Modelling the effect of active roots on soil organic matter turnover

The aim of this experiment was to study the effect of living roots on soil carbon metabolism at different decomposition stages during a long-term incubation. Plant material labelled with ¹⁴C and ¹⁵N was incubated in two contrasting soils under controlled laboratory conditions, over two years. Half the samples were cropped with wheat (Triticum aestivum) 11 times in succession. At earing time the wheat was harvested, the roots were extracted from the soil and a new crop was started. Thus the soils were continuously occupied by active root systems. The other half of the samples was maintained bare, without plants under the same conditions. Over the 2 years, pairs of cropped and bare soils were analysed at eight sampling occasions (total-, plant debris-, and microbial biomass-C and -¹⁴C). A five compartment (labile and recalcitrant plant residues, labile microbial metabolites, microbial biomass and stabilised humified compounds) decomposition model was fitted to the labelled and soil native organic matter data of the bare and cropped soils. Two different phases in the decomposition processes showed a different plant effect. (1) During the initial fast decomposition stage, labile ¹⁴C-material stimulated microbial activities and N immobilisation, increasing the ¹⁴C-microbial biomass. In the presence of living roots, competition between micro-organisms and plants for inorganic N weakly lowered the measured and predicted total-¹⁴C mineralisation and resulted in a lower plant productivity compared to subsequent growths. (2) In contrast, beyond 3–6 months, when the labile material was exhausted, during the slow decomposition stage, the presence of living roots stimulated the mineralisation of the recalcitrant plant residue-¹⁴C in the sandy soil and of the humified-¹⁴C in the clay soil. In the sandy soil, the presence of roots also substantially stimulated decomposition of old soil native humus compounds. During this slow decomposition stage, the measured and predicted plant induced decrease in total-¹⁴C and -C was essentially explained by the predicted decrease in humus-¹⁴C and -C. The ¹⁴C-microbial biomass (MB) partly decayed or became inactive in the bare soils, whereas in the rooted soils, the labelled MB turnover was accelerated: the MB-¹⁴C was replaced by unlabelled-C from C derived from living roots. At the end of experiment, the MB-C in the cropped soils was 2.5–3 times higher than in the bare soils. To sustain this biomass and activity, the model predicted a daily root derived C input (rhizodeposition), amounting to 5.4 and 3.2% of the plant biomass-C or estimated at 46 and 41% of the daily net assimilated C (shoot + root + rhizodeposition C) in the clay and sandy soil, respectively. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root carbon inputs to the rhizosphere stimulate extracellular enzyme activity and increase nitrogen availability in temperate forest soils

The exudation of carbon (C) by tree roots stimulates microbial activity and the production of extracellular enzymes in the rhizosphere. Here, we investigated whether the strength of rhizosphere processes differed between temperate forest trees that vary in soil organic matter (SOM) chemistry and associate with either ectomycorrhizal (ECM) or arbuscular mycorrhizal (AM) fungi. We measured rates of root exudation, microbial and extracellular enzyme activity, and nitrogen (N) availability in samples of rhizosphere and bulk soil influenced by four temperate forest tree species (i.e., to estimate a rhizosphere effect). Although not significantly different between species, root exudation ranged from 0.36 to 1.10 g C m^?2 day^?1, representing a small but important transfer of C to rhizosphere microbes. The magnitude of the rhizosphere effects could not be easily characterized by mycorrhizal associations or SOM chemistry. Ash had the lowest rhizosphere effects and beech had the highest rhizosphere effects, representing one AM and one ECM species, respectively. Hemlock and sugar maple had equivalent rhizosphere effects on enzyme activity. However, the form of N produced in the rhizosphere varied with mycorrhizal association. Enhanced enzyme activity primarily increased amino acid availability in ECM rhizospheres and increased inorganic N availability in AM rhizospheres. These results show that the exudation of C by roots can enhance extracellular enzyme activity and soil-N cycling. This work suggests that global changes that alter belowground C allocation have the potential to impact the form and amount of N to support primary production in ECM and AM stands.     

Response of long-, medium- and short-term processes of the carbon budget to overgrazing-induced crusts in the Tibetan Plateau

The Kobresia pastures of the Tibetan Plateau represent the world’s largest alpine grassland ecosystem. These pastures remained stable during the last millennia of nomadic animal husbandry. However, strongly increased herds’ density has promoted overgrazing, with unclear consequences for vegetation and soils, particularly for cycles of carbon (C), nutrients and water. Vegetation-free patches of dead root-mat covered by blue-green algae and crustose lichens (crusts) are common in overgrazed Kobresia pastures, but their effect on C turnover processes is completely unknown. We tested the hypothesis that the crusts strongly affect the C cycle by examining: (i) the long-term C stock measured as soil organic matter content; (ii) medium-term C stock as dead roots; (iii) recent C fluxes analyzed as living roots and CO₂ efflux; and (iv) fast decomposition of root exudates. Up to 7.5 times less aboveground and 1.9 times less belowground living biomass were found in crust patches, reflecting a much smaller C input to soil as compared with the non-crust Kobresia patches. A lower C input initially changed the long-term C stock under crusts in the upper root-mat horizon. Linear regression between living roots and CO₂ efflux showed that roots contributed 23% to total CO₂ under non-crust areas (mean July–August 5.4 g C m^?2 day^?1) and 18% under crusts (5.1 g C m^?2 day^?1). To identify differences in the fast turnover processes in soil, we added ¹³C labeled glucose, glycine and acetic acid, representing the three main groups of root exudates. The decomposition rates of glucose (0.7 day^?1), glycine (1.5 day^?1) and acetic acid (1.2 day^?1) did not differ under crusts and non-crusts. More ¹³C, however, remained in soil under crusts, reflecting less complete decomposition of exudates and less root uptake. This shows that the crust patches decrease the rates of medium-term C turnover in response to the much lower C input. Very high ¹³C amounts recovered in plants from non-crust areas as well as the two times lower uptake by roots under crusts indicate that very dense roots are efficient competitors with microorganisms for soluble organics. In conclusion, the altered C cycle in the overgrazing-induced crustose lichens and blue-green algae crusts is connected with strongly decreased C input and reduced medium-term C turnover.     

Spatial distribution and turnover of root-derived carbon in alfalfa rhizosphere depending on top- and subsoil properties and mycorrhization

Aims

This study analyzed the extent to which root exudates diffuse from the root surface towards the soil depending on topsoil and subsoil properties and the effect of arbuscular mycorrhizal fungal hyphae on root-derived C distribution in the rhizosphere.

Methods

Alfalfa was grown in three-compartment pots. Nylon gauze prevented either roots alone or roots and arbuscular mycorrhizal fungal hyphae from penetrating into the rhizosphere compartments. ¹⁴CO₂ pulse labeling enabled the measurement of ¹⁴C-labeled exudates in dissolved (DOC) and total organic carbon (TOC) in the rhizosphere, distributed either by diffusion alone or by diffusion, root hair and hyphal transport.

Results

Root exudation and microbial decomposition of exudates was higher in the rhizosphere with topsoil compared to subsoil properties. Exudates extended over 28 mm (DOC) and 20 mm (TOC). Different soil properties and mycorrhization, likely caused by the low arbuscular mycorrhizal colonization of roots (13?±?4 % (topsoil properties) and 18?±?5 % (subsoil properties)), had no effect.

Conclusions

Higher microbial decomposition compensated for higher root exudation into the rhizosphere with topsoil properties, which resulted in equal exudate extent when compared to the rhizosphere with subsoil properties. Higher ¹⁴C activity used for labeling compared with previous studies enabled the detection of low exudate concentrations at longer distances from the root surface.     

The invasive annual cheatgrass releases more nitrogen than crested wheatgrass through root exudation and senescence

Plant-soil feedbacks are an important aspect of invasive species success. One type of feedback is alteration of soil nutrient cycling. Cheatgrass invasion in the western USA is associated with increases in plant-available nitrogen (N), but the mechanism for this has not been elucidated. We labeled cheatgrass and crested wheatgrass, a common perennial grass in western rangelands, with ¹⁵N-urea to determine if differences in root exudates and turnover could be a mechanism for increases in soil N. Mesocosms containing plants were either kept moist, or dried out during the final 10 days to determine the role of senescence in root N release. Soil N transformation rates were determined using ¹⁵N pool dilution. After 75 days of growth, cheatgrass accumulated 30 % more total soil N and organic carbon than crested wheatgrass. Cheatgrass roots released twice as much N as crested wheatgrass roots (0.11 vs. 0.05 mg N kg^?1 soil day^?1) in both soil moisture treatments. This occurred despite lower root abundance (7.0 vs. 17.3 g dry root kg^?1 soil) and N concentration (6.0 vs. 7.6 g N kg^?1 root) in cheatgrass vs. crested wheatgrass. We propose that increases in soil N pool sizes and transformation rates under cheatgrass are caused by higher rates of root exudation or release of organic matter containing relatively large amounts of labile N. Our results provide the first evidence for the underlying mechanism by which the invasive annual cheatgrass increases N availability and establishes positive plant-soil feedbacks that promote its success in western rangelands.     

Response of root respiration and root exudation to alterations in root C supply and demand in wheat

Rising atmospheric CO₂ concentrations have highlighted the importance of being able to understand and predict C fluxes in plant-soil systems. We investigated the responses of the two fluxes contributing to below-ground efflux of plant root-dependent CO₂, root respiration and rhizomicrobial respiration of root exudates. Wheat (Triticum aestivum L., var. Consort) plants were grown in hydroponics at 20°C, pulse-labelled with ¹⁴CO₂ and subjected to two regimes of temperature and light (12 h photoperiod or darkness at either 15°C or 25°C), to alter plant C supply and demand. Root respiration was increased by temperature with a Q ₁₀ of 1.6. Root exudation was, in itself, unaltered by temperature, however, it was reduced when C supply to the roots was reduced and demand for C for respiration was increased by elevated temperature. The rate of exudation responded much more rapidly to the restriction of C input than did respiration and was approximately four times more sensitive to the decline in C supply than respiration. Although temporal responses of exudation and respiration were treatment dependent, at the end of the experimental period (2 days) the relative proportion of C lost by the two processes was conserved despite differences in the magnitude of total root C loss. Approximately 77% of total C and 67% of ¹⁴C lost from roots was accounted for by root respiration. The ratio of exudate specific activity to CO₂ specific activity converged to a common value for all treatments of 2, suggesting that exudates and respired CO₂were not composed of C of the same age. The results suggest that the contributions of root and rhizomicrobial respiration to root-dependent below-ground respiration are conserved and highlight the dangers in estimating short-term respiration and exudation only from measurements of labelled C. The differences in responses over time and in the age of C lost may ultimately prove useful in improving estimates of root and rhizomicrobial respiration.     

Defoliation alters the relative contributions of recent and non-recent assimilate to root exudation from Festuca rubra

The deposition of organic compounds from plant roots is a key determinant of rhizosphere microbial activity and community structure. Consequently, C-flow from roots to soil is an important process in coupling plant and microbial productivity, via impacts on microbial nutrient cycling in soil. Experimentally, isotopic tracers (¹³C or ¹⁴C) are used to track C inputs to soil and microbial communities. However, in many such studies the relationship between labelled C-flows and total C-flows are not established, limiting the interpretative value of the results. In this study, we applied steady-state near natural abundance ¹³CO₂ labelling to determine the impact of partial defoliation of Festuca rubra on root exudation. This approach in axenic culture facilitated determination of the contribution of pre- and post-defoliation assimilates both to root C-flow and plant tissues. The results demonstrated that total root exudation was increased in the two days following defoliation. This was concurrent with reduced net CO₂ assimilation and reduced allocation of post-defoliation assimilates below-ground and to active root meristems. Through determination of the δ¹³C of root exudates, it was established that the source of the increased root exudation was pre-defoliation assimilate. However, this response was transient, with reduced deposition of pre- and post-defoliation assimilates from roots during the period 2–4 d following defoliation. The results highlight the limitations of pulse-labelling approaches as a means of quantifying impacts of treatments on root exudation, particularly where the treatment is likely to affect plant C-partitioning or the balance between deposition to, and re-mobilization from, C-storage pools.     

A new approach for the quantification of root-cap mucilage exudation in the soil

Direct evidence on the functions of root-cap mucilage during plant root growth in soil is limited mainly due to the lack of a method for in situ measurements. In this paper, we offer a method that facilitates the measurement of mucilage exudation when roots are growing in soil. We observed the mucilage exudation directly through a transparent panel located on the side of a root box in which plant roots were growing. We used a CCD camera attached to a microscope to observe and record mucilage exudation. Using image analysis, the activity of mucilage exudation was evaluated based on the area occupied by the mucilage on the root tip. The area of mucilage observed on the root tips after 1-h growth in soil corresponded with the weight of mucilage that was originally observed on the tips before they were transplanted. This relationship suggests that the observed area on root tip relates to total exudation. The area of mucilage exudation on the root tips was high (0.48 mm²) at night and low (0.35 mm²) at midday, suggesting that the activity of mucilage exudation follows diurnal changes. Furthermore, the mucilage exudation positively correlated with the root elongation rate, implying that fast-growing roots exude more mucilage.     

No Differences in Soil Carbon Stocks Across the Tree Line in the Peruvian Andes

Reliable soil organic carbon (SOC) stock measurements of all major ecosystems are essential for predicting the influence of global warming on global soil carbon pools, but hardly any detailed soil survey data are available for tropical montane cloud forests (TMCF) and adjacent high elevation grasslands above (puna). TMCF are among the most threatened of ecosystems under current predicted global warming scenarios. We conducted an intensive soil sampling campaign extending 40 km along the tree line in the Peruvian Andes between 2994 and 3860 m asl to quantify SOC stocks of TMCF, puna grassland, and shrubland sites in the transition zone between the two habitats. SOC stocks from the soil surface down to the bedrock averaged (±standard error SE) 11.8 (±1.5, N = 24) kg C/m² in TMCF, 14.7 (±1.4, N = 9) kg C/m² in the shrublands and 11.9 (±0.8, N = 35) kg C/m² in the grasslands and were not significantly different (P > 0.05 for all comparisons). However, soil profile analysis revealed distinct differences, with TMCF profiles showing a uniform SOC distribution with depth, shrublands a linear decrease, and puna sites an exponential decrease in SOC densities with soil depth. Organic soil layer thickness reached a maximum (~70 cm) at the upper limit of the TMCF and declined with increasing altitude toward puna sites. Within TMCF, no significant increase in SOC stocks with increasing altitude was observed, probably because of the large variations among SOC stocks at different sites, which in turn were correlated with spatial variation in soil depth.     

<Emphasis Type="Italic">Gluconacetobacter diazotrophicus</Emphasis> exopolysaccharide protects bacterial cells against oxidative stress in vitro and during rice plant colonization

Aims

Increasing the input and turnover of root tissue is considered to be one method that may increase carbon (C) inputs and storage in soil. The use of herbicide during pasture renewal (periodic re-sowing of pasture) is expected to increase root inputs and turnover as plants die. The objective of this study was to quantify the short-term impact of pasture renewal on root turnover and C input to soil of ryegrass-clover pastures.

Methods

Pastures were labelled in the field using a ¹³C isotope pulse labelling method within 1 m² clear chambers. Five daily labelling events were carried out during one week in paired treatment plots within 3 replicate paddocks. One plot per paddock was sprayed with herbicide and then the pasture was renewed by direct drilling of seed. The ¹³C of roots and soil (0–100 mm) was measured at regular intervals over an 89-day period.

Results

Herbicide application caused an initial rapid turnover time of 17 days followed by a slower turnover time of 524 days, compared to unsprayed pasture which had a root turnover of 585 days. Faster root turnover following herbicide application resulted in greater cumulative C input to soil over 89 days with approximately double the C input in the sprayed treatment (3238 ± 378 kg C ha^?1) compared to the unsprayed treatment (1726 ± 540 kg C ha^?1).

Conclusions

The use of glyphosate during pasture renewal increased root turnover and resulted in a greater short term cumulative C input to soil. This study provides the first values of root turnover and C input to soil during a pasture renewal event in New Zealand pasture systems and contributes to the understanding of how pasture roots may influence the soil C input following plant death in grassland systems.

     

Seasonal dynamics of δ13C of C‐rich fractions from Picea abies (Norway spruce) and Fagus sylvatica (European beech) fine roots

The ^13/12C ratio in plant roots is likely dynamic depending on root function (storage versus uptake), but to date, little is known about the effect of season and root order (an indicator of root function) on the isotopic composition of C‐rich fractions in roots. To address this, we monitored the stable isotopic composition of one evergreen (Picea abies) and one deciduous (Fagus sylvatica), tree species' roots by measuring δ¹³C of bulk, respired and labile C, and starch from first/second and third/fourth order roots during spring and fall root production periods. In both species, root order differences in δ¹³C were observed in bulk organic matter, labile, and respired C fractions. Beech exhibited distinct seasonal trends in δ¹³C of respired C, while spruce did not. In fall, first/second order beech roots were significantly depleted in ¹³C, whereas spruce roots were enriched compared to higher order roots. Species variation in δ ¹³C of respired C may be partially explained by seasonal shifts from enriched to depleted C substrates in deciduous beech roots. Regardless of species identity, differences in stable C isotopic composition of at least two root order groupings (first/second, third/fourth) were apparent, and should hereafter be separated in belowground C‐supply‐chain inquiry.     

Exudation rates and δ<Superscript>13</Superscript>C signatures of tree root soluble organic carbon in a riparian forest

Tree root exudation (TRE) of water soluble organic carbon (WSOC) is an important but under-assessed component of net primary production, and is thought to strongly influence rhizosphere biogeochemistry. Riparian systems in particular are often viewed as biogeochemical hot spots fueled partially by root exudate WSOC. However, TRE rates have not been previously reported for these systems. The δ¹³C signatures of exudates may provide important insights into plant physiology and inform isotope-based methods to identify sources of soil CO₂ fluxes, but this information is also generally lacking. In the present study, root exudate WSOC was collected in situ to assess both net exudation rates and exudate δ¹³C values in a temperate riparian forest. Net TRE rates were found to be most strongly related to a combination of tree species, root characteristics and net ecosystem exchange (Adj. R² = 0.73; p < 0.001). In contrast, exudate δ¹³C values were correlated to time-lagged vapor pressure deficit (Adj. R² = 0.21; p < 0.05) and air temperature (Adj. R² = 0.43; p < 0.05), suggesting a rapid transfer of photosynthate from the canopy to the rhizosphere. Extrapolation of mean net TRE rates (13 µmol C g root^?1 day^?1) from a root mass basis to the entire sampling area suggests that TRE may account for as much as 3% of net annual C uptake and represents an important input of organic matter to riparian soils. Our findings of predictable TRE rates and exudate δ¹³C values in the present study suggest that future studies examining δ¹³C values of different plant components, soil organic matter and respired soil CO₂ would benefit by accounting for the impact of root exudates.     

Silicon Alleviates Cadmium Toxicity in Two Cypress Varieties by Strengthening the Exodermis Tissues and Stimulating Phenolic Exudation of Roots

The aim of this work was to investigate the effect of silicon (Si) on phenolic exudation of plant roots and cadmium (Cd) bioavailability in rhizospheres. For this purpose, pot experiments with two cypress varieties, Juniperus chinensis and Platycladus orientalis, each subjected to 100 mg kg^?1 Cd and/or 400 mg kg^?1 Si for 220 days, were conducted using a rhizobag technique. The results showed that P. orientalis accumulated a higher amount of Cd, hence caused higher growth inhibition on the leaves compared with J. chinensis. Si alleviated the growth inhibition induced by Cd toxicity on both varieties, but the mechanisms involved were species specific. For J. chinensis, Si did not affect the root exudation but enhanced the Cd retention of the roots by strengthening the exodermis tissues, restraining Cd translocation from the roots to the shoots. For P. orientalis, Si exposure significantly elevated the phenolic exudation (for example, ferulic acid, catechin, and gallic acid) of the roots, which caused greater Cd mobility in the rhizosphere and enhancement of Cd accumulation in the shoots compared with Cd treatment alone. These results suggest that Cd-chelating with the Si-induced phenolics in the rhizosphere is involved in the Cd detoxification in P. orientalis.     

Shoot P status regulates cluster-root growth and citrate exudation in Lupinus albus grown with a divided root system

The present study was carried out to investigate whether the P concentration in the roots or the shoots controls the growth and citrate exudation of cluster roots in white lupin (Lupinus albus L). Foliar P application indicated that low P concentration in the shoots enhanced cluster‐root growth and citrate‐exudation rate more so than low P concentration in the roots. In the split‐root study, the P concentration in the shoots increased with increased P supply (1, 25 or 75 mmol m^?3 P), to the ‘privileged’ root halves. Roots ‘deprived’ of P invariably had the same low P concentrations, whereas those in the ‘privileged’ roots increased with increasing P supply (1, 25 or 75 mmol m^?3 P). Nevertheless, the proportion of the total root mass allocated to cluster roots, and the citrate‐exudation rates from the root halves were always similar on both root halves, irrespective of P supply, and decreased with increasing shoot P concentrations. Peak citrate exudation rates from developing cluster roots were significantly faster from cluster roots on the ‘deprived’ root halves when the ‘privileged’ half was exposed to 1 mmol m^?3 P as compared with 25 or 75 mmol m^?3 P. The possibility that changes in the concentrations of P fractions in the root halves influenced cluster‐root growth and citrate exudation was discounted, because there were no significant differences in insoluble organic P, ester‐P and inorganic P among all ‘deprived’ root halves. The results indicate that cluster‐root proportions and citrate exudation rates were regulated systemically by the P status of the shoot, and that P concentrations in the roots had little influence on growth and citrate exudation of cluster roots in L. albus.     

Adenosine 5′-monophosphate decreases citrate exudation and aluminium resistance in Tamba black soybean by inhibiting the interaction between 14-3-3 proteins and plasma membrane H<Superscript>+</Superscript>-ATPase

Our previous study suggested that aluminium (Al) stress increased plasma membrane (PM) H⁺-ATPase activity and citrate secretion and simultaneously enhanced the interaction between 14-3-3 proteins and phosphorylated PM H⁺-ATPase in Al-resistant Tamba black soybean (RB). Adenosine 5′-monophosphate (AMP) is known as an inhibitor of the interaction between 14-3-3 proteins and PM H⁺-ATPases. To investigate the effects of AMP on Al resistance, PM H⁺-ATPase activity and citrate exudation, AMP was used to treat Al-stressed RB. The results showed that after treatment with either 100 μM AMP or 50 μM Al for 8 h, RB root growth was inhibited by approximately 50 and 30%, respectively. However, simultaneous treatment with 100 μM AMP and 50 μM Al for 8 h resulted in a 60% inhibition of RB root growth, indicating that the presence of AMP reduced Al tolerance in RB. The interaction of PM H⁺-ATPase and 14-3-3 proteins in the root tips of Al-treated RB was stronger than that in the untreated control. However, the interaction of the two proteins was greatly reduced (lower than that in the control) after co-treatment with Al and AMP, suggesting that the presence of AMP under Al stress reduced the Al-enhanced interaction between PM H⁺-ATPase and 14-3-3 proteins. Consequently, PM H⁺-ATPase activity decreased by approximately 50%, which led to a significant decrease in H⁺ efflux and citrate secretion in RB roots under Al stress. Collectively, these results indicate that AMP reduced citrate exudation and Al resistance in RB by inhibiting the interaction between 14-3-3 proteins and PM H⁺-ATPases under Al stress.     

Fine Root Morphology,Biochemistry and Litter Quality Indices of Fast- and Slow-growing Woody Species in Ethiopian Highland Forest

Fine root turnover of trees is a major C input to soil. However, the quality of litter input is influenced by root morphological traits and tissue chemical composition. In this study, fine roots of ten tropical woody species were collected from an Afromontane forest in the northern highlands of Ethiopia. The fine roots were analysed for root morphological traits and tissue chemistry measured as proxy carbon fractionations. Based on stem increment, the 10 species were divided into faster- and slower-growing species. Faster-growing species exhibited higher specific root length (1362 cm g^?1) than slower-growing species (923 cm g^?1). Similarly specific root area was higher in faster-growing species (223 cm² g^?1) than in slower-growing species (167 cm² g^?1). Among the carbon fractions, the acid-insoluble fraction (AIF) was the highest (44–51%). The carbon content, AIF, and the lignocellulose index were higher for slower-growing species. Root tissue density was lower in faster-growing species (0.33 g cm^?3) than slower-growing species (0.40 g cm^?3) and showed a strong positive correlation with carbon content (r ² = 0.84) and the AIF (r _pearson = 0.93). The morphological traits of fine roots between faster- and slower-growing species reflect the ecological strategy they employ. Slower-growing species have a higher tissue density which may reflect a greater longevity.     

Environmental Control of Root Exudation of Low-Molecular Weight Organic Acids in Tropical Rainforests

The availability of phosphorus (P) can limit net primary production (NPP) in tropical rainforests growing on highly weathered soils. Although it is well known that plant roots release organic acids to acquire P from P-deficient soils, the importance of organic acid exudation in P-limited tropical rainforests has rarely been verified. Study sites were located in two tropical montane rainforests (a P-deficient older soil and a P-rich younger soil) and a tropical lowland rainforest on Mt. Kinabalu, Borneo to analyze environmental control of organic acid exudation with respect to soil P availability, tree genus, and NPP. We quantified root exudation of oxalic, citric, and malic acids using in situ methods in which live fine roots were placed in syringes containing nutrient solution. Exudation rates of organic acids were greatest in the P-deficient soil in the tropical montane rainforest. The carbon (C) fluxes of organic acid exudation in the P-deficient soil (0.7?mol?C?m^?2?month^?1) represented 16.6% of the aboveground NPP, which was greater than those in the P-rich soil (3.1%) and in the lowland rainforest (4.7%), which exhibited higher NPP. The exudation rates of organic acids increased with increasing root surface area and tip number. A shift in vegetation composition toward dominance by tree species exhibiting a larger root surface area might contribute to the higher organic acid exudation observed in P-deficient soil. Our results quantitatively showed that tree roots can release greater quantities of organic acids in response to P deficiency in tropical rainforests.     

Root carboxylate exudation capacity under phosphorus stress does not improve grain yield in green gram

Key message

Genetic variability in carboxylate exudation capacity along with improved root traits was a key mechanism for P-efficient green gram genotype to cope with P-stress but it did not increase grain yield.

Abstract

This study evaluates genotypic variability in green gram for total root carbon exudation under low phosphorus (P) using ¹⁴C and its relationship with root exuded carboxylates, growth and yield potential in contrasting genotypes. Forty-four genotypes grown hydroponically with low (2 μM) and sufficient (100 μM) P concentrations were exposed to ¹⁴CO₂ to screen for total root carbon exudation. Contrasting genotypes were employed to study carboxylate exudation and their performance in soil at two P levels. Based on relative ¹⁴C exudation and biomass, genotypes were categorized. Carboxylic acids were measured in exudates and root apices of contrasting genotypes belonging to efficient and inefficient categories. Oxalic and citric acids were released into the medium under low-P. PDM-139 (efficient) was highly efficient in carboxylate exudation as compared to ML-818 (inefficient). In low soil P, the reduction in biomass was higher in ML-818 as compared to PDM-139. Total leaf area and photosynthetic rate averaged for genotypes increased by 71 and 41 %, respectively, with P fertilization. Significantly, higher root surface area and volume were observed in PDM-139 under low soil P. Though the grain yield was higher in ML-818, the total plant biomass was significantly higher in PDM-139 indicating improved P uptake and its efficient translation into biomass. The higher carboxylate exudation capacity and improved root traits in the later genotype might be the possible adaptive mechanisms to cope with P-stress. However, it is not necessary that higher root exudation would result in higher grain yield.     

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).     

A novel method of quantifying root exudation in the presence of soil microflora

A microcosm is described in which root exudation may be estimated in the presence of microorganisms. Ryegrass seedlings are grown in microcosms in which roots were spatially separated from a microbial inoculant by a Millipore membrane. Seedlings grown in the microcosms were labelled with [¹⁴C]-CO₂, and the fate of the label within the plant and rhizosphere was determined. Inoculation of the microcosms with Cladosporium resinae increased net fixation of the [¹⁴C] label compared to plants grown under sterile conditions. Inoculation also increased root exudation. The use of the microcosm was illustrated and its applications discussed.