Carbon input by roots into the soil: Quantification of rhizodeposition from root to ecosystem scale

Despite its fundamental role for carbon (C) and nutrient cycling, rhizodeposition remains ‘the hidden half of the hidden half’: it is highly dynamic and rhizodeposits are rapidly incorporated into microorganisms, soil organic matter, and decomposed to CO₂. Therefore, rhizodeposition is rarely quantified and remains the most uncertain part of the soil C cycle and of C fluxes in terrestrial ecosystems. This review synthesizes and generalizes the literature on C inputs by rhizodeposition under crops and grasslands (281 data sets). The allocation dynamics of assimilated C (after ¹³C‐CO₂ or ¹⁴C‐CO₂ labeling of plants) were quantified within shoots, shoot respiration, roots, net rhizodeposition (i.e., C remaining in soil for longer periods), root‐derived CO₂, and microorganisms. Partitioning of C pools and fluxes were used to extrapolate belowground C inputs via rhizodeposition to ecosystem level. Allocation from shoots to roots reaches a maximum within the first day after C assimilation. Annual crops retained more C (45% of assimilated ¹³C or ¹⁴C) in shoots than grasses (34%), mainly perennials, and allocated 1.5 times less C belowground. For crops, belowground C allocation was maximal during the first 1–2 months of growth and decreased very fast thereafter. For grasses, it peaked after 2–4 months and remained very high within the second year causing much longer allocation periods. Despite higher belowground C allocation by grasses (33%) than crops (21%), its distribution between various belowground pools remains very similar. Hence, the total C allocated belowground depends on the plant species, but its further fate is species independent. This review demonstrates that C partitioning can be used in various approaches, e.g., root sampling, CO₂ flux measurements, to assess rhizodeposits’ pools and fluxes at pot, plot, field and ecosystem scale and so, to close the most uncertain gap of the terrestrial C cycle.     

Cover crop root contributions to soil carbon in a no‐till corn bioenergy cropping system

Crop residues are potential biofuel feedstocks, but residue removal may reduce soil carbon (C). The inclusion of a cover crop in a corn bioenergy system could provide additional biomass, mitigating the negative effects of residue removal by adding to stable soil C pools. In a no‐till continuous corn bioenergy system in the northern US Corn Belt, we used ¹³CO₂ pulse labeling to trace plant C from a winter rye (Secale cereale) cover crop into different soil C pools for 2 years following rye cover crop termination. Corn stover left as residue (30% of total stover) contributed 66, corn roots 57, rye shoots 61, rye roots 50, and rye rhizodeposits 25 g C m^?2 to soil. Five months following cover crop termination, belowground cover crop inputs were three times more likely to remain in soil C pools than were aboveground inputs, and much of the root‐derived C was in mineral‐associated soil fractions. After 2 years, both above‐ and belowground inputs had declined substantially, indicating that the majority of both root and shoot inputs are eventually mineralized. Our results underscore the importance of cover crop roots vs. shoots and the importance of cover crop rhizodeposition (33% of total belowground cover crop C inputs) as a source of soil C. However, the eventual loss of most cover crop C from these soils indicates that cover crops will likely need to be included every year in rotations to accumulate soil C.     

Intercropping effect on root growth and nitrogen uptake at different nitrogen levels

Aims Intercropping legumes and non-legumes may affect the root growth of both components in the mixture, and the non-legume is known to be strongly favored by increasing nitrogen (N) supply. The knowledge of how root systems affect the growth of the individual species is useful for understanding the interactions in intercrops as well as for planning cover cropping strategies. The aim of this work was (i) to determine if different levels of N in the topsoil influence root depth (RD) and intensity of barley and vetch as sole crops or as an intercropped mixture and (ii) to test if the choice of a mixture or the N availability in the topsoil will influence the N uptake by deep roots.Methods In this study, we combined rhizotron studies with root extraction and species identification by microscopy with studies of growth, N uptake and 15 N uptake from deeper soil layers, for studying the root interactions of root growth and N foraging for barley (Hordeum vulgare L.) and vetch (Vicia sativa L.), frequently grown in mixtures as cover crops. N was added at 0 (N0), 50 (N1) and 150 (N2) kg N ha^-1. The roots discrimination relying on the anatomical and morphological differences observed between dicots and monocots proved to be a reliable method providing valuable data for the analysis.Important findings The intercrop and the barley attained slightly higher root intensity (RI) and RD than the vetch, with values around 150 crosses m^-1 and 1.4 m, respectively, compared to 50 crosses m^-1 and 0.9 m for the vetch. At deep soil layers, intercropping showed slightly larger RI values compared to the sole-cropped barley. The barley and the intercropping had larger root length density (RLD) values (200–600 m m ?3) than the vetch (25–130) at 0.8–1.2 m depth. The topsoil N supply did not show a clear effect on the RI, RD or RLD; however, increasing topsoil N favored the proliferation of vetch roots in the intercropping at deep soil layers, with the barley:vetch root ratio ranging from 25 at N0 to 5 at N2. The N uptake of the barley was enhanced in the intercropping at the expense of the vetch (from ~100mg plant^-1 to 200). The intercropped barley roots took up more labeled nitrogen (0.6mg 15 N plant^-1) than the sole-cropped barley roots (0.3mg 15 N plant^-1) from deep layers.     

Root systems and root:mass ratio-carbon allocation under current and projected atmospheric conditions in arable crops

Roots of annual crop plants are a major sink for carbon particularly during early, vegetative growth when up to one-half of all assimilated carbon may be translocated belowground. Flowering marks a particularly important change in resource allocation, especially in determinate species, with considerably less allocation to roots and, depending on environmental conditions, there may be insufficient for maintenance. Studies with ¹⁴C indicate the rapid transfer belowground of assimilates with typically 50% translocated in young cereal plants of which 50% is respired; exudation/rhizodeposition is generally <5% of the fixed carbon. Root: total plant mass decreases through the season and is affected by soil and atmospheric conditions. Limited water availability increased the allocation of ¹³C to roots of wheat grown in columns so that at booting 0.38 of shoot C (ignoring shoot respiration) was belowground compared to 0.31 in well-watered plants. Elevated CO₂ (700 mol CO₂ mol^–1 air) increased the proportion of root:total mass by 55% compared with normal concentration, while increasing the air temperature by a mean of 3 °C decreased the proportion from 0.093 in the cool treatment to 0.055 in the warm treatment.     

Fine root turnover of irrigated hedgerow intercropping in Northern Kenya

Fine root turnover (<2 mm) was determined from repeated measurements of root distribution up to 120 cm soil depth by core sampling in four month intervals. Sole cropped Sorghum bicolor and Acacia saligna were compared with the agroforestry combination in an alley cropping system in semiarid Northern Kenya. Three methods for the calculation of root production were used: the max-min, balancing-transfer and compartment-flow method. The highest root biomass was found in the topsoil for all cropping systems, though trees had a deeper root system. Trees and crops had a similar amount of below-ground biomass during the vegetation period (0.3 and 0.4 Mg DM ha^-1 120 cm^-1), but in the agroforestry combination root biomass was more than the sum of the sole cropped systems (1.1 Mg DM ha^-1 120 cm^-1). The tree system showed a very static root development with little fluctuation between seasons, whereas root biomasses were very dynamic in the crop and tree + crop systems. Root production was highest in the tree + crop combination with 2.1 Mg DM ha^-1 a^-1, with about 50% less in sole cropped trees and crops. Root N input to soil decreased in the order tree + crop>tree>crop system with 13.5, 11.0 and 3.2 kg N ha^-1 a^-1, and cannot be estimated from total below-ground biomass or carbon turnover, as N is accumulated in senescing roots. Such low N input to soil stresses the need for investigating other processes of nutrient input from roots to soil. Areas of highest N input were identified in the topsoil under the tree row in the tree system. Resource utilisation and C and N input to soil were highest with a combination of annual and perennial crops.     

东北三江平原覆盖作物种植效果

以土壤紧实度、冬前生物量、根系性状、植株氮累积量等为指标对供试12种覆盖作物(豆科:紫花苜蓿、光叶苕子、毛叶苕子、红三叶、白三叶、箭筈豌豆;非豆科:苏丹草、青萝卜、Nitro radish、甘蓝型油菜、羽衣甘蓝、菊苣)在东北三江平原地区的种植效果及应用潜力进行综合评价。结果表明: 12种覆盖作物在试验播期均能正常生长,不同覆盖作物与对照相比均有利于降低土壤紧实度,其中青萝卜、Nitro radish和苏丹草土壤紧实度下降最显著,分别较对照下降了47.1%、43.4%和33.4%;覆盖作物群体冬前鲜生物量为3.38～13.98 kg·m^-2,干生物量为0.78～2.43 kg·m^-2,非豆科覆盖作物的生物量显著高于豆科覆盖作物;覆盖作物的群体根系体积以萝卜、油菜、菊苣较大,尤其Nitro radish的根体积高达4018.5 cm³·m^-2,苏丹草的根系横向延展范围最宽;豆科覆盖作物的灰分含量显著低于非豆科覆盖作物,能提供更多易分解的有机物质;覆盖作物总氮积累量为18.72～53.09 g·m^-2,其中,羽衣甘蓝和菊苣的氮积累量最高,且生物量相对较大,有利于氮的积累和固定。在三江平原地区根据主栽作物的类型与冠层结构,选择豆科的三叶草、苕子、紫花苜蓿和非豆科的萝卜、羽衣甘蓝、苏丹草作为覆盖作物进行行间或行内混播的种植方式,可以在调控土壤结构的同时促进养分循环,有利于三江平原黑土地力的提升。     

冬季作物种植对双季稻根系酶活性及形态指标的影响

基于湖南长沙7a定位试验,以冬闲为对照,研究了冬种马铃薯、紫云英及油菜为前茬作物对早、晚稻根系酶活性、形态指标及产量的影响.结果表明,与冬闲相比,冬种作物后早、晚稻根系丙二醛(MDA)含量增加,但其根系的活性氧清除能力更强(SOD、POD和CAT活性高),能够在一定程度上缓解膜脂过氧化作用带来的伤害;冬种不同作物对早晚稻根系形态的影响表现不一.冬种马铃薯和紫云英处理在早稻生育后期的根系优势明显,并能在一定程度上促进晚稻根系生长,双季稻总产量较对照分别增加6.29％和7.76％,而冬种油菜抑制了晚稻根系生长,导致晚稻产量及双季稻总产分别降低6.31％和1.96％;相关性分析表明,灌浆期较高的根长、根数、根体积和根表面积是冬种作物改善双季稻产量的主要原因.综合来看,冬种马铃薯和紫云英对于促进双季稻根系生长,提高稻谷产量具有重要作用,而冬种油菜则不利于提高双季稻的稻谷生产力.     

Spatial and Temporal Variation of Roots, Arbuscular Mycorrhizal Fungi, and Plant and Soil Nutrients in a Mature Pinot Noir (Vitis vinifera L.) Vineyard in Oregon, USA

Soil and crop management practices may influence biomass growth and yields of cotton (Gossypium hirsutum L.) and sorghum (Sorghum bicolorL.) and sequester significant amount of atmospheric CO₂in plant biomass and underlying soil, thereby helping to mitigate the undesirable effects of global warming. This study examined the effects of three tillage practices [no-till (NT), strip till (ST), and chisel till (CT)], four cover crops [legume (hairy vetch) (Vicia villosa roth), nonlegume (rye) (Secale cerealeL), hairy vetch/rye mixture, and winter weeds orno covercrop], and three N fertilization rates (0, 60–65, and 120–130 kg N ha ^–1) on the amount of C sequestered in cotton lint (lint + seed), sorghum grain, their stalks (stems + leaves) and roots, and underlying soil from 2000 to 2002 in central Georgia, USA. A field experiment was conducted on a Dothan sandy loam (fine-loamy, kaolinitic, thermic, Plinthic Kandiudults). In 2000, C accumulation in cotton lint was greater in NT with rye or vetch/rye mixture but in stalks, it was greater in ST with vetch or vetch/rye mixture than in CT with or without cover crops. Similarly, C accumulation in lint was greater in NT with 60 kg N ha ^–1 but in stalks, it was greater in ST with 60 and 120 kg N ha ^–1 than in CT with 0 kg N ha ^–1. In 2001, C accumulation in sorghum grains and stalks was greater in vetch and vetch/rye mixture with or without N rate than in rye without N rate. In 2002, C accumulation in cotton lint was greater in CT with or without N rate but in stalks, it was greater in ST with 60 and 120 kg N ha ^–1 than in NT with or without N rate. Total C accumulation in the above- and belowground biomass in cotton ranged from 1.7 to 5.6 Mg ha ^–1 and in sorghum ranged from 3.4 to 7.2 Mg ha ^–1. Carbon accumulation in cotton and sorghum roots ranged from 1 to 14% of the total C accumulation in above- and belowground biomass. In NT, soil organic C at 0–10 cm depth was greater in vetch with 0 kg N ha ^–1 or in vetch/rye with 120–130 kg N ha ^–1 than in weeds with 0 and 60 kg N ha ^–1 but at 10–30 cm, it was greater in rye with 120–130 kg N ha ^–1 than in weeds with or without rate. In ST, soil organic C at 0–10 cm was greater in rye with 120–130 kg N ha ^–1 than in rye, vetch, vetch/rye and weeds with 0 and 60 kg N ha ^–1. Soil organic C at 0–10 and 10–30 cm was also greater in NT and ST than in CT. Since 5 to 24% of C accumulation in lint and grain were harvested, C sequestered in cotton and sorghum stalks and roots can be significant in the terrestrial ecosystem and can significantly increase C storage in the soil if these residues are left after lint or grain harvest, thereby helping to mitigate the effects of global warming. Conservation tillage, such as ST, with hairy vetch/rye mixture cover crops and 60–65 kg N ha ^–1 can sustain C accumulation in cotton lint and sorghum grain and increase C storage in the surface soil due to increased C input from crop residues and their reduced incorporation into the soil compared with conventional tillage, such as CT, with no cover crop and N fertilization, thereby maintaining crop yields, improving soil quality, and reducing erosion.     

Contribution of above‐ and belowground bioenergy crop residues to soil carbon

GHG mitigation by bioenergy crops depends on crop type, management practices, and the input of residue carbon (C) to the soil. Perennial grasses may increase soil C compared to annual crops because of more extensive root systems, but it is less clear how much soil C is derived from above‐ vs. belowground inputs. The objective of this study was to synthesize the existing knowledge regarding soil C inputs from above‐ and belowground crop residues in regions cultivated with sugarcane, corn, and miscanthus, and to predict the impact of residue removal and tillage on soil C stocks. The literature review showed that aboveground inputs to soil C (to 1‐m depth) ranged from 70% to 81% for sugarcane and corn vs. 40% for miscanthus. Modeled aboveground C inputs (to 30 cm depth) ranged from 54% to 82% for sugarcane, but were 67% for miscanthus. Because 50% of observed miscanthus belowground biomass is below 30 cm depth, it may be necessary to increase the depth of modeled soil C dynamics to reconcile modeled belowground C inputs with measured. Modeled removal of aboveground corn residue (25–100%) resulted in C stock reduction in areas of corn–corn–soybean rotation under conventional tillage, while no‐till management lessoned this impact. In sugarcane, soil C stocks were reduced when total aboveground residue was removed at one site, while partial removal of sugarcane residue did not reduce soil C stocks in either area. This study suggests that aboveground crop residues were the main C‐residue source to the soil in the current bioethanol sector (corn and sugarcane) and the indiscriminate removal of crop residues to produce cellulosic biofuels can reduce soil C stocks and reduce the environmental benefits of bioenergy. Moreover, a switch to feedstocks such as miscanthus with more allocation to belowground C could increase soil C stocks at a much faster rate.     

Estimation of rhizodeposition at field scale: upscaling of a 14C labeling study

Background and aims

Rhizodeposition of plants is the most uncertain component of the carbon (C) cycle. By existing approaches the amount of rhizodeposition can only roughly be estimated since its persistence in soil is very short compared to other organic C pools. We suggest an approach to quantify rhizodeposition at the field scale by assuming a constant ratio between rhizodeposited-C to root-C.

Methods

Maize plants were pulse-labeled with ¹⁴CO₂ under controlled conditions and the soil ¹⁴CO₂ efflux was separated into root and rhizomicrobial respiration. The latter and the ¹⁴C activity remaining in the soil corresponded to total rhizodeposition. By relating rhizodeposited-¹⁴C to root-¹⁴C a rhizodeposition-to-root ratio of 0.56 was calculated. This ratio was applied to the root biomass C measured in the field to estimate rhizodeposition under field conditions.

Results

Maize allocated 298 kg C ha^?1 as root-C and 166 kg C ha^?1 as rhizodeposited-C belowground, 50 % of which were recovered in the upper 10 cm. The fate of rhizodeposits was estimated based on the ¹⁴C data, which showed that 62 % of total rhizodeposition was mineralized within 16 days, 7 % and 0.3 % was incorporated into microbial biomass and DOC, respectively, and 31 % was recovered in the soil.

Conclusions

We conclude that the present approach allows for an improved estimation of total rhizodeposition, since it accounts not only for the fraction of rhizodeposits remaining in soil, but also for that decomposed by microorganisms and released from the soil as CO₂.     

How does nitrogen availability alter rhizodeposition in Lolium multiflorum Lam. during vegetative growth?

The objective of this work was to determine if the impact of nitrogen (N) on the release of organic carbon (C) into the soil by roots (rhizodeposition) correlated with the effect of this nutrient on some variables of plant growth. Lolium multiflorum Lam. was grown at two levels of N supply, either in sterile sand percolated with nutrient solution or in non-sterile soil. The axenic sand systems allowed continuous quantification of rhizodeposition and accurate analysis of root morphology whilst the soil microcosms allowed the study of ¹⁴C labelled C flows in physico-chemical and biological conditions relevant to natural soils. In the axenic sand cultures, enhanced N supply strongly increased the plant biomass, the plant N content and the shoot to root ratio. N supply altered the root morphology by increasing the root surface area and the density of apices, both being significantly positively correlated with the rate of organic C release by plant roots before sampling. This observation is consistent with the production of mucilage by root tips and with mechanisms of root exudation reported previously in the literature, i.e. the passive diffusion of roots solutes along the root with increased rate behind the root apex. We proposed a model of root net exudation, based on the number of root apices and on root soluble C that explained 60% of the variability in the rate of C release from roots at harvest. The effects of N on plant growth were less marked in soil, probably related to the relatively high supply of N from non-fertiliser soil-sources. N fertilization increased the shoot N concentration of the plants and the shoot to root ratio. Increased N supply decreased the partitioning of ¹⁴C to roots. In parallel, N fertilisation increased the root soluble ¹⁴C and the ¹⁴C recovered in the soil per unit of root biomass, suggesting a stimulation of root exudation by N supply. However, due to the high concentration of N in our unfertilised plants, this stimulation was assumed to be very weak because no significant effect of N was observed on the microbial C and on the bacterial abundance in the rhizosphere. Considering the difficulties in evaluating rhizodeposition in non sterile soil, it is suggested that the root soluble C, the root surface area and the root apex density are additional relevant variables that should be useful to measure along with the variables that are commonly determined when investigating how plant functioning impacts on the release of C by roots (i.e soil C, C of the microbial biomass, rhizosphere respiration).     

Induced carbon reallocation and compensatory growth as root herbivore tolerance mechanisms

Upon attack by leaf herbivores, many plants reallocate photoassimilates below ground. However, little is known about how plants respond when the roots themselves come under attack. We investigated induced resource allocation in maize plants that are infested by the larvae Western corn rootworm Diabrotica virgifera virgifera. Using radioactive ¹¹CO₂, we demonstrate that root‐attacked maize plants allocate more new ¹¹C carbon from source leaves to stems, but not to roots. Reduced meristematic activity and reduced invertase activity in attacked maize root systems are identified as possible drivers of this shoot reallocation response. The increased allocation of photoassimilates to stems is shown to be associated with a marked thickening of these tissues and increased growth of stem‐borne crown roots. A strong quantitative correlation between stem thickness and root regrowth across different watering levels suggests that retaining photoassimilates in the shoots may help root‐attacked plants to compensate for the loss of belowground tissues. Taken together, our results indicate that induced tolerance may be an important strategy of plants to withstand belowground attack. Furthermore, root herbivore‐induced carbon reallocation needs to be taken into account when studying plant‐mediated interactions between herbivores.     

Effects of experimental nitrogen deposition on soil organic carbon storage in Southern California drylands

Atmospheric nitrogen (N) deposition is enriching soils with N across biomes. Soil N enrichment can increase plant productivity and affect microbial activity, thereby increasing soil organic carbon (SOC), but such responses vary across biomes. Drylands cover ~45% of Earth's land area and store ~33% of global SOC contained in the top 1 m of soil. Nitrogen fertilization could, therefore, disproportionately impact carbon (C) cycling, yet whether dryland SOC storage increases with N remains unclear. To understand how N enrichment may change SOC storage, we separated SOC into plant-derived, particulate organic C (POC), and largely microbially derived, mineral-associated organic C (MAOC) at four N deposition experimental sites in Southern California. Theory suggests that N enrichment increases the efficiency by which microbes build MAOC (C stabilization efficiency) if soil pH stays constant. But if soils acidify, a common response to N enrichment, then microbial biomass and enzymatic organic matter decay may decrease, increasing POC but not MAOC. We found that N enrichment had no effect on C fractions except for a decrease in MAOC at one site. Specifically, despite reported increases in plant biomass in three sites and decreases in microbial biomass and extracellular enzyme activities in two sites that acidified, POC did not increase. Furthermore, microbial C use and stabilization efficiency increased in a non-acidified site, but without increasing MAOC. Instead, MAOC decreased by 16% at one of the sites that acidified, likely because it lost 47% of the exchangeable calcium (Ca) relative to controls. Indeed, MAOC was strongly and positively affected by Ca, which directly and, through its positive effect on microbial biomass, explained 58% of variation in MAOC. Long-term effects of N fertilization on dryland SOC storage appear abiotic in nature, such that drylands where Ca-stabilization of SOC is prevalent and soils acidify, are most at risk for significant C loss.     

Root characteristics of selected field crops: Data from the Wageningen Rhizolab (1990–2002)

Since being built in 1990, the rhizotron facility in Wageningen, the Wageningen Rhizolab, has been used for experiments on crops (e.g. Alfalfa, Brussels sprouts, common velvet grass, field bean, fodder radish, leeks, lupins, maize, potato, beetroot, ryegrass, spinach, spring wheat, winter rye and winter wheat). In the experiments, horizontal glass minirhizotron tubes combined with auger sampling were used to assess rooting characteristics. For this paper we took the root data from these experiments and looked for a general relationship between thermal time/time after planting and rooting depth, the velocity of the root front and root proliferation. For certain depths (fixed by the depth at which the horizontal minirhizotrons were installed) a simple linear regression was established between the average root number per cm² minirhizotron surface area and thermal time after planting. The compartments selected for each crop were those in which there had been a control treatment and/or in which conditions for rooting were considered to be optimal. We performed regression analyses per compartment and per depth, but only for the period after planting in which a linear increase of root numbers vs. thermal time was observed. After averaging the results, the regression procedure yielded two parameters of rooting for each crop: (a) the actual or thermal time at which the first root appeared at a certain depth and (b) the root proliferation rate after the first root had appeared. In this way, inherent crop differences in rooting behaviour (rooting depth and root proliferation) became apparent. For each crop, the velocity of the root front after planting could be established (calculated in cm(°C day)^–1). This parameter differed greatly between crops. Some crops (such as leeks and common velvet grass) explored the soil profile slowly: the root front moved at a velocity of only 0.07cm(°C day)^–1. Among the crops whose roots grew down much faster (0.18–0.26cm (°C day)^–1) were cereals and fodder radish. For a day with an average temperature of 15°C these rates would have corresponded with the root front travelling approximately 1–4cm per day. In the crops studied the root front velocity did not correlate with the root proliferation rate.     

Partitioning of belowground C in young sugar maple forest

Background and aims

Trees allocate a high proportion of assimilated carbon belowground, but the partitioning of that C among ecosystem components is poorly understood thereby limiting our ability to predict responses of forest C dynamics to global change drivers.

Methods

We labeled sugar maple saplings in natural forest with a pulse of photosynthetic ¹³C in late summer and traced the pulse over the following 3 years. We quantified the fate of belowground carbon by measuring ¹³C enrichment of roots, rhizosphere soil, soil respiration, soil aggregates and microbial biomass.

Results

The pulse of ¹³C contributed strongly to root and rhizosphere respiration for over a year, and respiration comprised about 75 % of total belowground C allocation (TBCA) in the first year. We estimate that rhizosphere carbon flux (RCF) during the dormant season comprises at least 6 % of TBCA. After 3 years, 3.8 % of the C allocated belowground was recovered in soil organic matter, mostly in water-stable aggregates.

Conclusions

A pulse of carbon allocated belowground in temperate forest supplies root respiration, root growth and RCF throughout the following year and a small proportion becomes stabilized in soil aggregates.     

Vertical and horizontal development of the root system of carrots following green manure

Cover crops grown as green manure or for other purposes will affect nitrogen (N) distribution in the soil, and may thereby alter root growth of a succeeding crop. During two years, experiments were performed to study effects of nitrogen supply by green manure on root development of carrots (Daucus carota L). Total root intensity (roots cm⁻² on minirhizotrons) was significantly affected by the green manures, and was highest in the control plots where no green manure had been grown. Spread of the root system into the interrow soil was also affected by green manure treatments, as the spread was reduced where spring topsoil N_min was high. Although N supply and distribution in the soil profile differed strongly among the treatments, no effect was observed on the rooting depth of the carrot crops. Across all treatments the rooting front penetrated at a rate of 0.82 and 0.68 mm day⁻¹ °C⁻¹ beneath the crop rows and in the interrow soil, respectively. The minirhizotrons only allowed measurements down to 1 m, and the roots reached this depth before harvest. Extrapolating the linear relationship between temperature sum and rooting depth until harvest would lead to rooting depths of 1.59 and 1.18 m under the crop rows and in the interrow soil respectively. Soil analysis showed that the carrot crop was able to reduce N_min to very low levels even in the 0.75 to 1.0 m soil layer, which is in accordance with the root measurements. Still, where well supplied, the carrots left up 90 kg N ha⁻¹ in the soil at harvest. This seemed to be related to a limited N uptake capacity of the carrots rather than to insufficient root growth in the top metre of the soil. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

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]     

Nitrogen increases soil organic carbon accrual and alters its functionality

Nitrogen (N) availability has been considered as a critical factor for the cycling and storage of soil organic carbon (SOC), but effects of N enrichment on the SOC pool appear highly variable. Given the complex nature of the SOC pool, recent frameworks suggest that separating this pool into different functional components, for example, particulate organic carbon (POC) and mineral-associated organic carbon (MAOC), is of great importance for understanding and predicting SOC dynamics. Importantly, little is known about how these N-induced changes in SOC components (e.g., changes in the ratios among these fractions) would affect the functionality of the SOC pool, given the differences in nutrient density, resistance to disturbance, and turnover time between POC and MAOC pool. Here, we conducted a global meta-analysis of 803 paired observations from 98 published studies to assess the effect of N addition on these SOC components, and the ratios among these fractions. We found that N addition, on average, significantly increased POC and MAOC pools by 16.4% and 3.7%, respectively. In contrast, both the ratios of MAOC to SOC and MAOC to POC were remarkably decreased by N enrichment (4.1% and 10.1%, respectively). Increases in the POC pool were positively correlated with changes in aboveground plant biomass and with hydrolytic enzymes. However, the positive responses of MAOC to N enrichment were correlated with increases in microbial biomass. Our results suggest that although reactive N deposition could facilitate soil C sequestration to some extent, it might decrease the nutrient density, turnover time, and resistance to disturbance of the SOC pool. Our study provides mechanistic insights into the effects of N enrichment on the SOC pool and its functionality at global scale, which is pivotal for understanding soil C dynamics especially in future scenarios with more frequent and severe perturbations.     

Soil aggregate sequestration of cover crop root and shoot-derived nitrogen

Cover crop roots and shoots release carbon (C) and nitrogen (N) compounds in situ during their decomposition. Depending upon the season, these C and N compounds may be sequestered, the C may be respired or the N may be leached below the root zone. A field study was established to identify the contributions of cover crop root and shoot N to different regions within aggregates in the A_p horizon of a Kalamazoo loam soil. Fall-planted rye plants (Secale cerealeL.) were labeled the next May with foliar applications of solutions containing 99% atom (¹⁵NH₄)₂SO₄. Isotopic enrichment of soil aggregates ranging from 2.0 to 4.0, 4.0–6.3 and 6.3–9.5 mm across was determined following plant residue applications. Concentric layers of aggregates were removed from each aggregate by newly designed meso soil aggregate erosion (SAE) chambers. Non-uniform distributions of total N and recently derived rye N in soil macroaggregates, across time, suggested that the formations and functions of macroaggregates are very dynamics processes and soil aggregates influence where N is deposited. Early in the season, more ¹⁵N migrated to the interior regions of the smallest aggregates, 2–4 mm across, but it was limited to only surfaces and transitional regions of the larger aggregates, 6.3–9.3 mm across. Exterior layers of aggregates between 6.0 and 9.5 mm retained 1.6% of the N_{derived from roots} in July 1999, which was three times more than their interior regions. This was slightly greater than the % N_{derived from shoot.} One month later, as the maize root absorption of N increased rapidly, % N_{derived from roots} and % N_{derived from shoot} were nearly equal in exterior layers and interior regions of soil aggregates. This equilibrium distribution may have been from either greater diffusion of N within the aggregates and/or maize root removal form aggregate exteriors. Results supported that most of roots grew preferentially around surfaces of soil aggregates rather than through aggregates. Cover crop roots contributed as much N as cover crop shoots to the total soil N pool. Subsequent crops use N from the most easily accessible zones of soil structure, which are surfaces of larger soil aggregates. Therefore maintaining active plant roots and aggregated soil structure in the soil enhances N sequestration and maximize soil N availability. These studies suggest that the rapid and perhaps bulk flow of soil N solutions may bypass many of the central regions of soil aggregates, resulting in greater leaching losses.