Measuring Fine Root Turnover in Forest Ecosystems

Development of direct and indirect methods for measuring root turnover and the status of knowledge on fine root turnover in forest ecosystems are discussed. While soil and ingrowth cores give estimates of standing root biomass and relative growth, respectively, minirhizotrons provide estimates of median root longevity (turnover time) i.e., the time by which 50% of the roots are dead. Advanced minirhizotron and carbon tracer studies combined with demographic statistical methods and new models hold the promise of improving our fundamental understanding of the factors controlling root turnover. Using minirhizotron data, fine root turnover (y⁻¹) can be estimated in two ways: as the ratio of annual root length production to average live root length observed and as the inverse of median root longevity. Fine root production and mortality can be estimated by combining data from minirhizotrons and soil cores, provided that these data are based on roots of the same diameter class (e.g., < 1 mm in diameter) and changes in the same time steps. Fluxes of carbon and nutrients via fine root mortality can then be estimated by multiplying the amount of carbon and nutrients in fine root biomass by fine root turnover. It is suggested that the minirhizotron method is suitable for estimating median fine root longevity. In comparison to the minirhizotron method, the radio carbon technique favor larger fine roots that are less dynamics. We need to reconcile and improve both methods to develop a more complete understanding of root turnover.     

Global patterns of root turnover for terrestrial ecosystems

Root turnover is a critical component of ecosystem nutrient dynamics and carbon sequestration and is also an important sink for plant primary productivity. We tested global controls on root turnover across climatic gradients and for plant functional groups by using a database of 190 published studies. Root turnover rates increased exponentially with mean annual temperature for fine roots of grasslands ( r ² = 0.48) and forests ( r ² = 0.17) and for total root biomass in shrublands ( r ² = 0.55). On the basis of the best-fit exponential model, the Q ₁₀ for root turnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. Surprisingly, after accounting for temperature, there was no such global relationship between precipitation and root turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), followed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for forest fine roots. Root turnover decreased from tropical to high-latitude systems for all plant functional groups. To test whether global relationships can be used to predict interannual variability in root turnover, we evaluated 14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site there was no correlation between interannual variability in mean annual temperature and root turnover. Rather, turnover was positively correlated with the ratio of growing season precipitation and maximum monthly temperature ( r ² = 0.61). We conclude that there are global patterns in rates of root turnover between plant groups and across climatic gradients but that these patterns cannot always be used for the successful prediction of the relationship of root turnover to climate change at a particular site.     

Estimating Root Production: Comparison of 11 Methods in Shortgrass Steppe and Review of Biases

Estimating root production has been difficult due to multiple potential biases associated with both old and new methods. This shortgrass steppe site is the only place we are aware of that can compare most methods including sequential coring, ingrowth cores, and ingrowth donuts, ¹⁴C pulse-isotope dilution, ¹⁴C pulse-isotope turnover, rhizotron windows, and minirhizotron, and indirect methods including nitrogen budget, carbon flux, simulation carbon flow model, and regression model. We used the studies at this site, other comparisons, a summary of potential directional biases, and different ways of calculating estimates in a logical, comparative approach of evaluating methods. Much of the literature for root production is based on sequential biomass coring, a method resulting in erroneous estimates. Root ingrowth estimates of production are generally conservative compared to minirhizotron and isotope turnover methods. The size of the ingrowth area may be the most important determinant of the underestimation. Estimates based on pulse-isotope dilution are also erroneous due to non-uniform labeling of tissues. Uniform labeling is not an assumption of the pulse-isotope turnover method, and this method has the least severe potential biases. Root production estimates from pulse-isotope turnover were lower than those using minirhizotron when the most common method of calculation was used. This agrees with literature concerning bomb ¹⁴C continuous-isotope labeling comparisons with minirhizotron, although some potential biases between isotope methods are different. However, good agreement between pulse-isotope turnover and minirhizotron were obtained when minirhizotron estimates were calculated from regression of decomposition versus production to equilibrium and when pulse-isotope turnover estimates were calculated from two-phase life-span regressions. This minirhizotron method bypasses biases associated with the artificial surface similar to root-cohort methods that may be practical only in mesic systems, and takes into account both short- and long-lived roots and corrects for soil-isotope contamination that the continuous-isotope labeling bomb ¹⁴C method is not able to account for. Comparisons of these direct methods are also made with four indirect methods.     

Allocation of carbon to fine root compounds and their residence times in a boreal forest depend on root size class and season

Fine roots play a key role in the forest carbon balance, but their carbon dynamics remain largely unknown. We pulse labelled 50 m(2) patches of young boreal forest by exposure to (13)CO(2) in early and late summer. Labelled photosynthates were traced into carbon compounds of < 1 and 1-3 mm diameter roots (fine roots), and into bulk tissue of these and first-order roots (root tips). Root tips were the most strongly labelled size class. Carbon allocation to all size classes was higher in late than in early summer; mean residence times (MRTs) in starch increased from 4 to 11 months. In structural compounds, MRTs were 0.8 yr in tips and 1.8 yr in fine roots. The MRT of carbon in sugars was in the range of days. Functional differences within the fine root population were indicated by carbon allocation patterns and residence times. Pronounced allocation of recent carbon and higher turnover rates in tips are associated with their role in nutrient and water acquisition. In fine roots, longer MRTs but high allocation to sugars and starch reflect their role in structural support and storage. Accounting for heterogeneity in carbon residence times will improve and most probably reduce the estimates of fine root production.     

帽儿山温带落叶阔叶林细根生物量、生产力和周转率

细根在森林生态系统能量流动与物质循环中占有重要地位,但其生物量、生产和周转测定尚存在很大的不确定性,而且局域尺度空间变异机制尚不清楚。本研究分析了帽儿山温带天然次生林活细根生物量和死细根生物量在0～100 cm剖面的垂直分布与0～20 cm细根的季节动态、生产力和周转率,对比了采用连续根钻法(包括决策矩阵法和极差法)和内生长袋(直径3和5 cm)估测细根生产力和细根周转率,并探讨了可能影响细根的林分因子。结果表明: 76.8%的活细根生物量和62.9%的死细根生物量均集中在0～20 cm土层,随着深度增加,二者均呈指数形式减少。活细根生物量和死细根生物量的季节变化不显著,可能与冬季几乎无降雪而夏季降雨异常多有关。2种直径内生长袋估计的细根生产力无显著差异;对数转换后决策矩阵、极差法和内生长法估计的细根生产力和细根周转率差异显著。随着土壤养分增加,活细根生物量和死细根生物量比值显著增加,死细根生物量显著减少,但活细根生物量、细根生产力和细根周转率均无显著变化;细根周转率与前一年地上木质生物量增长量呈显著正相关,但与当年地上木质生物量增长量无显著相关关系。     

树木根系碳分配格局及其影响因子

根系作为树木提供养分和水分的“源”和消耗C的“汇”,在陆地生态系统C平衡研究中具有重要的理论意义。尽管20多年来的研究已经认识到根系消耗净初级生产力占总净初级生产力较大的比例,但是,根系（尤其是细根）消耗C的机理以及C分配的去向一直没有研究清楚。主要原因是细根消耗光合产物的生理生态过程相当复杂,准确估计各个组分消耗的C具有很大的不确定性,常常受树种和环境空间和时间异质性、以及研究方法的限制。综述了分配到地下的C主要去向,即细根生产和周转、呼吸及养分吸收与同化、分泌有机物、土壤植食动物,及有关林木地下碳分配机理的几种假说,分析了地下碳分配估计中存在的不确定性。目的是在全球变化C循环研究中对生态系统地下部分根系消耗的C以及分配格局引起重视。     

Regional scale patterns of fine root lifespan and turnover under current and future climate

Fine root dynamics control a dominant flux of carbon from plants and into soils and mediate potential uptake and cycling of nutrients and water in terrestrial ecosystems. Understanding of these patterns is needed to accurately describe critical processes like productivity and carbon storage from ecosystem to global scales. However, limited observations of root dynamics make it difficult to define and predict patterns of root dynamics across broad spatial scales. Here, we combine species‐specific estimates of fine root dynamics with a model that predicts current distribution and future suitable habitat of temperate tree species across the eastern United States (US). Estimates of fine root lifespan and turnover are based on empirical observations and relationships with fine root and whole‐plant traits and apply explicitly to the fine root pool that is relatively short‐lived and most active in nutrient and water uptake. Results from the combined model identified patterns of faster root turnover rates in the North Central US and slower turnover rates in the Southeastern US. Portions of Minnesota, Ohio, and Pennsylvania were also predicted to experience >10% increases in root turnover rates given potential shifts in tree species composition under future climate scenarios while root turnover rates in other portions of the eastern US were predicted to decrease. Despite potential regional changes, the average estimates of root lifespan and turnover for the entire study area remained relatively stable between the current and future climate scenarios. Our combined model provides the first empirically based, spatially explicit, and spatially extensive estimates of fine root lifespan and turnover and is a potentially powerful tool allowing researchers to identify reasonable approximations of forest fine root turnover in areas where no direct observations are available. Future efforts should focus on reducing uncertainty in estimates of root dynamics by better understanding how climate and soil factors drive variability in root dynamics of different species.     

Fine root biomass and turnover in southern taiga estimated by root inclusion nets

Fine roots play an important part in forest carbon, nutrient and water cycles. The turnover of fine roots constitutes a major carbon input to soils. Estimation of fine root turnover is difficult, labour intensive and is often compounded by artefacts created by soil disturbance. In this work, an alternative approach of using inclusion nets installed in an undisturbed soil profile was used to measure fine root production and was compared to the in-growth core method. There was no difference between fine root production estimated by the two methods in three southern taiga sites with contrasting soil conditions and tree species compositions in the Central Forest State Biosphere Reserve, Russia. Expressed as annual production over standing biomass, Norway spruce fine root turnover was in the region of 0.10 to 0.24 y^-1. The inclusion net technique is suitable for field based assessment of fine root production. There are several advantages over the in-growth core method, due to non-disturbance of the soil profile and its potential for very high rate of replication.     

水曲柳和落叶松不同根序之间细根直径的变异研究

细根直径大小和根序高低对细根寿命和周转估计具有重要的影响,研究不同根序之间的直径变异对认识细根直径与根序的关系具有重要意义。该文根据Pregitzer等(2002)提供的方法,研究了位于东北林业大学帽儿山实验林场尖砬沟森林培育实验站17年生水曲柳(Fraxinus mandshurica)和落叶松(Larix gmelinii)人工林细根1~5级根序的平均直径的变化、直径的最小值和最大值范围、直径的变异系数。结果表明,水曲柳和落叶松细根直径<2 mm时,包含5个根序,随着根序由小到大的增加,细根直径也在增大。各根序平均直径之间,存在较大的差异。在同一根序内,细根直径范围很大,水曲柳和落叶松一级根最小直径均<0.20 mm,最大直径分别<0.50 mm(水曲柳)和<0.70 mm(落叶松)左右。2~3级根序直径最小值在0.20~0.30 mm之间,最大值≤1.0 mm。5级根直径最小值<1.0 mm,最大值超过2.0 mm。随着根序等级增加,直径变异系数增大。一级根序的直径平均变异系数<10%,2~3级根序直径平均变异系数在10%~15%左右,4~5级根序直径的平均变异系数在20%~30%之间。因此,在细根寿命与周转研究过程中,必须同时考虑直径和根序对细根的寿命估计的影响。     

Biofumigation potential of brassicas

The relationship of global climate change to plant growth and the role of forests as sites of carbon sequestration have encouraged the refinement of the estimates of root biomass and production. However, tremendous controversy exists in the literature as to which is the best method to determine fine root biomass and production. This lack of consensus makes it difficult for researchers to determine which methods are most appropriate for their system. The sequential root coring method was the most commonly used method to collect root biomass data in the past and is still commonly used. But within the last decade the use of minirhizotrons has become a favorite method of many researchers. In addition, due to the high labor-intensive requirements of many of the direct approaches to determine root biomass, there has been a shift to develop indirect methods that would allow fine root biomass and production to be predicted using data on easily monitored variables that are highly correlated to root dynamics. Discussions occur as to which method should be used but without gathering data from the same site using different methods, these discussions can be futile. This paper discusses and compares the results of the most commonly used direct and indirect methods of determining root biomass and production: sequential root coring, ingrowth cores, minirhizotrons, carbon fluxes approach, nitrogen budget approach and correlations with abiotic resources. No consistent relationships were apparent when comparing several sites where at least one of the indirect and direct methods were used on the same site. Until the different root methods can be compared to some independently derived root biomass value obtained from total carbon budgets for systems, one root method cannot be stated to be the best and the method of choice will be determined from researcher's personal preference, experiences, equipment, and/or finances.     

Fine root heterogeneity by branch order: exploring the discrepancy in root turnover estimates between minirhizotron and carbon isotopic methods

Fine roots constitute a large and dynamic component of the carbon cycles of terrestrial ecosystems. The reported fivefold discrepancy in turnover estimates between median longevity (ML) from minirhizotrons and mean residence time (MRT) using carbon isotopes may have global consequences. Here, a root branch order-based model and a simulated factorial experiment were used to examine four sources of error. Inherent differences between ML, a number-based measure, and MRT, a mass-based measure, and the inability of the MRT method to account for multiple replacements of rapidly cycling roots were the two sources of error that contributed more to the disparity than did the improper choice of root age distribution models and sampling bias. Sensitivity analysis showed that the rate at which root longevity increases as order increases was the most important factor influencing the disparity between ML and MRT. Assessing root populations for each branch order may substantially reduce the errors in longevity estimates of the fine root guild. Our results point to the need to acquire longevity estimates of different orders, particularly those of higher orders.     

三峡库区马尾松细根生产和周转及其影响因子

采用连续根钻法、分解袋法、分室通量模型法计算三峡库区马尾松细根的年生产量和周转率,分析细根生产量和周转率与各影响因子的关系.结果表明: 马尾松<0.5、0.5～1和1～2 mm细根年均生物量分别为0.29、0.59、0.76 t·hm^-2,细根年生产量分别为0.13、0.49、0.37 t·hm^-2,细根年周转率分别为1.49、1.01、0.40 a^-1.各影响因子对不同径级细根生产与周转的影响不同.土壤温度、土壤钙含量显著影响<0.5 mm细根生产量与细根周转,且土壤温度解释生产量和周转率32.8%和25.0%的变异,土壤钙含量解释65.6%和73.1%的变异;细根生物量与细根生产量呈显著正相关,细根生物量分别解释<0.5、0.5～1和1～2 mm细根生产量41.0%、41.1%和54.5%的变异;细根P、K含量与<0.5 mm细根生产量具有显著相关性,分别解释<0.5 mm细根生产量32.2%、39.2%的变异.<0.5 mm细根与各影响因子的关系最为密切,土壤温度、土壤钙含量是细根生物量的主要影响因子.     

Variety specific relationships between effects of rhizobacteria on root exudation,growth and nutrient uptake of soybean

Aims

It has been increasingly recognized that only distal lower order roots turn over actively within the <2 mm fine root system of trees. This study aimed to estimate fine root production and turnover rate based on lower order fine roots and their relations to soil variables in mangroves.

Methods

We conducted sequential coring in five natural mangrove forests at Dongzhai Bay, China. Annual fine root production and turnover rate were calculated based on the seasonal variations of the biomass and necromass of lower order roots or the whole fine root system.

Results

Annual fine root production and turnover rate ranged between 571 and 2838 g m^?2 and 1.46–5.96 yr^?1, respectively, estimated with lower order roots, and they were increased by 0–30 % and reduced by 13–48 %, respectively, estimated with the whole fine root system. Annual fine root production was 1–3.5 times higher than aboveground litter production and was positively related to soil carbon, nitrogen and phosphorus concentrations. Fine root turnover rate was negatively related to soil salinity.

Conclusions

Mangrove fine root turnover plays a more important role than aboveground litter production in soil C accumulation. Sites with higher soil nutrients and lower salinity favor fine root production and turnover, and thus favor soil C accumulation.

     

Impacts of plant roots on soil CO cycling and soil–atmosphere CO exchange

Carbon monoxide (CO) plays a major role in tropospheric chemical dynamics. Accordingly, global CO budgets have been reasonably well documented. Atmospheric CO consumption by soils contributes significantly to these budgets, with the magnitude of the sink generally considered to reflect a balance between microbial uptake and abiological production. However, assays of live fine roots showed that diverse intact plants produced carbon monoxide at net rates ranging from 2 to 3000 mµg gdw⁻⁻¹ d⁻⁻¹. CO production was greater for legumes than nonlegumes, and primarily associated with nodules. Excised roots from woody and herbaceous plants produced CO at comparable rates. CO production rates were similar for roots of intact plants and roots excised from those plants. The magnitude of net CO fluxes from roots was determined in part by the balance between simultaneous production and consumption. Surface sterilization of roots indicated that CO consumption was due, in part, rhizoplane CO-oxidizing bacteria, but maximum CO consumption rates were typically only a small fraction of net production rates. Assays in a Brazilian agroecosystem indicated that root CO production affects soil–atmosphere CO exchange. Estimates of global CO production rates indicated that roots contribute about 170–260 Tg CO to the soil atmosphere annually, an amount comparable to current estimates of atmospheric CO uptake by soils, and much larger than estimates of net abiological soil CO production.     

Seasonal patterns of fine root demography in a cool-temperate deciduous forest in central Japan

In forest ecosystems, fine roots have a considerable role in carbon cycling. To investigate the seasonal pattern of fine root demography, we observed the fine root production and decomposition processes using a minirhizotron system in a Betula-dominated forest with understory evergreen dwarf bamboo. The length density of fine roots decreased with increasing soil depth. The seasonal patterns of each fine root demographic parameter (length density of visible roots, rates of stand-total fine root production and decomposition) were almost the same at different soil depths. The peak seasons of the fine root demographic parameters were observed in the order: stand-total fine root production rate (late summer) > length density of the visible roots (early autumn) > stand-total fine root decomposition rate (autumn, and a second small peak in spring). The fine root production rate was high in the latter part of the plant growing season. Fine root production peaked in late summer and remained high until the end of the tree defoliation season. The higher stand-total fine root production rate in autumn suggests the effect of understory evergreen bamboo on the stand-total fine root demography. The stand-total fine root decomposition rate was high in late autumn. In the snow-cover period, the rates of both fine root production and decomposition were low. The fine root demographic parameters appeared to show seasonal patterns. The fine root production rate had a clearer seasonality than the fine root decomposition rate. The seasonal pattern of stand-total fine root production rate could be explained by both overstory and understory above-ground productivities.     

高寒草甸植被细根生产和周转的比较研究

植物根系是陆地生态系统重要的碳汇和养分库,细根周转过程是陆地生态系统地下部分碳氮循环的核心环节,在陆地生态系统如何响应全球变化中起着关键作用。在全球变化敏感地区之一的青藏高原,对该地区的主要植被类型矮嵩草草甸同时采用根钻法、内生长袋法和微根管法3种观测方法研究细根生产和周转速率,并探讨了极差法、积分法、矩阵法和Kaplan-Meier法等数据处理方法对计算值的影响。研究结果显示:在估算细根净初级生产力时,根钻法宜采用积分法,内生长袋法宜选用矩阵法;由此进一步以最大细根生物量为基础,根钻法和内生长袋法估测的细根年周转速率分别为0.36 a-1和0.52 a-1,内生长袋法的估算结果是根钻法的1.44倍。对于微根管法,将其观测得到的细根长度转换为单位面积的生物量值后,采用积分法计算出细根周转速率为0.84 a-1,远高于传统方法的估算结果;若采用Kaplan-Meier生存分析方法,则计算出的细根周转速率更高达3.41 a-1。     

应用微根管法测定细根指标方法评述

树木细根(直径<2mm)在森林生态系统能量流动和物质循环中起着重要的作用。原有的细根生产周转研究中常采用的土钻法、内生长法、挖掘法、根室法和土柱法等,均不能直接观察到细根的动态变化。微根管法是一种非破坏性、可定点直接观察和研究植物根系的方法,为研究细根的生长、衰老、死亡、分解和再生长的过程提供了有效的工具,尤其适用于细根周转、寿命和分解等方面的研究。但该技术不能直接测定单位面积的细根生物量、细根化学组成及细根周转对土壤碳和养分循环的影响,需要与土钻法结合。本文就运用微根管法对细根生物量、生产、周转和寿命等指标的研究方法进行了评述。     

氮添加对温带森林细根长期分解的影响

细根分解是森林生态系统碳循环的重要过程之一,其分解速率受到大气氮沉降增加的潜在影响。利用长期模拟氮沉降样地（2009年至今）,采用凋落物分解袋方法,研究了氮添加对温带常见的5个森林树种长期细根分解的影响。结果表明：细根分解呈现先快后慢的趋势,在分解第516天质量损失达30%~50%,之后质量残留率变化较为平缓。总体上,渐近线分解模型可以更准确的反应各处理细根分解速率。氮添加对细根分解具有阶段性影响,分解前期促进细根分解,分解后期抑制分解。在细根分解后期氮添加减缓分解速率,一方面是因为木质素等较难分解的物质所占比例升高所带来的直接影响,另一方面,是因为氮添加改变了微生物活动所带来的间接影响。     

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.     

Estimating Fine Root Turnover in Tropical Forests along an Elevational Transect using Minirhizotrons

Growth and death of fine roots represent an important carbon sink in forests. Our understanding of the patterns of fine root turnover is limited, in particular in tropical forests, despite its acknowledged importance in the global carbon cycle. We used the minirhizotron technique for studying the changes in fine root longevity and turnover along a 2000-m-elevational transect in the tropical mountain forests of South Ecuador. Fine root growth and loss rates were monitored during a 5-mo period at intervals of four weeks with each 10 minirhizotron tubes in three stands at 1050, 1890, and 3060 m asl. Average root loss rate decreased from 1.07 to 0.72 g/g/yr from 1050 to 1890 m, indicating an increase in mean root longevity with increasing elevation. However average root loss rate increased again toward the uppermost stand at 3060 m (1.30 g/g/yr). Thus, root longevity increased from lower montane to mid-montane elevation as would be expected from an effect of low temperature on root turnover, but it decreased further upslope despite colder temperatures. We suggest that adverse soil conditions may reduce root longevity at high elevations in South Ecuador, and are thus additional factors besides temperature that control root dynamics in tropical mountain forests.