对应土壤全氮的空间分布则各不相同。(4)CART研究结果表明,该区植被类型分布格局可大致划分为四大部分。可初步确定海拔725m,TN含量4.23 g/kg和5.69 g/kg为影响该区植被分布格局的重要参考值。     

中国四种森林类型主要优势植物的C:N:P化学计量学特征

为了评价不同森林类型的生态化学计量特征的差异, 以吉林长白山温带针阔混交林、广东鼎湖山亚热带常绿阔叶林、云南西双版纳热带季雨林和江西千烟洲亚热带人工针叶林为研究对象, 通过对2007年4月-2008年5月4种典型区域森林植物叶片和凋落物的碳(C)氮(N)磷(P)元素质量比与N、P再吸收率的分析, 探讨了4种森林类型N、P养分限制和N、P养分再吸收的内在联系。结果表明: 1)从森林类型上看, 温带针阔混交林叶片的C : N : P为321 : 13 : 1, 亚热带常绿阔叶林叶片的C : N : P为561 : 22 : 1, 热带季雨林叶片的C : N : P为442 : 19 : 1, 亚热带人工针叶林叶片的C : N : P为728 : 18 : 1; 凋落物的C : N : P也是亚热带人工林最高, 达1 950 : 27 : 1, 温带针阔混交林的最低, 为552 : 14 : 1, 热带季雨林的为723 : 24 : 1, 亚热带常绿阔叶林的为1 305 : 35 : 1, 不同森林类型凋落物的C : N : P的计量大小关系与叶片的结果一致; 2)从植物生活型上看, 常绿针叶林叶片的C : N均显著高于常绿阔叶林及落叶阔叶林; 叶片C : P与森林类型的关系并不十分密切; 常绿阔叶林叶片的N : P最高, 常绿针叶林次之, 落叶阔叶林最低; 3)植物叶片的N : P与月平均气温有显著的负相关关系, 但叶片的C : P基本不受月平均气温影响, 叶片的C、N、P计量比与降水的线性关系不显著; 4)高纬度地区的植物更易受N元素限制, 而低纬度地区植物更易受P元素的限制; 但受N或P限制的植物并不一定具有高的N或P再吸收率。研究结果表明, 不同类型森林的叶片与凋落物的化学计量特征具有一致性, 但是环境因子对不同类型植物化学计量比的影响并不相同。     

Litter decomposition of a pine plantation is affected by species evenness and soil nitrogen availability

Background and aims

Litter decomposition is a key process controlling flows of energy and nutrients in ecosystems. Altered biodiversity and nutrient availability may affect litter decomposition. However, little is known about the response of litter decomposition to co-occurring changes in species evenness and soil nutrient availability.

Methods

We used a microcosm experiment to evaluate the simultaneous effects of species evenness (two levels), identity of the dominant species (three species) and soil N availability (control and N addition) on litter decomposition in a Mongolian pine (Pinus sylvestris var. mongolica) plantation in Northeast China. Mongolian pine needles and senesced aboveground materials of two dominant understory species (Setaria viridis and Artemisia scoparia) were used for incubation.

Results

Litter evenness, dominant species identity and N addition significantly affected species interaction and litter decomposition. Higher level of species evenness increased the decomposition rate of litter mixtures and decreased the incidence of antagonistic effects. A. scoparia-dominated litter mixtures decomposed faster than P. sylvestris var. mongolica- and S. viridis-dominated litter mixtures. Notably, N addition increased decomposition rate of both single-species litters and litter mixtures, and meanwhile altered the incidence and direction of non-additive effects during decomposition of litter mixtures. The presence of understory species litters stimulated the decomposition rate of pine litters irrespective of N addition, whereas the presence of pine litters suppressed the mass loss of A. scoparia litters. Moreover, N addition weakened the promoting effects of understory species litters on decomposition of pine litters.

Conclusions

Pine litter retarded the decomposition of understory species litters whereas its own decomposition was accelerated in mixtures. Nitrogen addition and understory species evenness altered species interaction through species-specific responses in litter mixtures and thus affected litter decomposition in Mongolian pine forests, which could produce a potential influence on ecosystem C budget and nutrient cycling.     

Correlations among indices of forest soil nutrient availability in fertilized and unfertilized loblolly pine plantations

Summary Various laboratory indices of N and P availability in forest soils correlated poorly among themselves and with on-site ion exchange resin (IER) estimates in both unfertilized and N+P fertilized loblolly pine plantations. IER nutrient availability estimates had greatest within-site variability than laboratory indices. Net nitrification was minimal in laboratory incubation of the mineral soil despite high rates of ammonification. In contrast, IER NO₃–N values were usually of the same magnitude as IER NH₄–N values. In both fertilized and unfertilized stands, at least one N availability index was negatively correlated with at least one P index. Soil N and P availabilities were generally higher on fertilized plots than on unfertilized plots 3.5 years after fertilization, and IER estimates showed the greatest number of plots with increased levels. The greater ability of the IER method to distinguish between fertilized and unfertilized plots indicated the method was affected by on-site factors that the laboratory methods do not assess.     

京东板栗主产区土壤氮磷钾的空间变异

采用经典统计学和地统计学相结合的方法,对京东板栗主产区迁西县土壤氮、磷、钾的空间变异进行了分析.结果表明：迁西县土壤全量氮、磷、钾和速效氮、钾的含量普遍较低,但表层土壤速效磷含量较高;氮、磷、钾的空间分布均表现为中等变异,且以磷的变异最大.表层土壤全氮和碱解氮的分布符合高斯模型,其空间变异主要由结构性因素决定;全钾和速效钾的分布分别符合球状模型和高斯模型,前者空间变异受结构性因素和随机性因素的共同影响,空间相关性为中等;后者主要受结构性因素的影响,空间相关性强烈.氮、钾在全县范围内的空间分布特征相似,高值区均出现在县域南部和西北部,而中部和东北部的含量较低.全磷的高值区主要分布在该县北部,速效磷高值区则分布在县区南部.氮、钾全量和有效态含量之间存在极显著正相关关系,但全磷和速效磷含量的相关性不显著.     

Regulation of soil phosphatase and chitinase activityby N and P availability

Soil microorganisms and plants produce enzymes thatmineralize organically bound nutrients. When nutrientavailability is low, the biota may be able to increase production ofthese enzymes to enhance the supply of inorganicnitrogen (N) and phosphorus (P). Regulation of enzyme productionmay be a point where N and P cyclesinteract. We measured acid phosphatase and chitinase(N-acetyl ß-D-glucosaminide) activity in soilacross a chronosequence in Hawaii where N and Pavailability varies substantially among sites and longterm fertilizer plots had been maintained for over 4years.Phosphatase activity was high at all sites. Chitinaseactivity decreased significantly as age and Navailability increased across the chronosequence.Phosphorus addition suppressed phosphatase activity atall sites, while N addition increased phosphataseactivity at the young, N-limited site. In contrast,N addition repressed chitinase activity only at the Nlimited young site, and P additions had no effect onchitinase activity. These results suggest that theregulatory relationship between nutrient supply andnutrient mineralization are asymmetric for N and P,and that the differences could help to explaindifferences observed in patterns of N and Pavailability.     

Concordance between plant species,oribatid mites and soil in a Mediterranean stone pine forest

Biological interactions between above-ground and below-ground organisms are not clearly defined among communities with regard to compositional patterns. The study investigates the concordance of species assemblages between vascular plants and oribatid mites and soil chemical properties with special attention to the role of vegetation structure, i.e. tree, shrub and herbaceous cover, for biological components. Data were collected in a Mediterranean coastal Nature Reserve using sampling design based on random selection of plots with cover of stone pine (Pinus pinea L.) exceeding 15%. Agreement of distribution patterns was verified by Spearman’s rank correlation coefficient applied to pairs of matrices of plot scores by principal component analysis (plants, mites and soil) and the Mantel test. The feasible role of vegetation cover on plant and mite assemblages was tested by redundancy analysis (RDA). Significant correlations were found for biological assemblages, indicating congruent plant–mite compositional patterns. On the other hand, the hypothesis of concordance between biological communities and soil was rejected. Moreover, RDA showed that vegetation cover was a driver of both plant and oribatid mite assemblages. In particular, herbaceous cover proved to be a good proxy for the two biological communities investigated, with different taxa linked to forest clearings and to areas with denser tree cover. Our results indicate that soil features were not of primary importance for below-ground and above-ground community assemblages in the study area. In the light of our findings and ongoing threats in coastal areas, we recommend that management measures be directed at maintenance of diversified vegetation structure, which may ensure above-ground and below-ground biodiversity with diverse biological community assemblages.