Seasonal dynamics of fine root biomass,root length density,specific root length,and soil resource availability in a Larix gmelinii plantation

Fine root tumover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors.Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past,our understanding of it remains limited.This is because the dynamics processes associated with soil resources availability are still poorly understood.Soil moisture,temperature,and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level.In temperate forest ecosystems,seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground.Therefore,fine root biomass,root length density(RLD)and specific root length(SRL)vary during the growing season.Studying seasonal changes of fine root biomass,RLD,and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover.The objective of this study was to understand whether seasonal variations of fine root biomass,RLD and SRL were associated with soil resource availability,such as moisture,temperature,and nitrogen,and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation.We used a soil coring method to obtain fine root samples(≤2 mm in diameter)every month from Mav to October in 2002 from a 17-year-old L.gmelinii plantation in Maoershan Experiment Station,Northeast Forestry University,China.Seventy-two soil cores(inside diameter 60 mm;depth intervals:0-10 cm,10-20 cm,20-30 cm)were sampled randomly from three replicates 25 m×30 m plots to estimate fine root biomass(live and dead),and calculate RLD and SRL.Soil moisture,temperature,and nitrogen(ammonia and nitrates)at three depth intervals were also analyzed in these plots.Results showed that the average standing fine root biomass(live (32.2 g.m-2.a-1)in the middle(10-20 cm)and deep layer (20-30cm),respectively.Live and dead fine root biomass was the highest from May to July and in September,but lower in August and October.The live fine root biomass decreased and dead biomass increased during the growing soil layer.RLD and SRL in May were the highestthe other months,and RLD was the lowest in Septemberdynamics of fine root biomass,RLD,and SRL showed a close relationship with changes in soil moisture,temperature,and nitrogen availability.To a lesser extent,the temperature could be determined by regression analysis.Fine roots in the upper soil layer have a function of absorbing moisture and nutrients,while the main function of deeper soil may be moisture uptake rather than nutrient acquisition.Therefore,carbon allocation to roots in the upper soil layer and deeper soil layer was different.Multiple regression analysis showed that variation in soil resource availability could explain 71-73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass.These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability,which resulted in an increased allocation of carbohydrate to these roots,but a lower allocation of carbohydrate to those in soil with lower resource availability.     

The dependence of root system properties on root system biomass of 10 North American grassland species

Dependence of the properties of root systems on the size of the root system may alter conclusions about differences in plant growth in different environments and among species. To determine whether important root system properties changed as root systems aged and accumulated biomass, we measured three important properties of fine roots (tissue density, diameter, and C:N) and three biomass ratios (root:shoot, fine:coarse, and shallow:deep) of monocultures of 10 North American grassland species five times during their second and third years of growth. With increasing belowground biomass, root tissue density increased and diameter decreased. This may reflect cortical loss associated with the aging of roots. For non-legumes, fine root C:N decreased with increasing root biomass, associated with decreases in soil solution NO^{3 –} concentrations. No changes in fine root C:N were detected with increasing belowground biomass for the two legumes we studied. Among all 10 species, there were generally no changes in the relative amounts of biomass in coarse and fine roots, root:shoot, or the depth placement of fine roots in the soil profile as belowground biomass increased. Though further research is needed to separate the influence of root system size, age of the roots, and changes in nutrient availability, these factors will need to be considered when comparing root functional traits among species and treatments.     

落叶松人工林细根动态与土壤资源有效性关系研究

树木细根在森林生态系统C和养分循环中具有重要的作用。由于温带土壤资源有效性具有明显的季节变化, 导致细根生物量、根长密度 (Rootlengthdensity, RLD) 和比根长 (Specificrootlength, SRL) 的季节性变化。以 17年生落叶松 (Larixgmelini) 人工林为研究对象, 采用根钻法从 5月到 10月连续取样, 研究了不同土层细根 (直径≤ 2mm) 生物量、RLD和SRL的季节动态, 以及这些根系指标动态与土壤水分、温度和N有效性的关系。结果表明 :1) 落叶松细根年平均生物量 (活根 +死根 ) 为 189.1g·m-2 ·a-1, 其中 5 0 %分布在表层 (0~ 10cm), 33%分布在亚表层 (11~ 2 0cm), 17%分布在底层 (2 1~ 30cm) 。活根和死根生物量在 5~ 7月以及 9月较高, 8月和 10月较低。从春季 (5月 ) 到秋季 (10月 ), 随着活细根生物量的减少, 死细根生物量增加 ;2 ) 土壤表层 (0~ 10cm) 具有较高的RLD和SRL, 而底层 (2 1~ 30cm) 最低。春季 (5月 ) 总RLD和SRL最高, 分别为 10 6 2 1.4 5m·m-3 和 14.83m·g-1, 到秋季 (9月 ) 树木生长结束后达到最低值, 分别为 2 198.2 0m·m-3 和 3.77m·g-1;3) 细根生物量、RLD和SRL与土壤水分、温度和有效N存在不同程度的相关性。从单因子分析来看, 土壤水分和有效N对细根的影响明显大于温度, 对活根的影响大于死根。由于土壤资源有效性的季节变化, 使得C的地下分配格局发生改变。各土层细根与有效性资源之间的相关性反映了细根功能季节性差异。细根 (生物量、RLD和SRL) 的季节动态 (5 8%~ 73%的变异 ) 主要由土壤资源有效性的季节变化引起。     

Fine-root biomass and soil properties in a semideciduous and a lower montane rain forest in Panama

The distribution of root biomass and physical and chemical properties of the soils were studied in a semideciduous and in a lower montane rain forest in Panama. Roots and soil samples were taken by means of soil cores (25 cm deep) and divided into five, 5-cm deep sections. Soils were wet-sieved to retrieve the roots that were classified in four diameter classes: very fine roots (<1 mm), fine roots (1–2 mm), medium roots (2–5 mm) and coarse roots (5–50 mm). Soil samples were analyzed for organic carbon, total nitrogen, available phosphorus, exchangeable bases, cation exchange capacity, pH, aluminium and exchangeable acidity. Total root biomass measured with the soil corer (roots <50 mm in diameter) was not different between the forests (9.45 t ha^-1), while biomass of very fine roots was larger in the mountains (2.00 t ha^-1) than in the lowlands (1.44 t ha^-1). The soils in the semideciduous forest were low in available phosphorus, while in the mountains, soils had low pH, high exchangeable aluminium and exchangeable acidity, and low concentration of exchangeable bases. Phosphorus was in high concentration only in the first 5 cm of the soil. In both forests, there was an exponential reduction of root biomass with increasing depth, and most of the variation in the vertical distribution of roots less than 2 mm in diameter was explained by the concentration of nitrogen in the soils. The results of this study support the hypothesis that a large root biomass in montane forests is related to nutrients in low concentration and diluted in organic soils with high CEC and low bulk density, and that fine root biomass in tropical forests in inversely related to calcium availability but not a phosphorus as has been suggested for other forests.     

贵州中部喀斯特山地不同植被生态系统细根生态特征及养分储量

为了解喀斯特生态系统退化过程中树木细根生物量和土壤养分的变化,选择贵州中部喀斯特山地乔木林、灌木林和灌草丛3种植被生态系统,比较分析不同深度(0～5 cm、5～10 cm和10～15 cm)土壤细根数量及其养分情况.结果表明:树木细根主要分布在0～10 cm土层,并随土层加深而减少.在0～10 cm土层中,乔木林、灌木林和灌草丛的活细根生物量分别占0～15 cm总细根生物量的42.78%、56.75%和53.38%,总活细根生物量的83.36%、86.91%和93.79%.不同植被下优势种植物细根生物量存在差异.0～5 cm土层乔木林活细根氮素和磷素储量均显著高于灌草丛和灌木林(P0.05),但灌木林和灌草丛间没有差异;5～10 cm土层乔木林活细根氮和磷储量显著高于灌草丛和灌木林(P0.05),灌木林下又显著高于灌草丛下(P0.05).0～10 cm土层的活细根生物量与植株地上部分生物量呈正相关,植物叶片氮、磷养分含量与细根比根长呈显著的负相关,说明细根的养分储量对地上生物量的建成和生态系统功能的发挥具有重要作用.     

The root to shoot ratio of trees from open- and closed-canopy cerrado in south-eastern Brazil

Background: The Brazilian savanna, or Cerrado, has been described as an ‘upside-down forest’, with higher below-ground than above-ground biomass. The cerrado vegetation, ranging from open grasslands to forests, comprises a wide range of ecological conditions and plant biomass.

Aims: To determine if and how root:shoot ratio in 102 trees differed between open- (cerrado sensu stricto) and closed-canopy cerrado (cerradão) within the same region in south-eastern Brazil.

Methods: Differences in root:shoot ratios and environmental conditions between the two cerrado types were examined, by uprooting and weighing trees from different species and functional groups.

Results: Root:shoot ratio was higher in the open than in the closed cerrado, especially among deciduous species. Root:shoot ratio in the open cerrado was lower than reported for the same cerrado type in central Brazil. Soil fertility did not differ between cerrado types, but soil water was lower and light availability was higher in the open cerrado.

Conclusions: The lower root:shoot ratio in closed than in open cerrado is probably a response to lower light and higher soil water availability, and/or to less frequent fires. Estimates of above-ground carbon storage alone significantly underestimate the carbon stock in open relative to closed cerrado.     

Soil freezing alters fine root dynamics in a northern hardwood forest

The retention of nutrients within an ecosystem depends on temporal andspatial synchrony between nutrient availability and nutrient uptake, anddisruption of fine root processes can have dramatic impacts on nutrientretention within forest ecosystems. There is increasing evidence thatoverwinter climate can influence biogeochemical cycling belowground,perhaps by disrupting this synchrony. In this study, we experimentallyreduced snow accumulation in northern hardwood forest plots to examinethe effects of soil freezing on the dynamics of fine roots (< 1 mm diameter)measured using minirhizotrons. Snow removal treatment during therelatively mild winters of 1997–1998 and 1998–1999 induced mild freezingtemperatures (to –4 °C) lasting approximately three months atshallow soil depths (to –30 cm) in sugar maple and yellow birch stands.This treatment resulted in elevated overwinter fine root mortality in treatedcompared to reference plots of both species, and led to an earlier peak infine root production during the subsequent growing season. These shiftsin fine root dynamics increased fine root turnover but were not largeenough to significantly alter fine root biomass. No differences inmorality response were found between species. Laboratory tests on pottedtree seedlings exposed to controlled freezing regimes confirmed that mildfreezing temperatures (to –5 °C) were insufficient to directlyinjure winter-hardened fine roots of these species, suggesting that themarked response recorded in our forest plots was caused indirectly bymechanical damage to roots in frozen soil. Elevated fine root necromass intreated plots decomposed quickly, and may have contributed an excess fluxof about 0.5 g N/m²·yr, which is substantial relative tomeasurements of N fluxes from these plots. Our results suggest elevatedoverwinter mortality temporarily reduced fine root length in treatmentplots and reduced plant uptake, thereby disrupting the temporalsynchrony between nutrient availability and uptake and enhancing ratesof nitrification. Increased frequency of soil freezing events, as may occurwith global change, could alter fine root dynamics within the northernhardwood forest disrupting the normally tight coupling between nutrientmineralization and uptake.     

Seasonal dynamics of fine root biomass, root length density, specific root length, and soil resource availability in a Larix gmelinii plantation

Fine root turnover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors. Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past, our understanding of it remains limited. This is because the dynamics processes associated with soil resources availability are still poorly understood. Soil moisture, temperature, and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level. In temperate forest ecosystems, seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground. Therefore, fine root biomass, root length density (RLD) and specific root length (SRL) vary during the growing season. Studying seasonal changes of fine root biomass, RLD, and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover. The objective of this study was to understand whether seasonal variations of fine root biomass, RLD and SRL were associated with soil resource availability, such as moisture, temperature, and nitrogen, and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation. We used a soil coring method to obtain fine root samples (⩽2 mm in diameter) every month from May to October in 2002 from a 17-year-old L. gmelinii plantation in Maoershan Experiment Station, Northeast Forestry University, China. Seventy-two soil cores (inside diameter 60 mm; depth intervals: 0–10 cm, 10–20 cm, 20–30 cm) were sampled randomly from three replicates 25 m × 30 m plots to estimate fine root biomass (live and dead), and calculate RLD and SRL. Soil moisture, temperature, and nitrogen (ammonia and nitrates) at three depth intervals were also analyzed in these plots. Results showed that the average standing fine root biomass (live and dead) was 189.1 g·m⁻²·a⁻¹, 50% (95.4 g·m⁻²·a⁻¹) in the surface soil layer (0–10 cm), 33% (61.5 g·m⁻²·a⁻¹), 17% (32.2 g·m⁻²·a⁻¹) in the middle (10–20 cm) and deep layer (20–30cm), respectively. Live and dead fine root biomass was the highest from May to July and in September, but lower in August and October. The live fine root biomass decreased and dead biomass increased during the growing season. Mean RLD (7,411.56 m·m⁻³·a⁻¹) and SRL (10.83 m·g⁻¹·a⁻¹) in the surface layer were higher than RLD (1 474.68 m·m⁻³·a⁻¹) and SRL (8.56 m·g⁻¹·a⁻¹) in the deep soil layer. RLD and SRL in May were the highest (10 621.45 m·m⁻³ and 14.83m·g⁻¹) compared with those in the other months, and RLD was the lowest in September (2 198.20 m·m⁻³) and SRL in October (3.77 m·g⁻¹). Seasonal dynamics of fine root biomass, RLD, and SRL showed a close relationship with changes in soil moisture, temperature, and nitrogen availability. To a lesser extent, the temperature could be determined by regression analysis. Fine roots in the upper soil layer have a function of absorbing moisture and nutrients, while the main function of deeper soil may be moisture uptake rather than nutrient acquisition. Therefore, carbon allocation to roots in the upper soil layer and deeper soil layer was different. Multiple regression analysis showed that variation in soil resource availability could explain 71–73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass. These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability, which resulted in an increased allocation of carbohydrate to these roots, but a lower allocation of carbohydrate to those in soil with lower resource availability. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(3): 403–410 [译自: 植物生态学报, 2005, 29(3): 403–410]     

三峡库区马尾松根系生物量的空间分布

以三峡库区主要植被马尾松人工林为研究对象,用内径为10 cm的根钻,分别在马尾松中龄林、近熟林和成熟林内,据树干0.5、1.0、1.5 m和2.0 m处设置取样点,各样点按0-10、10-20、20-30、30-40、40-60 cm将土壤分为5个垂直层次,对马尾松根系的空间分布格局进行调查。结果表明:(1)三峡库区马尾松总根系生物量(0-10 mm)为中龄林(4.72 t/hm²)显著高于成熟林(2.94 t/hm²)和近熟林(2.40 t/hm²)(P<0.05)。细根(0-2 mm)生物量随年龄增加而递减,差异不显著(P>0.05);(2)马尾松3个林龄中根系生物量表现出一定的水平分布特征,但具体趋势表现各异,细根生物量最大值均出现在距离样木1.0 m处;(3)细根主要分布在土壤上层,其中47.53%-71.73%的活细根集中在0-20 cm土壤深度内,且随土层的加深,其生物量明显减少。粗根(2-10 mm)则主要分布于20-60 cm土层范围内;(4)根系直径越小,受环境变化越明显。马尾松细根生物量分布主要受土壤深度的影响,树龄和不同水平距离对细根分布格局影响不显著(P>0.05),各因素对粗根生物量的影响均未达到显著水平(P>0.05)。     

长白山阔叶红松林表层土壤木本植物细根生物量及其空间分布

2008年在长白山北坡原始阔叶红松林内选择3块50 m×50 m样地,采用地统计学方法对表层土壤中木本植物细根生物量及其分布特征进行了定量研究.结果表明：3块样地0～20 cm土层中木本植物活细根生物量分别为3.195、1.930和2.058 t·hm^-2,死细根生物量分别为0.971、0.581和0.790 t·hm^-2,0～10 cm土层中,死、活细根生物量之间无显著相关关系,而10～20 cm土层中,二者呈显著正相关关系(r=0.352,P<0.05）,死、活细根生物量的实际变异函数大多符合球状理论模型.空间自相关引起的空间异质性占总空间异质性的百分比平均大于70%,各样地活、死细根生物量变程分别为5.2、14.6、9.8 m和4.3、20.4、20.1 m.采用贝叶斯统计方法对3块样地活细根生物量空间自相关范围进行估计的结果与地统计学方法的统计结果一致.     

岷江干旱河谷25种植物一年生植株根系功能性状及相互关系

相同条件下相同生长期的植物根系生长与适应策略及其差异性还不清楚。因此,采集岷江干旱河谷地区25种乡土植物(木本15/草本10种)的种子于2009年3月播种在同一干旱环境中,9月测定了1年生植株的最大根深(RDmax)、根幅(RW)与根生物量(RB),计算了总根长(TRL)、比根长(SRL)及细/粗根生物量比(RBf/c),分析了它们之间的关系,进行了根系功能组划分。结果表明:1)25种植物1年生植株RDmax与RW变异较小,总变异率为14.9%和20.7%;TRL和SRL变异相对较大,分别为28.5%和34.7%,草本植物SRL明显大于木本植物;RB和RBf/c种间变异较大,总变异率分别为50.1%和70.5%;2)25种植物的RDmax、RW、RB和TRL间呈显著正相关关系,表明根系较深的物种RW较大,TRL和RB也较高;SRL与RDmax呈极显著负相关关系,与RBf/c呈极显著正相关关系,表明根系垂直分布较浅的物种细根发达,SRL较大;3)主成分分析显示,25种植物可分为3个功能组:第1组具有较大RDmax、RW和RB,资源利用持续时间较长;第2组具有较大TRL、SRL和RBf/c,资源利用效率较高;第3组根系功能性状没有一致的突出特点,可能通过降低自身生理机能适应生存条件。综合分析表明,岷江干旱河谷区25种植物1年生植株根系的功能性状变异明显,可塑性大,历经长期自然选择压力而形成了不同的环境适应策略,但生长型并不必然表达出1年生植株根系功能性状的差异性。     

Compensation among root classes in Phaseolus vulgaris L.

The activity of soil pathogens, competition for assimilates, and the changing availability of below-ground resources make root systems subject to a continuous and dynamic process of formation and loss of both fine and coarse roots. As hypocotyl borne roots appear later than other root classes, they may serve to functionally replace basal and primary roots lost to biotic and abiotic stress. Using common bean (Phaseolus vulgaris L.), we conducted experiments in solution and solid media culture with treatments involving the removal of part of the root system (basal, hypocotyl borne or primary roots), phosphorus availability, and depth of seeding to test the hypothesis that there are compensation mechanisms among basal, hypocotyl borne and primary roots to cope with the loss of part of the root system. The root system was highly plastic in response to root excision, which resulted in the maintenance of below-ground biomass accumulation. In most cases, this compensation among root classes was enough to maintain plant performance in both phosphorus sufficient and phosphorus stressed plants. Removal of a specific root class induced an increase in the growth of the remaining root classes. All root classes, but especially the primary root, contributed to the compensation mechanism in some way. Primary roots represented around 10% of the root system in control plants and this proportion increased dramatically (up to 50%) when other root classes were removed. In contrast, negligible compensatory re-growth was observed following removal of the primary root. Greater planting depth increased the production of hypocotyl borne roots at the expense of basal roots. The proportion of hypocotyl borne roots increased from 25% of the whole root system when seeds were placed at a depth of 2 cm to 30% when they were placed at 5 cm and to 38% when placed at 8 cm, with corresponding decreases in the proportion represented by basal roots. The common feature of our observations is the innate ability of the root system for its own regeneration. Total root biomass maintained strict allometric relationships with total shoot biomass in all treatments. Re-stabilization of root to shoot balance after partial root loss is governed by overall plant size following allometric relationships similar to undisturbed plants. However, the pattern of this root regeneration was not uniform since the way the three root classes compensated each other after the removal of any one of them varied among the different growth media and phosphorus supply conditions. The resulting changes in root architecture could have functional significance for soil resource acquisition.     

Interactions between repeated fire, nutrients, and insect herbivores affect the recovery of diversity in the southern Amazon

Surface fires burn extensive areas of tropical forests each year, altering resource availability, biotic interactions, and, ultimately, plant diversity. In transitional forest between the Brazilian cerrado (savanna) and high stature Amazon forest, we took advantage of a long-term fire experiment to establish a factorial study of the interactions between fire, nutrient availability, and herbivory on early plant regeneration. Overall, five annual burns reduced the number and diversity of regenerating stems. Community composition changed substantially after repeated fires, and species common in the cerrado became more abundant. The number of recruits and their diversity were reduced in the burned area, but burned plots closed to herbivores with nitrogen additions had a 14 % increase in recruitment. Diversity of recruits also increased up to 50 % in burned plots when nitrogen was added. Phosphorus additions were related to an increase in species evenness in burned plots open to herbivores. Herbivory reduced seedling survival overall and increased diversity in burned plots when nutrients were added. This last result supports our hypothesis that positive relationships between herbivore presence and diversity would be strongest in treatments that favor herbivory—in this case herbivory was higher in burned plots which were initially lower in diversity. Regenerating seedlings in less diverse plots were likely more apparent to herbivores, enabling increased herbivory and a stronger signal of negative density dependence. In contrast, herbivores generally decreased diversity in more species rich unburned plots. Although this study documents complex interactions between repeated burns, nutrients, and herbivory, it is clear that fire initiates a shift in the factors that are most important in determining the diversity and number of recruits. This change may have long-lasting effects as the forest progresses through succession.     

模拟氮沉降对中亚热带杉木幼树根系生物量的影响

植物根系是全球陆地生态系统碳储量的重要组成部分,在全球生态系统碳循环中起着重要作用,日益加剧的氮沉降会影响根系生物量在空间和不同径级的分配,进而影响森林生态系统的生产力和土壤养分循环。以杉木幼树为研究对象,通过野外氮沉降模拟实验,研究氮沉降四年后对不同土层、不同径级根系生物量的影响。结果发现：（1）低氮和高氮处理总细根生物量较对照均无显著差异（P > 0.05）,高氮处理粗根生物量及总根系生物量较对照分别增加45%和40%（P < 0.05）;（2）与对照相比,施氮处理显著增加20-40 cm与40-60 cm土层细根和粗根生物量,且在低氮处理下,20-40 cm土层细根、粗根在总土层细根与粗根生物量的占比显著提高。（3）与对照相比,高氮处理显著增加了2-5 mm、5-10 mm及10-20 mm径级的根系生物量,低氮处理显著增加2-5 mm、5-10 mm径级根系生物量,且显著降低20-50 mm径级根系生物量。综上所述表明：氮沉降后杉木幼树通过增加较粗径级根系来增加对养分及水分的输送,同时通过增加深层根系生物量及其比例的策略来维持杉木幼树的快速生长;而根系生物量的增加,在一定程度上会增加根系碳源的输入,影响土壤碳循环过程。     

Positive tree diversity effect on fine root biomass: via density dependence rather than spatial root partitioning

The importance of species richness to ecosystem functioning and services is a central tenet of biological conservation. However, most of our theory and mechanistic understanding is based on diversity found aboveground. Our study sought to better understand the relationship between diversity and belowground function by studying root biomass across a plant diversity gradient. We collected soil cores from 91 plots with between 1 and 12 aboveground tree species in three natural secondary forests to measure fine root (≤ 2 mm in diameter) biomass. Molecular methods were used to identify the tree species of fine roots and to estimate fine root biomass for each species. This study tested whether the spatial root partitioning (species differ by belowground territory) and symmetric growth (the capacity to colonize nutrient-rich hotspots) underpin the relationship between aboveground species richness and fine root biomass. All species preferred to grow in nutrient-rich areas and symmetric growth could explain the positive relationship between aboveground species richness and fine root biomass. However, symmetric growth only appeared in the nutrient-rich upper soil layer (0–10 cm). Structural equation modelling indicated that aboveground species richness and stand density significantly affected fine root biomass. Specifically, fine root biomass depended on the interaction between aboveground species richness and stand density, with fine root biomass increasing with species richness at lower stand density, but not at higher stand density. Overall, evidence for spatial (i.e. vertical) root partitioning was inconsistent; assumingly any roots growing into deeper unexplored soil layers were not sufficient contributors to the positive diversity–function relationship. Alternatively, density-dependent biotic interactions affecting tree recruitment are an important driver affecting productivity in diverse subtropical forests but the usual root distribution patterns in line with the spatial root partitioning hypothesis are unrealistic in contexts where soil nutrients are heterogeneously distributed.     

Does soil acidity reduce subsoil rooting in Norway spruce (Picea abies)?

Increasing evidence suggests that forest soils in central and northern Europe as well as in North America have been significantly acidified by acid deposition during the last decades. The present investigation was undertaken to examine the effect of soil acidity on rooting patterns of 40-year-old Norway spruce trees by comparing fine and coarse roots among four stands which differed in soil acidity and Mg (and Ca) nutrition. The coarse root systems of four to five 40-year-old Norway spruce trees per stand were manually excavated. The sum of cross sectional area (CSA) at 60 cm soil depth and below of all vertical coarse roots, as a measure of vertical rooting intensity, was strongly reduced with increasing subsoil acidity of the stands. This pattern was confirmed when 5 additional acidic sites were included in the analysis. Fine root biomass in the mineral soil estimated by repeated soil coring was strongly reduced in the heavily acidified stands, but increased in the humic layer. Using ingrowth cores and a screen technique, we showed that the higher root biomass in the humic layer of the more acidic stands was a result of higher root production. Thus, reduced fine root biomass and coarse root CSA in deeper soil layers coincided with increased root growth in the humic layer. Root mineral analysis showed Ca/Al ratios decreased with decreasing base saturation in the deeper mineral soil (20–40 cm). In the top mineral soil, only minor differences were observed among stands. In general, low Ca/Al ratios coincided with low fine root biomass. Calcium/aluminum ratios determined in cortical cell walls using X-ray microanalysis showed a similar pattern as Ca/Al ratios based on analysis of whole fine roots, although the amplitude of changes among the stands was much greater. Aluminum concentrations and Ca/Al ratios in cortical cell walls were at levels found to inhibit root growth of spruce seedlings in laboratory experiments. The data support the idea that Al (or Ca/Al ratios) and acid deposition-induced Mg (and possibly Ca) deficiency are important factors influencing root growth and distribution in acidic forest soils. Changes in carbon partitioning within the root system may contribute to a reduction in deep root growth.     

Biomass and distribution of fine and coarse roots from blue oak (Quercus douglasii) trees in the northern Sierra Nevada foothills of California

This research adds to the limited data on coarse and fine root biomass for blue oak (Quercus douglasii Hook and Arn.), a California deciduous oak species found extensively throughout the interior foothills surrounding the Central Valley. Root systems of six blue oak trees were analyzed using three methods — backhoe excavation, quantitative pits, and soil cores. Coarse root biomass ranged from 7 to 177 kg per tree. Rooting depth for the main root system ranged from 0.5 to 1.5 m, with an average of 70% of excavated root biomass located above 0.5 m. Of the total biomass in excavated central root systems, primary roots (including burls) accounted for 56% and large lateral roots (> 20 mm diameter) accounted for 36%. Data from cores indicated that most biomass outside of the root crown was located in fine roots and that fine root biomass decreased with depth. At surface depths (0–20 cm), small-fine (< 0.5 mm diameter) roots accounted for 71%, large-fine (0.5–2.0 mm) for 25%, and coarse (> 2 mm) for 4% of total root biomass collected with cores. Mean fine root biomass density in the top 50 cm was 0.43 kg m⁻³. Fine root biomass did not change with increasing distance from the trees (up to approximately 5 m). Thus, fine roots were not concentrated under the tree canopies. Our results emphasize the importance of the smallest size class of roots (<0.5 mm), which had both higher N concentration and, in the area outside the central root system, greater biomass than large fine (0.5–2.0 mm) or coarse (> 2.0 mm) roots. This revised version was published online in June 2006 with corrections to the Cover Date.     

青海省森林细根生物量及其影响因子

细根是植物吸收水分和养分的主要器官。全球变暖背景下,研究森林细根生物量及其环境因子的变化对生态系统碳平衡、碳收支及其贡献率具有重要意义。采用土钻法和室内分析法对青海省森林6个海拔梯度上5种林分类型的细根生物量和土壤理化性质进行测定,并分析了与环境因子之间的相互关系。结果表明:(1)青海省森林0—40 cm土层总细根生物量平均为8.50 t/hm~2,随着海拔梯度的增加先降低后升高,不同海拔梯度细根生物量差异显著(P0.05),最大值出现在2100—2400 m处。(2)5种林分0—40 cm土层总细根生物量为:白桦白杨云杉圆柏山杨,不同林分间细根生物量差异不显著。(3)细根垂直分布随土层深度增加而减少,且70%的细根集中在表层(0—20 cm)。(4)土壤容重深层(20—40 cm)显著大于表层(P0.05),并随海拔梯度逐步增加,且林分间差异较大。(5)全碳(Total carbon, TC)、全氮(Total nitrogen, TN)、全磷(Total phosphorus, TP)含量表层显著高于深层。TC、TN随海拔升高先增后降低,TP则随海拔逐步降低。不同林分间土壤养分差异较明显。(6)结构方程模型分析得到海拔、土层、容重直接影响细根生物量,细根生物量直接影响土壤养分。林分类型通过土壤容重间接影响细根生物量。因此,林分和海拔通过影响土壤微环境而影响到细根生物量及其空间分布格局。     

中亚热带森林植物多样性增加导致细根生物量“超产”

细根在森林生态系统C分配和养分循环过程中发挥着重要作用, 但对地下细根与植物多样性之间关系的研究相对较少。该研究选择中亚热带从单一树种的杉木(Cunninghamia lanceolata)人工林到多树种的常绿阔叶林(青冈(Cyclobalanopsis glauca)-石栎(Lithocarpus glaber)林)的不同植物多样性梯度, 用根钻法采集细根并测定其生物量, 用Win-RHIZO 2005C根系分析系统测定细根形态参数, 以验证以下3个假设: 1)植物种类丰富度高的林分其细根生产存在“地下超产”现象; 2)根系空间生态位的分离水平是否随着植物多样性增多而增大? 3)细根是否通过形态可塑性对林木竞争做出响应?结果显示: 从单一树种的杉木人工林到植物种类较复杂的青冈-石栎常绿阔叶林, 0-30 cm土层的林分细根总生物量和活细根生物量均呈增加的趋势, 即细根总生物量为杉木林(305.20 g·m^-2) <马尾松(Pinus massoniana)林(374.25 g·m^-2) <南酸枣(Choerospondias axillaris)林(537.42 g·m^-2) <青冈林(579.33 g·m^-2), 活细根生物量为杉木林(268.74 g·m^-2) <马尾松林(299.15 g·m^-2) <南酸枣林(457.32 g·m^-2) <青冈林(508.47 g·m^-2), 各森林类型之间的细根总生物量差异显著(p < 0.05), 但活细根生物量差异不显著。土壤垂直剖面上, 除杉木林细根生物量随土层变化不显著外, 其他森林类型的活细根生物量和总细根生物量均随土层变化显著, 表层细根生物量随树种多样性的升高呈减小趋势, 据此推测树种间的生态位分离水平逐渐增大。植物多样性的不同对林分的细根形态及空间分布格局影响不显著, 细根形态可塑性对生物量变化响应不明显。     

Phylogeny, traits, environment, and space in cerrado plant communities at Emas National Park (Brazil)

Soil, drought, and fire are abiotic factors that may act as environmental filters in the cerrado, the Brazilian savanna. We used a framework to analyze environmental filtering in geographic and phylogenetic context, sampling woody species in one of the largest cerrado reserves. In 100 quadrats, we measured 10 functional traits on each woody individual. We also measured several soil variables, altitude and slope as a rough surrogate of water availability, interval between fires, and time since last fire. Almost all environmental variables were spatially auto-correlated. We found an overall trait clustering, but not an overall phylogenetic clustering. Nevertheless, we found a phylogenetic signal for some traits. Linking phylogeny, traits, environment, and space, we were able to detect a major dichotomy between two geomorphological units. The flat tableland was positively related with altitude, fire frequency, and nutrient-richer soil. Environmental filtering caused by water availability and fire lead to trait clustering, with smaller shrubs and trees that presented thicker barks, denser woods, sclerophyllous leaves, highlighted by the prevalance of Myrtaceae. The other geomorphological unit, hilly terrain, was positively related with slope, low fire frequency, and nutrient-poorer soil. Environmental filtering was caused especially by nutrient-poor soil that lead to trait clustering, assembling taller trees, with thinner barks, lighter woods, and compound, large, tender, nutrient-richer leaves, distributed across many lineages, including Fabaceae. Hence, the high environmental variability in space with different environmental filters assembled different combination of plant traits and lineages, increasing the overall diversity in cerrado.