Carbon Turnover in the Rhizosphere

Considerable progress has been made during the last decade towards understanding and quantifying the input and turnover of plant carbon in the rhizosphere. This was made possible by the development (partially by the authors) and combination of appropriate new methods, such as:

• –homogeneous labelling of whole plants with ¹⁴C
• –distinction between root and microbial respiration
• –separation of soil zones of known distances from the roots
• –determination of microbial soil biomass.

These methods were applied to study the following aspects:

• –release of organic plant carbon into the soil by growing roots
• –utilization of this plant carbon by the microbial biomass in the rhizosphere
• –related influence on the turnover of soil organic matter, and
• –spatial range of such root influence in the soil.

About 19% of the total photosynthetic production of the investigated plants was released into the rhizosphere as organic material. Most of this (15%) was transformed by the rhizosphere microorganisms into CO₂, while only a small fraction (4%) remained in the soil, mainly as microbial cells (2.5%). As a result, microbial rhizosphere biomass increased considerably. Relative to the organic C-input, however, the incorporation of root derived carbon by the microbial biomass was remarkably low (13%). Along with the increase in microbial rhizosphere biomass, the presence of plant roots also enhanced the decomposition of soil organic matter and affected soil aggregate stability. Root carbon and root influences were even detected up to 20 mm away from the roots. This may be partially attributed to the contribution of root derived volatiles. Accordingly, both the actual volume of the rhizosphere and its metabolic significance is greater than what has so far been assumed. Possible interactions involving root, soil and microbial carbon are discussed.     

Budgets for root-derived C and litter-derived C: comparison between conventional tillage and no tillage soils

Placement of plant residues in conventional tillage (CT) and no-tillage (NT) soils affects organic matter accumulation and the organization of the associated soil food webs. Root-derived C inputs can be considerable and may also influence soil organic matter dynamics and soil food web organization. In order to differentiate and quantify C contributions from either roots or litter in CT and NT soils, a ¹⁴C tracer method was used.To follow root-derived C, maize plants growing in the field were ¹⁴C pulse-labeled, while the plant litter in those plots remained unlabeled. The ¹⁴C was measured in NT and CT soils for the different C pools (shoots, roots, soil, soil respiration, microbial biomass). Litter-derived C was followed by applying ¹⁴C labeled maize litter to plots which had previously grown unlabeled maize plants. The ¹⁴C pools measured for the litter-derived CT and NT plots included organic matter, microbial biomass, soil respiration, and soil organic C.Of the applied label in the root-derived C plots, 35–55, 6–8, 3, 1.6, and 0.4–2.4% was recovered in the shoots, roots, soil, cumulative soil respiration, and microbial biomass, respectively. The ¹⁴C recovered in these pools did not differ between CT and NT treatments, supporting the hypothesis that the rhizosphere microbial biomass in NT and CT may be similar in utilization of root-derived C. Root exudates were estimated to be 8–13% of the applied label. In litter-derived C plots, the percentage of applied label recovered in the particulate organic matter (3.2–82%), microbial biomass (4–6%), or cumulative soil respiration (12.5–14.7%) was the same for CT and NT soils. But the percentage of ¹⁴C recovered in CT soil organic C (18–69%) was higher than that in NT (12–43%), suggesting that particulate organic matter (POM) leaching and decomposition occurred at a higher rate in CT than in NT. Results indicate faster turnover of litter-derived C in the CT plots.     

Glucose uptake by maize roots and its transformation in the rhizosphere

The flow of carbon from roots into the rhizosphere represents a significant C loss from plants. However, roots have the capacity to recapture low molecular weight C from soil although this is in direct competition with soil microorganisms. The aim of this study was to investigate the behaviour of glucose in rhizosphere and non-rhizosphere soil, the plant's potential to recapture sugars from soil and translocation and utilization of the recaptured sugars. In microcosms containing maize plants we injected ¹⁴C-glucose into the rhizosphere and followed its uptake into plants, upward and downward transport in the plant and soil, evolution as ¹⁴CO₂ and incorporation into the soil microbial biomass. These fluxes were compared with non-rhizosphere soil. Glucose was rapidly mineralized in soil and the rate of turnover was significantly greater in the rhizosphere in comparison to non-rhizosphere soil. The amount of glucose captured by the maize plants was low (<10% of the total ¹⁴C-glucose added) in comparison to that captured by the soil microbial biomass. Only small amounts of the ¹⁴C-glucose were transported to the shoot (0.6% of the total). The degree of glucose capture by maize roots whilst in competition with soil microorganisms was similar to similar experiments performed for amino acids. We conclude that while plant roots can recapture low molecular weight C from the rhizosphere, intense competition from soil microorganisms may reduce the efficiency of this process.     

Influence of the rhizosphere on microbial biomass and recently formed organic matter

We have aimed to quantify the effect of roots on the size of the soil microbial biomass, and their influence on the turnover of soil organic matter and on the extent of the rhizosphere. We sampled a sandy clay loam topsoil from two subplots with different treatment histories. One had a normal arable fertilization record, the other had received only inorganic nitrogen fertilizer but no phosphorus and potassium for 30 years. Glucose labelled with ¹⁴C was added to both samples which were then incubated for 4 weeks before the soil was packed in cylinders and planted with ryegrass. In both soils, microbial biomass at the root surface doubled during the first 8 days. At day 15, the microbial biomass had further increased in the fertile soil, and the rhizosphere effect had extended 2.5 mm into the fertile soil, but to only 1 mm in the infertile soil. The microbial ¹⁴C increased threefold near the roots in the fertile soil as a result of assimilation of previously formed microbial residues, but in the infertile soil there was no increase. There was a close relation between the increase in microbial ¹⁴C and a decrease in ¹⁴C soluble in 2 m KCl, indicating that the microbial residues were more weakly adsorbed in the fertile soil. We conclude that the increased microbial population living near the root surfaces re‐assimilated part of the ¹⁴C‐labelled microbial residues in the fertile soil. In the infertile soil, microbial residues resisted decomposition because they were more strongly sorbed on to soil surfaces.     

Mobilization and turnover of soil phosphorus in the rhizosphere

Recent progress in methods enables a better understanding of the turnover of P in the rhizosphere. Examples of this progress are the separation of soil layers differing in proximity to the roots, improved methods for extraction and fractionation of soil P, application of ³²P isotope dilution analysis to follow P fluxes between various fractions and direct determination of microbially bound P and of root phosphatases.

• These methods were combined to investigate the following aspects
• –labile P pools, the P fluxes between these pools and their contribution to the P supply to growing maize roots
• –the role of microbial biomass in these interactions and the partition of mobilized P between plants and microorganisms
• –modifications of sorption and transport of P in the rhizosphere
• –plant availability of native and added organic phosphates, and the relative significance of root and soil phosphatases.

There is a significant transformation of P in the rhizosphere with a corresponding redistribution among fractions of different plant availability. About 9% of the inorganic ³²P added to soil were incorporated within 2 weeks into microbial and organic fractions. The transfer of P from non-exchangeable forms exceeded the depletion of the exchangeable P by a factor of 5. About 53% of the mobilized P originated from inorganic, the remaining 47% from organic fractions. Of the mobilized P 80% was taken up by the plants and 20% was found in the microbial biomass. Up to 90% of the P in the rhizosphere soil solution was organic with a maximum just outside the root zone. Soluble inositol hexaphosphate modified the sorption of inorganic P, thus shifting its equilibrium solution concentration. The phosphatase activity of the roots is considerable. Both root phosphatase activity and the utilization of inositol hexaphosphate depend on the P supply and nutritional status of plants with regard to P. It is concluded that the rhizosphere is a key site of P transformation with a significant mobilization of P from the non-exchangeable inorganic and organic fractions. Organic P fractions not only play a significant role as a P source but also modify important soil parameters related to the sorption and transport of P in the rhizosphere.     

Interaction between rhizosphere microorganisms and plant roots: 13C fluxes in the rhizosphere after pulse labeling

The input dynamics of labeled C into pools of soil organic matter and CO₂ fluxes from soil were studied in a pot experiment with the pulse labeling of oats and corn under a ¹³CO₂ atmosphere, and the contribution of the root and microbial respiration to the emission of CO₂ from the soil was determined from the fluxes of labeled C in the microbial biomass and the evolved carbon dioxide. A considerable amount of ¹³C (up to 96% of the total amount of the label found in the rhizosphere soil) was incorporated into the biomass of the rhizosphere microorganisms. The diurnal fluctuations of the labeled C pools in the microbial biomass, dissolved organic carbon, and CO₂ released in the rhizosphere of oats and corn were related to the day/night changes, i.e., to the on and off periods of the photosynthetic activity of the plants. The average contribution of the corn root respiration (70% of the total CO₂ emission from the soil surface) was higher than that of the oats roots (44%), which was related to the lower incorporation of rhizodeposit carbon into the microbial biomass in the soil under the corn plants than in the soil under the oats plants.     

水稻根际和周围土壤中微生物功能基因丰度与碳、氮状况及其通量之间的关系

Rapid nitrogen（N） transformations and losses occur in the rice rhizosphere through root uptake and microbial activities. However,the relationships between rice roots and rhizosphere microbes for N utilization are still unclear. We analyzed different N forms（NH＋4,NO-3, and dissolved organic N）, microbial biomass N and C, dissolved organic C, CH4 and N2O emissions, and abundance of microbial functional genes in both rhizosphere and bulk soils after 37-d rice growth in a greenhouse pot experiment. Results showed that the dissolved organic C was significantly higher in the rhizosphere soil than in the non-rhizosphere bulk soil, but microbial biomass C showed no significant difference. The concentrations of NH＋4, dissolved organic N, and microbial biomass N in the rhizosphere soil were significantly lower than those of the bulk soil, whereas NO-3in the rhizosphere soil was comparable to that in the bulk soil. The CH4 and N2O fluxes from the rhizosphere soil were much higher than those from the bulk soil. Real-time polymerase chain reaction analysis showed that the abundance of seven selected genes, bacterial and archaeal 16 S rRNA genes, amoA genes of ammonia-oxidizing archaea and ammonia-oxidizing bacteria, nosZ gene, mcrA gene, and pmoA gene, was lower in the rhizosphere soil than in the bulk soil, which is contrary to the results of previous studies. The lower concentration of N in the rhizosphere soil indicated that the competition for N in the rhizosphere soil was very strong, thus having a negative effect on the numbers of microbes. We concluded that when N was limiting, the growth of rhizosphere microorganisms depended on their competitive abilities with rice roots for N.     

Microbial immobilisation of C rhizodeposits in rhizosphere and root-free soil under continuous C labelling of oats

A greenhouse experiment was conducted by growing oats (Avenasativa L.) in a continuously ¹³CO₂ labeled atmosphere. The allocation of ¹³C-labeled photosynthates in plants, microbial biomass in rhizosphere and root-free soil, pools of soil organic C, and CO₂ emissions were examined over the plant's life cycle. To isolate rhizosphere from root-free soil, plant seedlings were placed into bags made of nylon monofilament screen tissue (16 μm mesh) filled with soil. Two peaks of ¹³C in rhizosphere pools of microbial biomass and dissolved organic carbon (DOC), as well as in CO₂ emissions at the earing and ripeness stages were revealed. These ¹³C maxima corresponded to: (i) the end of rapid root growth and (ii) beginning of root decomposition, respectively. The δ¹³C values of microbial biomass were higher than those of DOC and of soil organic matter (SOM). The microbial biomass C accounted for up to 56 and 39% of ¹³C recovered in the rhizosphere and root-free soil, respectively. Between 4 and 28% of ¹³C assimilated was recovered in the root-free soil. Depending on the phenological stage, the contribution of root-derived C to total CO₂ emission from soil varied from 61 to 92% of total CO₂ evolved, including 4-23% attributed to rhizomicrobial respiration. While 81-91% of C substrates used for microbial growth in the root-free soil and rhizosphere came from SOM, the remaining 9-19% of C substrates utilized by the microbial biomass was attributable to rhizodeposition. The use of continuous isotopic labelling and physical separation of root-free and rhizosphere soil, combined with natural ¹³C abundance were effective in gaining new insight on soil and rhizosphere C-cycling.     

Turnover of root-derived material and related microbial biomass formation in soils of different texture

Wheat plants were grown on two soils of different texture, a sandy soil and a silty clay loam, in an atmosphere containing ¹⁴CO₂. The ¹⁴C and total C content of the shoots, roots, soil rhizosphere CO₂ and soil microbial biomass were measured 21, 28, 35 and 42 days after germination. There was a pronounced effect of soil texture on the turnover of root-derived C through the microbial biomass. Turnover was relatively fast and at a constant rate in the sandy soil but slowed down in the clay soil, following an initial high assimilation of root products into the microbial biomass.Four percent of the total fixed ¹⁴C was retained in the clay loam after 6 weeks compared with a corresponding value of 1.2% for the sandy soil. The proportion of fixed ¹⁴C recovered as rhizosphere CO₂ at each of the sampling times was relatively constant for the sandy soil (ca 19%) but decreased from 17% at day 28 to 11% at day 42 in the clay soil. The proportion of total fixed ¹⁴C in the soil biomass as measured by a fumigation technique increased to a maximum value of 20% after 6 weeks in the sandy soil but decreased in the clay soil from 86% at day 21 to 26% after 42 days plant growth.     

Rhizodeposition: Its contribution to microbial growth and carbon and nitrogen turnover within the rhizosphere

A greenhouse rhizobox experiment was carried out to investigate the fate and turnover of ¹³C‐ and ¹⁵N‐labeled rhizodeposits within a rhizosphere gradient from 0–8 mm distance to the roots of wheat. Rhizosphere soil layers from 0–1, 1–2, 2–3, 3–4, 4–6, and 6–8 mm distance to separated roots were investigated in an incubation experiment (42 d, 15°C) for changes in total C and N and that derived from rhizodeposition in total soil, in soil microbial biomass, and in the 0.05 M K₂SO₄–extractable soil fraction. CO₂‐C respiration in total and that derived from rhizodeposition were measured from the incubated rhizosphere soil samples. Rhizodeposition C was detected in rhizosphere soil up to 4–6 mm distance from the separated roots. Rhizodeposition N was only detected in the rhizosphere soils up to 3–4 mm distance from the roots. Microbial biomass C and N was increased with increasing proximity to the separated roots. Beside ¹³C and ¹⁵N derived from rhizodeposits, unlabeled soil C and N (native SOM) were incorporated into the growing microbial biomass towards the roots, indicating a distinct acceleration of soil organic matter (SOM) decomposition and N immobilization into the growing microbial biomass, even under the competition of plant growth. During the soil incubation, microbial biomass C and N decreased in all samples. Any decrease in microbial biomass C and N in the incubated rhizosphere soil layers is attributed mainly to a decrease of unlabeled (native) C and N, whereas the main portion of previously incorporated rhizodeposition C and N during the plant growth period remained immobilized in the microbial biomass during the incubation. Mineralization of native SOM C and N was enhanced within the entire investigated rhizosphere gradient. The results indicate complex interactions between substrate input derived from rhizodeposition, microbial growth, and accelerated C and N turnover, including the decomposition of native SOM (i.e., rhizosphere priming effects) at a high spatial resolution from the roots.     

Production of root-derived material and associated microbial growth in soil at different nutrient levels

Summary Maize plants were grown for 42 days in a sandy soil at two different mineral nutrient levels, in an atmosphere containing ¹⁴CO₂. The ¹⁴C and total carbon contents of shoots, roots, soil and soil microbial biomass were measured 28, 35 and 42 days after germination. Relative growth rates of shoots and roots decreased after 35 days at the lower nutrient level, but were relatively constant at the higher nutrient level. In the former treatment, 2% of the total ¹⁴C fixed was retained as a residue in soil at all harvests while at the higher nutrient level up to 4% was retained after 42 days. Incorporation of ¹⁴C into the soil microbial biomass was close to its maximum after 35 days at the lower nutrient level, but continued to increase at the higher level. Generally a good agreement existed between microbial biomass, ¹⁴C contents and numbers of fluorescent pseudomonads in the rhizosphere. Numbers of fluorescent pseudomonads in the rhizosphere were maximal after 35 days at the lower nutrient level and continued to increase at the higher nutrient level. The proportions of the residual ¹⁴C in soil, incorporated in the soil microbial biomass, were 28% to 41% at the lower nutrient level and 20%6 – 30% at the higher nutrient level. From the lower nutrient soil 18%6 – 52%6 of the residual soil ¹⁴C could be extracted with 0.5 N K₂SO₄, versus 14%6 – 16% from the higher nutrient soil.Microbial growth in the rhizosphere seemed directly affected by the depletion of mineral nutrients while plant growth and the related production of root-derived materials continued.     

Microbial activity in the rhizosphere soil of six herbaceous species cultivated in a greenhouse is correlated with shoot biomass and root C concentrations

The stimulation of rhizosphere microorganisms by exudates released from roots is important for nutrient cycling and differs between plant species. The reasons for this between-species variability are poorly understood. We studied correlations between shoot biomass, soluble and non-soluble root C concentrations and rhizosphere bacterial abundance (CFU: colony forming units) and an index of microbial activity (in vitro utilization of [U-¹⁴C]glucose by soil microorganisms). We studied Briza media and Rumex acetosella (nutrient-poor habitats), Epilobium hirsutum, Eupatorium cannabinum, Rumex obtusifolius and Urtica dioica (nutrient rich habitats) cultivated in a greenhouse for 5 weeks in a forest soil. We found significant differences among species for the bacterial abundance and microbial activity in the rhizosphere. These differences poorly reflected the nutrient richness of the common habitats for these species, possibly because the soil conditions were not optimal. Nevertheless, microbial activity was positively correlated with root soluble C concentration and shoot biomass and negatively correlated with the concentration of non-soluble C in roots. These preliminary results suggest that the carbon economy could be an important control of the between-species variability of microbial activity in the rhizosphere.     

Fate of C- and N-labelled rhizodeposition of Lolium perenne as function of the distance to the root surface

A greenhouse rhizobox experiment was carried out to quantify the incorporation of ¹³C- and ¹⁵N-labelled rhizodeposits into different soil pools, especially into the rhizosphere microbial biomass, with increasing distances to the root surface of Lolium perenne. Five layers were analysed over 0-4.2 mm distance to an artificial root surface. C and N derived from rhizodeposition were 4.2% of total C and 2.8% of total N in soil at 0-1.0 mm distance and decreased rapidly with increasing distance. Microbial biomass C and N increased significantly towards the roots. At 0-1.0 mm distance microbial biomass C and N accounted for 66% and 29% of C and N derived from rhizodeposition, respectively. These percentages declined with increasing distance to the roots, but were still traceable up to 4.2 mm distance. Only small amounts of root released C and N were found in the 0.05 M K₂SO₄-extractable fraction. Extractable C and N derived from rhizodeposition varied around means of 4% of total C and N derived from rhizodeposition and increased only marginally with increasing distance to the roots. C derived from rhizodeposition in the non-extractable soil organic matter increased from 65 to 89% of total C derived from rhizodeposition at 0-3.4 mm distance. Conversely, microbial biomass C derived from rhizodeposition decreased from 33 to 4%. N derived from rhizodeposition in the non-extractable soil organic matter increased from 61 to 79% of total N derived from rhizodeposition at 0-2.6 mm distance, followed by a decline to roughly 55% in the two outer layers. Microbial biomass N decreased from 37 to 16% at 0-2.6 mm distance, followed by an increase to roughly 41% in the two outer layers. The C/N ratio of total C and N derived from rhizodeposition as well as that of extractable C and N derived from rhizodeposition increased with increasing distance to the roots to values above 30. In contrast, the C/N ratio of incorporated rhizodeposition C and N into the microbial biomass decreased to values less than 5 at 2.6-4.2 mm distance. The data indicate differential microbial response to C and N derived from rhizodeposition at a high spatial resolution from the root surface. The turnover of C and N derived from rhizodeposition in the rhizosphere as a function of the distance to the root surface is discussed.     

Relationships between microbial indices in roots and silt loam soils forming a gradient in soil organic matter

Perennial rye grass (Lolium perenne) was grown in a greenhouse pot experiment on seven soils to answer the question whether the microbial colonisation of roots is related to existing differences in soil microbial indices. The soils were similar in texture, but differed considerably in soil organic matter, microbial biomass, and microbial community structure. Ergosterol and fungal glucosamine were significantly interrelated in the root material. This ergosterol was also significantly correlated with the average ergosterol content of bulk and rhizosphere soil. In addition, the sum of fungal C and bacterial C in the root material revealed a significant linear relationship with microbial biomass C in soil. The colonisation of roots with microorganisms increased apparently with an increase in soil microbial biomass. In the root material, microbial tissue consisted of 77% fungi and 23% bacteria. In soil, the fungal dominance was slightly, but significantly lower, with 70% fungi and 30% bacteria. Fungal glucosamine in the root material was significantly correlated with that in soil (r=0.65). This indicates a close relationship between the composition of dead microbial remains in soil and the living fraction in soil and root material for unknown reasons.     

Effects of pine roots on microorganisms,fauna, and nitrogen availability in two soil horizons of a coniferous forest spodosol

Summary We investigated the effects of pitch pine seedling roots on extractable N, microbial growth rate, biomass C and N, and nematodes and microarthropods in microcosms with either organic (41% C, 1.14% N) or mineral (0.05% C, 0.01% N) horizon soils of a spondosol. Root quantity was manipulated by varying plant density (0, 1, 2, or 4 seedlings) and rhizosphere soil was separated from non-rhizosphere soil by a 1.2 m mesh fabric. In the rhizosphere of organic soil horizons, moisture, microbial growth rate, biomass C and N, and extractable N declined as root density was increased, but there was little effect on nematodes or microarthropods. High levels of extractable N remained after 5 months, suggesting that N mineralization was stimulated during the incubation. In the rhizosphere of mineral soil horizons, microbial growth rate, and nematode and microarthropod abundances increased at higher root density, and in the absence of roots faunal abundance approached zero. Faunal activity was concentrated in the rhizosphere compared to non-rhizosphere soil. In organic soil horizons, roots may limit microbial activity by reducing soil moisture and/or N availability. However, in mineral soil horizons, where nutrient levels are very low, root inputs can stimulate microbial growth and faunal abundance by providing important substrates for microbial growth. Our results demonstrate a rhizosphere effect for soil fauna in the mineral soil, and thus extends the rhizosphere concept to components of the soil community other than microbes for forest ecosystems. Although our results need to be verified by field manipulations, we suggest that the effects of pine roots on nutrient cycling processes in coniferous forests can vary with soil nutrient content and, therefore, position in the soil profile.     

Carbon turnover in the rhizosphere under continuous plant labeling with <Superscript>13</Superscript>CO<Subscript>2</Subscript>: Partitioning of root,microbial, and rhizomicrobial respiration

The input of labeled C into the pool of soil organic matter, the CO₂ fluxes from the soil, and the contribution of root and microbial respiration to the CO₂ emission were studied in a greenhouse experiment with continuous labeling of oat plants with ¹³CO₂ using the method of the natural ¹³C abundance in the air. The carbon of the microbial biomass composed 56 and 39% of the total amounts of ¹³C photoassimilates in the rhizosphere and in the bulk soil, respectively. The contribution of root respiration to the CO₂ emission from the soil reached 61–92%, including 4–23% of the rhizomicrobial respiration. The contribution of the microbial respiration to the total CO₂ emission from the soil varied from 8 to 39%. The soil organic matter served as the major carbon-containing substrate for microorganisms in the bulk soil and in the rhizosphere: 81–91% of the total amount of carbon involved in the microbial metabolism was derived from the soil organic matter.     

Microbial colonisation of roots as a function of plant species

Fifteen plants species were grown in the greenhouse on the same soil and sampled at flowering to obtain rhizosphere soil and root material. In both fractions, the data on fungal and bacterial tissue obtained by amino sugar analysis were compared with the total microbial biomass based on fumigation-extraction and ergosterol data. The available literature on glucosamine concentrations in fungi and on muramic acid concentrations in bacteria was reviewed to prove the possibility of generating conversion values for general use in root material. All microbial properties analysed revealed strong species-specific differences in microbial colonisation of plant roots. The root material contained considerable amounts of microbial biomass C and biomass N, reaching mean levels of 10.9 and 1.4 mg g⁻¹ dry weight, respectively. However, the majority of CHCl₃ labile C and N, i.e. 89 and 55% was root derived. The average amount of ergosterol was 13 μg g⁻¹ dry weight and varied between 0.0 for Phacelia roots and 45.5 μg g⁻¹ dry weight for Vicia roots. The ergosterol content in root material of mycorrhizal and non-mycorrhizal plant species did not differ significantly. Fungal glucosamine was converted to fungal C by multiplication by 9 giving a range of 7.1-25.9 mg g⁻¹ dry weight in the root material. Fungal C and ergosterol were significantly correlated. Bacterial C was calculated by multiplying muramic acid by 45 giving a range from 1.7 to 21.6 mg g⁻¹ dry weight in the root material. In the root material of the 15 plant species, the ratio of fungal C-to-bacterial C ranged from 1.0 in mycorrhizal Trifolium roots to 9.5 in non-mycorrhizal Lupinus roots and it was on average 3.1. These figures mean that the microbial tissue in the root material consists on average of 76% fungal C and 24% bacterial C. The differences in microbial colonisation of the roots were reflected by differences in microbial indices found in the rhizosphere soil, most strongly for microbial biomass C and ergosterol, but to some extent also for glucosamine and muramic acid.     

Carbon flows in the rhizosphere of ryegrass (Lolium perenne)

This study addresses the issue of carbon (C) fluxes through below ground pools within the rhizosphere of Lolium perenne using the ¹⁴C pulse labeling. Lolium perenne was grown in plexiglas chambers on topsoil of a Haplic Luvisol under controled laboratory conditions. ¹⁴C‐CO₂ efflux from soil, as well as ¹⁴C content in shoots, roots, soil, dissolved organic C (DOC), and microbial biomass were monitored for 11 days after the pulsing. Lolium allocates about 48 % of the total assimilated ¹⁴C below the soil surface, and roots were the primary sink for this C. Maximum ¹⁴C content in the roots was observed 12 hours after the labeling and it amounts to 42 % of the assimilated C. Only half of the ¹⁴C amount was found in the roots at the end of the monitoring period. The remainder was lost through root respiration, root decomposition, and rhizodeposition. Six hours after the ¹⁴C pulse labeling soil accounted for 11 %, DOC for 1.1 %, and microbial biomass for 4.9 % of assimilated C. ¹⁴C in CO₂ efflux from soil was detected as early as 30 minutes after labeling. The maximum ¹⁴C‐CO₂ emission rate (0.34 % of assimilated ¹⁴C h^—1) from the soil occurred between four and twelve hours after labeling. From the 5^th day onwards, only insignificant changes in carbon partitioning occurred. The partitioning of assimilated C was completed after 5 days after assimilation. Based on the ¹⁴C partitioning pattern, we calculated the amount of assimilated C during 47 days of growth at 256 g C m^—2. Of this amount 122 g C m^—2 were allocated to below ground, shoots retained 64 g C m^—2, and 70 g C m^—2 were lost from the shoots due to respiration. Roots were the main sink for below ground C and they accounted for 74 g C m^—2, while 28 g C m^—2 were respired and 19 g C m^—2 were found as residual ¹⁴C in soil and microorganisms.     

Coffee mucilage impact on young coffee seedlings and soil microorganisms

A greenhouse pot experiment was carried out to assess the effects of fermented coffee mucilage applied as mulch together with maize leaves on the growth of young coffee plants of two different varieties and on soil microbial biomass indices. The coffee variety Catuai required 32% more water per g plant biomass than the variety Yellow Caturra, but had a 49% lower leaf area, 34% less shoot and 46% less root biomass. Maize and mucilage amendments did not affect leaf area, shoot and root yield, or the N concentration in shoot and root dry matter. The amendments always reduced the water use efficiency values, but this reduction was only significant in the maize+mucilage‐14 (= 14 g mucilage pot^?1) treatment. Soil pH significantly increased from 4.30 in the control to 4.63 in the maize+mucilage‐14 treatment. Microbial biomass C increased by 18.5 µg g^?1 soil, microbial biomass N by 3.1 µg g^?1 soil, and ergosterol by 0.21 µg g^?1 soil per g mucilage added pot^?1. The presence of mucilage significantly reduced the microbial biomass‐C/N ratio from a mean of 13.4 in the control and maize treatments to 9.3, without addition rate and coffee variety effects. The application of non‐composted mucilage is recommended in areas where drought leads to economic losses and in coffee plantations on low fertility soils like Oxisols, where Al toxicity is a major constraint.     

Three-source partitioning of CO2 efflux from soil planted with maize by C natural abundance fails due to inactive microbial biomass

A theoretical approach to the partitioning of carbon dioxide (CO₂) efflux from soil with a C₃ vegetation history planted with maize (Zea mays), a C₄ plant, into three sources, root respiration (RR), rhizomicrobial respiration (RMR), and microbial soil organic matter (SOM) decomposition (SOMD), was examined. The δ¹³C values of SOM, roots, microbial biomass, and total CO₂ efflux were measured during a 40-day growing period. A three-source isotopic mass balance based on the measured δ¹³C values and on assumptions made in other studies showed that RR, RMR, and SOMD amounted to 91%, 4%, and 5%, respectively. Two assumptions were thoroughly examined in a sensitivity analysis: the absence of ¹³C fractionation and the conformity of δ¹³C of microbial CO₂ and that of microbial biomass. This approach strongly overestimated RR and underestimated RMR and microbial SOMD. CO₂ efflux from unplanted soil was enriched in ¹³C by 2.0‰ compared to microbial biomass. The consideration of this ¹³C fractionation in the mass balance equation changed the proportions of RR and RMR by only 4% and did not affect SOMD. A calculated δ¹³C value of microbial CO₂ by a mass balance equation including active and inactive parts of microbial biomass was used to adjust a hypothetical below-ground CO₂ partitioning to the measured and literature data. The active microbial biomass in the rhizosphere amounted to 37% to achieve an appropriate ratio between RR and RMR compared to measured data. Therefore, the three-source partitioning approach failed due to a low active portion of microbial biomass, which is the main microbial CO₂ source controlling the δ¹³C value of total microbial biomass. Since fumigation-extraction reflects total microbial biomass, its δ¹³C value was unsuitable to predict δ¹³C of released microbial CO₂ after a C₃-C₄ vegetation change. The second adjustment to the CO₂ partitioning results in the literature showed that at least 71% of the active microbial biomass utilizing maize rhizodeposits would be necessary to achieve that proportion between RR and RMR observed by other approaches based on ¹⁴C labelling. The method for partitioning total below-ground CO₂ efflux into three sources using a natural ¹³C labelling technique failed due to the small proportion of active microbial biomass in the rhizosphere. This small active fraction led to a discrepancy between δ¹³C values of microbial biomass and of microbially respired CO₂.