水曲柳和落叶松细根形态及母根与子根比例关系

细根（直径〈2mm）的分枝是根系重要的结构特征,不同根序等级的细根在养分和水分吸收、C的消耗和寿命方面具有较大的差异,定量研究各根序等级之间的比例关系对认识细根死亡的顺序具有重要的理论意义。根据Pregitzer等2002年提供的方法,研究了17年生水曲柳（Fraxinus mandshurica Rupr．）和落叶松（Larix gmelinii Rupr．）人工纯林1-5级细根的直径、长度、比根长、生物量和数量。结果表明,两树种细根中1级根序的数量占总根系数量80％-90％,它们直径小、长度短、比根长高。随着根序等级（1级-5级）的增加细根直径增粗和长度增加、比根长减小。细根的数量和生物量在上下土层的分布受土壤资源有效性的影响。水曲柳5级根序-2级根序之间母根与子根的数量关系是1：3,落叶松是1：2-3。2级根序与1级根序之间母根与子根的数量关系,水曲柳是1：10—12,落叶松是1：8。如果当年生长的1级细根当中保持1：3的比例,将有65％-75％的1级细根死亡,占根系总数的55％～65％,总长度的40％-50％,以及总生物量的20％-30％。     

Effects of liming on rhizosphere chemistry and growth of fine roots and of shoots of sessile oak (Quercus petraea)

Soil acidification can be detrimental to root growth and nutrient uptake, and liming may alleviate such acidification. In the following study, seedlings of sessile oak (Quercus petraea Liebl. M.) were grown in rhizotrons and subjected to liming (L) or gypsum (G) treatments and compared with the control (C). In order to study and interpret the impact of these calcium rich treatments on fine root development and tree growth, the following parameters were assessed: fine root biomass, fine root length, seedling development (height, diameter, leaves), seedling biomass, nutrient content of roots and seedlings, bulk soil and soil solution chemistry and rhizosphere soil chemistry. The results show that liming increased bulk soil pH, exchangeable Mg, Ca and the Ca/Al molar ratio, and decreased exchangeable Al, mainly in the A-horizon. Gypsum had a similar but smaller impact on exchangeable Al, Ca, H⁺ and the Ca/Al molar ratio in the A-horizon, but reacted with depth, so that exchangeable Mn, Mg and Ca were increased in the B-horizon. In the rhizosphere, the general pattern was determined by the treatment effects of the bulk soil. Most elements were more concentrated in the rhizosphere than in bulk soil, except for Ca which was less concentrated after liming or gypsum application. In the B-horizon rhizosphere pH was increased by the treatments (L > G,C) close to the root tips. Furthermore, the length of the zone with a positive root-induced pH increase was greater for the limed roots as compared with both the other treatments. Fine root growth was stimulated by liming (L > G,C) both in terms of biomass and length, whereas specific root length was not obviously affected apart from the indication of some stimulation after liming at the beginning. The live:dead ratio of fine roots was significantly higher in the limed rhizotrons as compared to the control (G not assessed), indicating lower mortality (higher longevity). Shoot growth showed greater lime-induced stimulation (L > G,C) as compared to root growth. As a result the shoot:root ratio was higher in the limed rhizotrons than in the control (L > G,C). Liming induced a higher allocation of P, S, Mg, Ca and K to the leaves, stem and twigs. Gypsum showed similar effects, but was only significant for S. Liming increased the foliar Ca/Al ratio by both increasing foliar Ca and decreasing foliar Al, whereas gypsum did not clearly improve foliar nutrition. This study suggests that a moderate application of lime can be successful in stimulating seedling growth, but that gypsum had no effect on seedling growth. It can be concluded that this lime-induced growth stimulation is directly related to the improved soil fertility status, and the alleviation of Al toxicity and acid stress, resulting in better foliar nutrition. The impact of liming on fine roots, as a consequence, was not limited to a stimulation of the total amount of fine roots, but also improved the root uptake performance. This revised version was published online in June 2006 with corrections to the Cover Date.     

Seasonal development of loblolly pine lateral roots in response to stand density and fertilization

In 1989, two levels each of stand density and fertilization treatments were factorially established in a 9-year-old loblolly pine plantation on a P-deficient Gulf Coastal Plain site in Rapides Parish, Louisiana, USA. In 1995, a second thinning was conducted on the previously thinned plots and fertilizer was re-applied to the previously fertilized plots. The morphology of new long lateral roots was evaluated at 2-week intervals in five Plexiglas rhizotrons per plot of two replications. The overall objective of this study was to evaluate the seasonal initiation of six morphological categories of long lateral roots ( 2.5 cm in length) in response to stand density and fertilization. Lateral root development exhibited a seasonal pattern with the initiation of branched lateral roots predominantly occurring in spring and summer. The initiation of non-branched lateral roots occurred throughout the year regardless of season. Stand density did not affect lateral root morphological development. However, fertilization stimulated the initiation of branched lateral roots that were greater than 1 mm in diameter.     

杨树细根及草根的生产力与周转的研究

对北方杨树人工林0－40cm土层中杨树细根和草根（≤2mm）年生物量、分解量、死亡量、生长量和周转率进行观察研究。结果表明,杨树细根的年生物量为2．062t.hm^-2,死根生物量为0．746t.hm^-2,分解量为0．158t.hm^2,生长量为2．351t.hm^－2,周转率为每年1．14次,活草根的年生物量、死根生物量、分解量、生长量和周转率分别0．501、0．035、0．023、0．691t.hm^-2和1．38,同时给出了杨树细根干重损失随分解时间变化的方程：1nx/x0=0.9515e^-0.0014t。     

Fine root production and nutrient content in wet and moist arctic tundras as influenced by chronic fertilization

We used ingrowth cores to estimate fine root production in organic soils of wet sedge and moist tundra ecosystems near Toolik Lake on Alaska's North Slope. Root-free soil cores contained in nylon mesh tubes (5 cm diameter, 20–30 cm long) were placed in control and chronically fertilized (N plus P) plots in mid-August 1994 and were retrieved 1 year later. Estimated fine root production in control plots was 75 g m^–2 year^–1 in wet sedge and 56 g m^–2 year^–1 in moist tussock tundra. Fine root production in fertilized plots was 85 g m^–2 year^–1 in wet sedge and 67 g m^–2 year^–1 in moist tussock tundra. Although our estimates of fine root production were higher on fertilized than control plots, differences were not statistically significant within either tundra type. Comparisons between our estimates of fine root production and other estimates of aboveground (plus rhizome) production on the same (wet sedge tundra) or similar (moist tussock tundra) plots suggest that fine root production was about one-third of total net primary production (NPP) under non-fertilized conditions and about one-fifth of total NPP under chronic fertilization. Fine root N and P concentrations increased with fertilization in both tundra types, but P concentrations increased more than N concentrations in wet sedge tundra, whereas relative increases in N and P concentrations in moist tundra roots were similar. These data are consistent with other studies suggesting that NPP in wet sedge tundra is often P limited and that co-limitation by N and P is more important in moist tussock tundra.     

杉木,火力楠纯林及混交林细根周转的研究

系统研究了杉木、火力楠纯林及混交林细根生物量、生产力及年周转率。结果表明,杉木、火力楠纯林及混交林活细根生物量分别为８８０、３０３５和１５６０ｋｇ．ｈａ＾－１;死细根生物量为２３５、３９８和５６５ｋｇ．ｈａ＾－１;细根年周转率为１．２９、１．４２和１．４０;年生产量为１１３７、４３１８和２１７９ｋｇ．ｈａ＾－１;年死亡量为４９７．５９５和１１４９ｋｇ．ｈａ＾－１,分别相当于林分枯枝落叶年凋落量为３     

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.     

Automatic discrimination of fine roots in minirhizotron images

Minirhizotrons provide detailed information on the production, life history and mortality of fine roots. However, manual processing of minirhizotron images is time-consuming, limiting the number and size of experiments that can reasonably be analysed. Previously, an algorithm was developed to automatically detect and measure individual roots in minirhizotron images. Here, species-specific root classifiers were developed to discriminate detected roots from bright background artifacts. Classifiers were developed from training images of peach (Prunus persica), freeman maple (Acer x freemanii) and sweetbay magnolia (Magnolia virginiana) using the Adaboost algorithm. True- and false-positive rates for classifiers were estimated using receiver operating characteristic curves. Classifiers gave true positive rates of 89-94% and false positive rates of 3-7% when applied to nontraining images of the species for which they were developed. The application of a classifier trained on one species to images from another species resulted in little or no reduction in accuracy. These results suggest that a single root classifier can be used to distinguish roots from background objects across multiple minirhizotron experiments. By incorporating root detection and discrimination algorithms into an open-source minirhizotron image analysis application, many analysis tasks that are currently performed by hand can be automated.     

Fine root dynamics in a loblolly pine forest are influenced by free-air-CO2-enrichment: a six-year-minirhizotron study

Efforts to characterize carbon (C) cycling among atmosphere, forest canopy, and soil C pools are hindered by poorly quantified fine root dynamics. We characterized the influence of free‐air‐CO₂‐enrichment (ambient +200 ppm) on fine roots for a period of 6 years (Autumn 1998 through Autumn 2004) in an 18‐year‐old loblolly pine (Pinus taeda) plantation near Durham, NC, USA using minirhizotrons. Root production and mortality were synchronous processes that peaked most years during spring and early summer. Seasonality of fine root production and mortality was not influenced by atmospheric CO₂ availability. Averaged over all 6 years of the study, CO₂ enrichment increased average fine root standing crop (+23%), annual root length production (+25%), and annual root length mortality (+36%). Larger increase in mortality compared with production with CO₂ enrichment is explained by shorter average fine root lifespans in elevated plots (500 days) compared with controls (574 days). The effects of CO₂‐enrichment on fine root proliferation tended to shift from shallow (0–15 cm) to deeper soil depths (15–30) with increasing duration of the study. Diameters of fine roots were initially increased by CO₂‐enrichment but this effect diminished over time. Averaged over 6 years, annual fine root NPP was estimated to be 163 g dw m^?2 yr^?1 in CO₂‐enriched plots and 130 g dw m^?2 yr^?1 in control plots (P= 0.13) corresponding to an average annual additional input of fine root biomass to soil of 33 g m^?2 yr^?1 in CO₂‐enriched plots. A lack of consistent CO₂× year effects suggest that the positive effects of CO₂ enrichment on fine root growth persisted 6 years following minirhizotron tube installation (8 years following initiation of the CO₂ fumigation). Although CO₂‐enrichment contributed to extra flow of C into soil in this experiment, the magnitude of the effect was small suggesting only modest potential for fine root processes to directly contribute to soil C storage in south‐eastern pine forests.     

Earthworm effect on root morphology in a split root system

Plants respond to their environment through adaptations such as root proliferation in nutrient-rich patches. Through their burrows and casts production in soil, earthworms create heterogeneity which could lead to local root adaptations or systemic effects. To investigate the effect of earthworms on root system morphology and determine whether earthworm effect is local or systemic, we set up two independent split root experiments with rice or barley, (i) without earthworm (CC), (ii) with earthworms in both compartments (EE), and (iii) with earthworms in one single compartment (CE). Earthworms had an effect on belowground plant biomass. The relative length of thick roots decreased with an increasing abundance of earthworms. Some root diameter classes responded to earthworm number in a linear or curvilinear way, making simple conclusions difficult. We found no difference in root biomass or morphology between the two compartments of the split root system in the CE treatment, but a positive effect of earthworm biomass on root biomass, volume, surface area, and length at the whole plant level. Results supported a systemic effect dependent on earthworm abundance. Modification of nutrient mineralization, soil physical structure, and/or the concentration of signal molecules could all be responsible for this systemic effect.     

幼龄柠条细根的空间分布和季节动态

以晋西北黄土高原区5年生柠条(Caragana korshinskii)人工林为研究对象,应用Minirhizotron技术,分别在距茎干水平距离0 cm和50 cm处设点(以下简称为0 cm位点和50 cm位点),对林地0—100 cm土层深度范围内的柠条细根进行了观测。以2009年生长季(4—10月)的细根根长密度(RLD,mm/cm2)和表面积密度(RAD,mm2/cm2)数据为基础,结合同期环境因子(气温、降雨量、土壤温度和土壤含水量等)数据,对0 cm和50 cm两个位点的细根动态特点进行了比较研究。结果表明:(1)两个水平位点的细根垂直分布和季节变化趋势均具有一定差异,主要差异是0 cm位点0—60 cm各土层的RLD均大于50 cm位点,前者各测定期的RLD(RAD)均大于后者。因此,0 cm位点的细根分布量(4.04 mm/cm2和4.67 mm2/cm2)显著大于50 cm位点(3.07 mm/cm2和2.99 mm2/cm2)。(2)就整体(两个位点平均值)而言,RLD(RAD)的垂直分布以40—50cm土层最大,以60—70cm土层最小。RLD(RAD)的季节变化具有由小变大再变小的趋势。年生长季幼龄柠条细根的RLD和RAD总平均值分别为3.55 mm/cm2和3.83 mm2/cm2。(3)就0 cm位点、50 cm位点或整个林地而言,细根RLD的季节变化与气温和土壤温度的季节变化均具有显著正相关性。以上结果表明,幼龄柠条细根的水平分布具有"近主根"特点;RLD的季节变化与温度因子的季节变化具有高度一致性。     

Fine roots, arbuscular mycorrhizal hyphae and soil nutrients in four neotropical rain forests: patterns across large geographic distances

* It is commonly hypothesized that stand-level fine root biomass increases as soil fertility decreases both within and among tropical forests, but few data exist to test this prediction across broad geographic scales. This study investigated the relationships among fine roots, arbuscular mycorrhizal (AM) fungi and soil nutrients in four lowland, neotropical rainforests. * Within each forest, samples were collected from plots that differed in fertility and above-ground biomass, and fine roots, AM hyphae and total soil nutrients were measured. * Among sites, total fine root mass varied by a factor of three, from 237+/-19 g m-2 in Costa Rica to 800+/-116 g m-2 in Brazil (0-40 cm depth). Both root mass and length were negatively correlated to soil nitrogen and phosphorus, but AM hyphae were not related to nutrients, root properties or above-ground biomass. * These results suggest that understanding how soil fertility affects fine roots is an additional factor that may improve the representation of root functions in global biogeochemical models or biome-wide averages of root properties in tropical forests.     

The effect of nitrogen fertilization on the phenology of roots in a barrier island sand dune community

Little work has been done on the phenology of root growth and senescence largely due to methodological difficulties. The application of minirhizotron technology has enabled the tracking of individual roots through an entire growing season. As a result, direct measures of mortality, root growth, and an analysis of cohorts can be obtained. This study examined the belowground response of vegetation in a nutrient limited system to nitrogen addition. Small plots on a 36 year old dune on Hog Island, a barrier island in the Virginia Coast Reserve Long Term Ecological Research Site, were fertilized with nitrogen. Minirhizotron tubes were installed in each fertilized and control plot. Each tube was sampled monthly for nine months, March through October of 1992. Root length density increased throughout the growing season with the greatest root length density in the top 20 cm of the soil profile. The fertilized plots had greater root length densities (14.1 mm cm^-2) than the unfertilized plots (2.9 mm cm^-2). There was no significant depth × treatment interaction. Root mortality did not significantly change with fertilization. The largest loss of roots for a cohort occurred within the first month. The dune grassland community did not respond to fertilization with large changes in root distribution or increases in mortality in this study.     

Rapid decline in nitrate uptake and respiration with age in fine lateral roots of grape: implications for root efficiency and competitive effectiveness

Changes in function as an individual root ages has important implications for understanding resource acquisition, competitive ability and optimal lifespan. Both nitrate uptake and respiration rates of differently aged fine roots of grape (Vitis rupestris x V. riparia cv. 3309 C) were measured. The resulting data were then used to simulate nitrate uptake efficiency and nutrient depletion as a function of root age. Both nitrate uptake and root respiration declined remarkably quickly with increasing root age. The decline in both N uptake and root respiration corresponded with a strong decline in root N concentration, suggesting translocation of nitrogen out of the roots. For simulations where no nutrient depletion occurs at the root surface, daily uptake efficiency was maximal at root birth and lifetime nitrate uptake efficiency slowly increased as the roots aged. Simulations of growth of roots into unoccupied soil using a solute transport model indicated the advantage of high uptake capacity in new roots under competitive conditions where nitrate availability is very transitory.     

Anatomical patterns of condensed tannin in fine roots of tree species from a cool-temperate forest

Background and AimsCondensed tannin (CT) is an important compound in plant biological structural defence and for tolerance of herbivory and environmental stress. However, little is known of the role and location of CT within the fine roots of woody plants. To understand the role of CT in fine roots across diverse species of woody dicot, we evaluated the localization of CT that accumulated in root tissue, and examined its relationships with the stele and cortex tissue in cross-sections of roots in 20 tree species forming different microbial symbiotic groups (ectomycorrhiza and arbuscular mycorrhiza).MethodsIn a cool-temperate forest in Japan, cross-sections of sampled roots in different branching order classes, namely, first order, second to third order, fourth order, and higher than fourth order (higher order), were measured in terms of the length-based ratios of stele diameter and cortex thickness to root diameter. All root samples were then stained with ρ-dimethylaminocinnamaldehyde solution and we determined the ratio of localized CT accumulation area to the root cross-section area (CT ratio).Key ResultsStele ratio tended to increase with increasing root order, whereas cortex ratio either remained unchanged or decreased with increasing order in all species. The CT ratio was significantly positively correlated to the stele ratio and negatively correlated to the cortex ratio in second- to fourth-order roots across species during the shift from primary to secondary root growth. Ectomycorrhiza-associated species mostly had a higher stele ratio and lower cortex ratio than arbuscular mycorrhiza-associated species across root orders. Compared with arbuscular mycorrhiza species, there was greater accumulation of CT in response to changes in the root order of ectomycorrhiza species.ConclusionsDifferent development patterns of the stele, cortex and CT accumulation along the transition from root tip to secondary roots could be distinguished between different mycorrhizal associations. The CT in tissues in different mycorrhizal associations could help with root protection in specific branching orders during shifts in stele and cortex development before and during cork layer formation.     

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.