Applications of functional classification methods for tree fine root biomass estimation: Advancements and synthesis

The morphology of fine root branching of woody plants is highly variable in their forms and functions. In the past two decades, researchers have increasingly recognized that the root-diameter-based method, using an arbitrary size of root diameter, failed to precisely characterize the physiological and ecological processes involved in finest roots. The number of publications using root-order-based approaches has increased regardless the fact that root trait-measurements based on root order are time-consuming and labor-intensive. A new approach—root functional classification method—was proposed and had been applied in the literature. The functional classification of fine roots separates roots of < 2 mm to absorptive and transport pools, making it more feasible for studies on root biomass and turnover. This new concept redefines fine root guild and has great potentials for future studies. Our literature review of the topic indicates that less is known about the inter-specific differences in estimates of biomass of absorptive and/or transport roots, with a large variation of absorptive roots on global scale. In addition, our review emphasizes the importance in: a) precision estimating of the absorptive biomass of fine roots, and b) proper definition of the range of the transport roots within and among forest ecosystems. Finally, after compare the strengths and weaknesses of the functional classification method, we propose several specific suggestions to improve the applications of this approach.     

Root strength,growth, and rates of decay: root reinforcement changes of two tree species and their contribution to slope stability

Information on live root-wood strength, rates of root decay and root growth of both radiata pine (Pinus radiata D. Don) and kanuka (Kunzea ericoides (A. Rich.) Joy Thomps. var. ericoides) are combined to form a generalized conceptual model of changes in nett root reinforcement. The model provides an initial opportunity to rank the plant species having specific below-ground rooting habits that can be used to control erosion, and when linked with extreme flood probability can be used to indicate the risk of a storm likely to cause slope instability in the period between clear-felling and regrowth. Erosion-susceptible slopes planted 1 year after clearfelling in radiata pine at 1250 stems ha^-1 regain root site-occupancy in 4.7 years, an interval during which there is an 80% chance of experiencing an extreme flood. Similarly for radiata planted at 800 and 400 stems ha^-1, root site-occupancy is regained in 5.6 and 7.5 years, and the probability of occurrence of an extreme event within these periods is 85 and 90%, respectively. For erosion-susceptible slopes on which kanuka has become established, the probability of a significant event within the 2.8 years prior to root site-occupancy is 60%. Slopes felled of radiata pine are potentially more vulnerable to the stresses promoting slope instability, at least in the earlier years. This revised version was published online in June 2006 with corrections to the Cover Date.     

Linking root respiration to chemistry and morphology across species

Root respiration is a critical physiological trait involved in root resource acquisition strategies, yet it is less represented in root trait syndrome. Here we compiled a large dataset of root respiration associated with root chemical and morphological traits from 245 plant species. Our results demonstrated that root respiration correlated positively with root nitrogen concentration (RNC) and negatively with root tissue density (RTD) across and within woody and non‐woody species. However, the relationships between root respiration and specific root length (SRL) and root diameter (RD) were weak or even insignificant. Such root respiration–traits relationships were not completely in line with predictions by the root economics spectrum (RES). Furthermore, the principal component analysis showed that root trait syndrome was multidimensional. Root respiration was associated more strongly with the RNC‐RTD axis (the classical RES) than with the orthogonal SRL‐RD axis for woody species, but not for non‐woody species. Collectively, the linkages of root physiological, chemical, and morphological traits provide a better understanding of root trait covariation and root resource acquisition strategies.     

Relationship between root elongation rate and diameter and duration of growth of lateral roots of maize

The objective of this work was to describe the relationship between elongation rate and diameter of maize roots and to estimate the length and growth duration of lateral roots of maize. Diameters and elongation rates of roots were measuredin situ on plants grown 5 weeks in small rhizotrons under greenhouse conditions. At the end of the experimental period the roots were harvested and diameters of axile and lateral roots were measured. The frequency distribution of diameters of harvested roots was bimodal with a minimum at 0.6 mm; 97% of axile roots were larger than this value and 98% of the lateral roots were smaller. Root elongation per day increased as diameter increased but the slope of the relationship with lateral roots was about 2.5 times that with axile roots when separate linear regressions were fitted to the two populations. The length of lateral roots found on axillary roots between the base and about 30 cm from the apex was approximately 2.2 cm. All of the data was consistent with the hypothesis that the lateral roots grew for about 2.5 days and then ceased growing. The axillary roots continued to grow throughout the experimental period at a rate of about 3 cm day⁻¹. Contribution from the Department of Agronomy, New York State College of Agriculture and Life Sciences, Cornell University, Ithaca, NY 14853. Agronomy paper No. 1661. This research is part of the program of the Center for Root-Soil Research.     

Influence of Fusarium solani on citrus root rot caused by Phytophthora parasitica and Phytophthora citrophthora

Interactions between Fusarium solani and Phytophthora parasitica or F. solani and P. citrophthora influenced the development of root rot of citrus but depended on the temporal order of inoculation with F. solani or the two Phytophthora spp. Inoculation of citrus with either Fusarium solani and Phytophthora parasitica or Phytophthora citrophthora increased root rot compared to inoculation with P. parasitica or P. citrophthora alone when plants were inoculated with Phytophthora by dipping their roots in zoospore suspensions and subsequently transplanted into soil infested with F. solani. However, root rot was not increased by simultaneous co-inoculation of P. parasitica and F. solani or when plants were inoculated with F. solani first. Root rot was not increased when heat-stressed or non-stressed plants were inoculated with P. parasitica 30 days after transplanting into soil infested with F. solani. In most but not all experiments, F. solani alone reduced growth of tops or roots a small but significant amount.Co-inoculation of citrus by root-dipping into zoospore suspensions of P. parasitica and transplanting into soil infested with F. solani reduced feeder root length by 62% and root weight by 61% but did not significantly reduce the percentage of living roots when compared to inoculation with P. parasitica alone. When citrus roots were immersed in zoospore suspensions of P. citrophthora and transplanted into soil infested with F. solani, feeder root length was reduced by 68%, but feeder root weight and the percentage of living roots were not significantly reduced when compared to plants inoculated with P. citrophthora alone.Propagule densities of both P. parasitica and P. citrophthora in the rhizosphere of plants inoculated by root-immersion and then transplanting into soil infested with F. solani were not significantly different than propagule densities from plants transplanted into non-infested soil. Propagule densities of P. parasitica were suppressed an average of 41% when citrus was inoculated with P. parasitica 30 days after transplanting into soil infested with F. solani and by 41% when citrus was co-inoculated by transplanting into soil infested with both F. solani and P. parasitica.     

不同分类系统下油松幼苗根系特征的差异与联系

植物根序和径级不仅反映细根的形态结构, 而且能反映根系的一些生理特征, 如细根寿命和周转等。该文以二年生油松(Pinus tabulaeformis)幼苗根系为研究对象, 系统比较了根序分类方法和径级分类方法在描述根系特征上的优缺点, 探索了两者之间的内在联系。结果表明: 二年生油松幼苗最多可包括6级根序, 直径的变化范围为0.169-3.877 mm。按根序划分, I-VI级根序的总根长和总根表面积主要集中在前3级根序, 这3级根序的根占总根长的78.77%和总根表面积的62.72%。前3级根序的比根长是后3级根序比根长的1.3-3.0倍, 比根面积是后3级比根面积的1.0-1.5倍。按常用的径级(以0.5、1.0、1.5和2.0 mm为阈值)划分方法, 油松幼苗大部分根系直径≤1.5 mm, 此区间细根的根长和根表面积占总根长的93.76%和总根表面积的84.35%。直径≤1.5 mm的细根平均比根长是>1.5 mm细根比根长的3-7倍, 比根面积的1.5-3.0倍。由于油松根序和径级之间有显著的指数关系, 依据径级最大程度反映根序的原则, 提出了新的径级划分方法, 即以0.4、0.8、1.3和2.0 mm为阈值对油松幼苗根系径级重新进行划分。此时, 上述区间可分别包括I级、II级、III级、IV级、V级根序中根尖数的93.22%、86.37%、75.96%、70.47%和76.67%。同时也可分别涵盖各径级根长的89.34%-70.83%、根面积的86.01%-76.12%以及体积的87.73%-76.12%。此时, 根系不同径级与根序之间可以建立起良好的对应关系。这些结果表明, 通过合理划分径级区间可以较好地反映根序 特征。     

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。     

Sample preparation and scanning protocol for computerised analysis of root length and diameter

Root length and diameter distribution are important characteristics to be considered when describing and comparing root systems. Root length and root-diameter distribution may be obtained in two ways: by microscopical measurements, which are laborious, or by computerised analysis, which is fast but sensitive to the scanning protocol. Although scanning protocols vary widely between laboratories, papers rarely report the details. Using two commercially available root analysis software packages (Delta-T Scan and WinRHIZO), we performed a sensitivity analysis of scanning protocols for estimating root length and diameter distribution. The results are shown graphically, rather than numerically, because the interactions between different parameters in the overall scanning protocol are most clearly illustrated by the shape of the curves. The present analysis clearly demonstrated the sensitivity of the two scanning methods with regard to staining period, maximum root density, scanning resolution and transformation threshold. For example, estimating the root-diameter distribution versus measuring root length, puts opposite constraints on the transformation threshold settings. We suggest the following settings for the most sensitive parameters: a staining period of 24 h, a root density of less than 0.5 mm root per mm² surface, a resolution of 400 dpi and the automatic threshold for WinRHIZO and a brightness of 200 for Delta-T Scan. According to this protocol, comparison of computerised analyses with microscopic measurements showed good total root length and diameter distribution agreement for three contrasting root systems. We suggest to always start with the present protocol when studying other species. If validation indicates that the proposed scanning protocol needs to be modified for other species, the present sensitivity analysis may be used as a guideline for changing the most critical parameters. Similarly, the use of another stain than neutral red may also require modifications of the scanning protocol. In general, a long staining period (e.g., 24 h) is recommended for all stains, as small differences in staining period have the least effect when the root tissue is saturated. To enhance comparability of results in the literature, the staining period, stain, (maximum) root density, scanner resolution and threshold should always be listed when root data obtained by scanning are presented. This revised version was published online in August 2006 with corrections to the Cover Date.     

落叶松和水曲柳不同根序细根形态结构、组织氮浓度与根呼吸的关系

根系具有高度的形态和生理功能异质性,在森林生态系统碳和养分循环中起重要作用。根系分枝的顺序构成根序,是根系最基本的构型特征,根序代表根系不同的发育阶段。然而,目前直接测定不同根序细根生理功能的研究很少。以落叶松（Larix gmelinii）和水曲柳（Fraxinus mandshurica）的细根为研究对象,使用气相氧电极测定不同根序细根的呼吸速率,探讨根系呼吸速率与其形态、结构和组织氮浓度的关系。结果表明：落叶松和水曲柳细根的直径、根长和维管束直径均随着根序的增加（1–5级）而增加,而比根长、组织氮浓度和呼吸速率随着根序的增加而降低,各根序之间差异显著（P〈0.05）;1级根比根长最大、皮层组织发达、组织氮浓度最高且呼吸速率也最高,其呼吸速率分别为17.57nmolO2·g^–1·s^–1（落叶松）和18.80 nmolO2·g^–1·s^–1（水曲柳）,比5级根分别高148%（落叶松）和124%（水曲柳）;并且,落叶松根的呼吸速率几乎有96%与根系组织氮浓度相关,而水曲柳根的呼吸速率则有89%与根系组织氮浓度相关。上述结果说明,细根的形态和生理功能异质性是紧密相连的,低级根的形态、结构决定其功能是吸收养分和水,而高级根的形态、结构决定其功能是运输和贮存养分。     

Development and validation of a model to describe root length density of maize from root counts on soil profiles

Root length density (RLD) is an important determinant of crop water and nutrient acquisition, but is difficult to measure in the field. On a soil profile, in-situ counts of root impacts per unit surface on soil profiles (NI) can be used to calculate RLD if crop-specific parameters for preferential root orientation (anisotropy) are known. An improved method for field determinations of RLD was developed and validated for maize at sites in Côte d'Ivoire and Burkina Faso. Root anisotropy was measured with cubes of undisturbed soil with 0.1 m sidelength, based on NI observed on three planes oriented perpendicularly to each other. RLD was also measured for the enclosed volume. Repetition of such measurements enabled estimation of the robustness across sites of empirical and geometric models for the relationship between RLD and NI:RLD = NI CO, with CO being the coefficient of root orientation, theoretically equals 2 for an isotropic distribution. Root systems were found to be nearly isotropic, except near the root front (0.3 to 0.5 m), where roots had a preferentially orthotropic orientation. Measured RLD was generally about 50% larger than RLD calculated from observed NI and CO, indicating that at least one of the measurement techniques had a systematic error. The ratio between measured and calculated RLD (CE), which ranged from 0.8 to 2, increased with the age of the plants and decreased with soil depth. CE was therefore introduced as an additional coefficient, resulting in RLD = NI CO CE. The empirical value for CO CE was between 2 and 5. The empirical coefficients CO and CE were the same for the sites in Cote d'Ivoire (oxisol with an iron pan at 0.6 to 0.9 m) and Burkina Faso (alfisol with an iron pan at 0.4 to 0.8 m). The model was validated with independent data sets at both sites, and gave satisfactory predictions of RLD on the basis of NI obtained from single soil planes, which can be easily measured in the field.     

Drip irrigation,soil characteristics and the root distribution and root activity of olive trees

A study was carried out on the root distribution and root activity of the olive tree (Olea Europaea, L., var. manzanillo) as influenced by drip irrigation and by several soil characteristics such as texture and depth. The experiments were conducted in two plots within a drip-irrigated grove of 20-year-old trees planted at 7×7 m spacing. One soil was a sandy loam, the other a clay-loam. Both cylinder and trench methods were used to determine root distribution. Labelling with ³²P was used to determine root activity. Under dryland conditions the adult tree adapted its rooting system, following the installation of a drip system, by concentrating the roots within the wet soil zones near the drippers. The highest root densities occur in those zones, down to a 0.6 m depth, the most abundant being the <0.5 mm diameter roots. The most intensive root activity was also found in that zone. For a given irrigation system, wet soil bulbs are more extensive and therefore root distribution expands to a larger soil volume when the soil is more clayey and with a hard calcareous pan present at about 0.8 m depth which prevents deep drainage.     

Using coloured roots to study root interaction and competition in intercropped legumes and non-legumes

Aims Root interactions between neighbour plants represent a fundamental aspect of the competitive dynamics in pure stand and mixed cropping systems. The comprehension of such phenomena places big methodological challenges, and still needs clarification. The objectives of this work were (i) to test if a species with coloured roots can be used to examine the interaction in a legume-non-legume intercropping system; (ii) to verify the importance of initial root growth on the successive root development of mixture component plants; (iii) to test if the root interaction in the shallow layers has consequences for deep root growth and (iv) to compare the effect of intraspecific and interspecific competition on root development and biomass growth.Methods A detailed study on root growth and interaction was carried out using rhizotron tubes where two legume species were grown in pure stands or were intercropped with red beet, a variety of Beta vulgaris L. with clear red roots. Within the rhizotrons, the three species were grown either without competitors, with two plants of the same species to measure intraspecific competition or with one legume and one red beet plant to study interspecific competition. The use of mixtures where one component has clearly coloured roots, together with several scalar measurements of root depth and proliferation, allowed the measurement of the root system of each species when grown in the mixtures.Important findings The use of rhizotron tubes coupled with species with coloured roots represented a valuable method to study the belowground interaction in mixed cropping systems. The initial root growth was a very important feature for the subsequent dominance of a species and it was not related to seed dimension. Initial root growth was also important because the root interactions in the shallower soil layers were found to influence the root growth in deeper soil. The root system of the red beet showed much faster and deeper growth than that of the legumes, and made red beet the dominant component in the mixtures while the legume root system was confined to the shallower soil layer. Intraspecific competition was well tolerated by the legumes, but it was limiting for the highly competitive red beet. The outcome of root interaction between neighbour plants was confirmed to be species-specific as it changed according to the intensity of the competitive effect/response of each species of the mixture: both legumes were slightly affected by the intraspecific and highly affected by interspecific competition while red beet was more affected by intraspecific competition but strongly dominant when intercropped with legumes.     

CO2 enrichment increases carbon and nitrogen input from fine roots in a deciduous forest

* Greater fine-root production under elevated [CO2] may increase the input of carbon (C) and nitrogen (N) to the soil profile because fine root populations turn over quickly in forested ecosystems. * Here, the effect of elevated [CO)] was assessed on root biomass and N inputs at several soil depths by combining a long-term minirhizotron dataset with continuous, root-specific measurements of root mass and [N]. The experiment was conducted in a CO(2)-enriched sweetgum (Liquidambar styraciflua) plantation. * CO2) enrichment had no effect on root tissue density or [N] within a given diameter class. Root biomass production and standing crop were doubled under elevated [CO2]. Though fine-root turnover declined under elevated [CO2], fine-root mortality was also nearly doubled under CO2 enrichment. Over 9 yr, root mortality resulted in 681 g m(-2) of extra C and 9 g m(-2) of extra N input to the soil system under elevated [CO2]. At least half of these inputs were below 30 cm soil depth. * Increased C and N input to the soil under CO2 enrichment, especially below 30 cm depth, might alter soil C storage and N mineralization. Future research should focus on quantifying root decomposition dynamics and C and N mineralization deeper in the soil.     

亚热带6种树种细根序级结构和形态特征

以福建省建瓯市万木林自然保护区内占优势的6种天然林树种(沉水樟Cinnamomum micranthum,CIM;观光木Tsoongiodendron odorum Chun,TOC;浙江桂Cinnamomum chekiangense,CIC;罗浮栲Castanopsis fabri,CAF;细柄阿丁枫Altingiagracilipes,ALG;米槠Castanopsis carlesii,CAC)为研究对象,对其1—5级细根的结构,形态特征及生物量进行了分析。结果表明:沉水樟,细柄阿丁枫和米槠细根分支比表现出在1,2级(4倍以上)明显大于其它序级(3倍左右);其余3种树种则是在3,4级的细根分支比最大,其中浙江桂达到8.65倍,其它序级则大致为3倍左右。6种树种1,2级细根数量占到总数的70%—90%。6种树种细根直径,根长,组织密度随序级升高逐渐增大,比根长减小,生物量未表现出一致的变化规律,6种树种生物量主要集中在高级根部分。方差分析表明,树种对细根分支比例有显著影响(P<0.05),浙江桂和米槠细根分支水平对分支比例有极显著影响(P<0.01),其余4种树种分支水平对分支比例有显著影响(P<0.05),树种和分支水平的交互作用对6种树种细根分支比均有极显著的影响(P<0.01);树种对细根根长,直径以及生物量均有极显著影响(P<0.01),对比根长有显著影响(P<0.05),而对组织密度的影响则不显著(P>0.05);树种和序级的交互作用对细根根长,直径以及生物量均有极显著影响(P<0.01),对组织密度有显著影响(P<0.05),对比根长影响不显著(P>0.05)。序级对6种树种细根根长,直径,比根长以及生物量的影响并未达到一致,对6种树种细根组织密度有极显著影响(P<0.01)。树种间1—4级根的比根长变异主要由组织密度引起,而5级根的比根长变异则由直径引起,同时在1级根中组织密度与直径呈现出权衡的关系。6种树种细根数量,直径,根长,比根长,组织密度以及生物量与序级之间回归分析发现它们与序级之间具有指数函数,线性函数,二次函数,三次函数或者幂函数关系。     

Genotype and Planting Density Effects on Rooting Traits and Yield in Cotton （Gossypium hirsutum L,）

Root density distribution of plants is a major Indicator of competition between plants and determines resource capture from the solh This experiment was conducted in 2005 at Anyang, located in the Yellow River region, Henan Province, China. Three cotton （Gossyplum hlrsutum L.） cultivars were chosen： hybrid Btcultlvar CRI46, conventional Btcultlvars CRI44 and CRI45. Six planting densities were designed, ranging from 1.5 to 12.0 plants/m^2. Root parameters such as surface area, diameter and length were analyzed by using the DT-SCAN Image analysis method. The root length density （RLD）, root average diameter and root area Index （RAI）, root surface area per unit land area, were studied. The results showed that RLD and RAI differed between genotypes; hybrid CRI46 had significantly higher （P 〈0.05） RLD and RAI values than conventlonal cultlvars, especially under low planting densities, less than 3.0 plants/m^2. The root area index （RAI） of hybrid CRI46 was 61% higher than of CRI44 and CRI45 at the flowering stage. The RLD and RAI were also significantly different （P = 0.000） between planting densities. The depth distribution of RAI showed that at Increasing planting densities RAI was Increasingly distributed in the soil layers below 50 cm. The RAI of hybrid CRI46 was for all planting densities, obviously higher than other cultivars during the flowering and boll stages. It was concluded that the hybrid had a strong advantage in root maintenance preventing premature senescence of roots. The root diameter of hybrid CRI46 had a genetically higher root diameter at planting densities lower than 6.0 plants/m^2. Good associations were found between yield and RAI In different stages. The optimum planting density ranged from 4.50 plants/m^2 to 6.75 plants/m^2 for conventional cultlvars and around 4.0-5.0 plants/m^2 for hybrids.     

Quantifying relationships between rooting traits and water uptake under drought in Mediterranean barley and durum wheat

In Mediterranean regions drought is the major factor limiting spring barley and durum wheat grain yields. This study aimed to compare spring barley and durum wheat root and shoot responses to drought and quantify relationships between root traits and water uptake under terminal drought.One spring barley（Hordeum vulgare L. cv. Rum） and two durum wheat Mediterranean cultivars（Triticum turgidum L. var durum cvs Hourani and Karim） were examined in soil‐column experiments under well watered and drought conditions. Root system architecture traits, water uptake, and plant growth were measured. Barley aerial biomass and grain yields were higher than for durum wheat cultivars in well watered conditions. Drought decreased grain yield more for barley（47%） than durum wheat（30%, Hourani）. Root‐to‐shoot dry matter ratio increased for durum wheat under drought but not for barley, and root weight increased for wheat in response todrought but decreased for barley. The critical root length density（RLD） and root volume density（RVD） for 90% available water capture for wheat were similar to（cv. Hourani） or lower than（cv. Karim） for barley depending on wheat cultivar. For both species, RVD accounted for a slightly higher proportion of phenotypic variation in water uptake under drought than RLD.