128 RBCIAMB | v.56 | n.1 | Mar 2021 | 128-136 - ISSN 2176-9478 A B S T R A C T The objective of this work was to evaluate the distribution of fine roots and its influence on the soil organic carbon stock, at a depth of 20 cm, in a Grevillea robusta and Coffea arabica agroforestry system. The study was conducted in an agroforestry system established 15 years ago in a transition area of Caatinga and Atlantic Forest biomes in Brazil. G. robusta trees representing the most frequent diameter class were selected, and three distances of these trees (0, 0.75 and 1.50 m) and two soil collection depths (0–10 and 10–20 cm) were defined. The root samples were scanned and quantified using a software program. There was a general predominance of roots with a diameter of 0.6 mm at the shortest distance from the surface layer, while there was a predominance of roots with a diameter of 0.4 mm in the 10–20 cm layer. The root carbon stock at a distance of 0.75 m was higher at a depth of 0–10 cm (0.60 Mg ha-1). The soil organic carbon stock also showed higher results in the 0–10 cm layer compared to the 10–20 cm layer, although with significant variation only in the distance of 1.5 m. There was a higher concentration of fine roots in the topsoil, probably influenced by a greater availability of water and nutrients from plant residues. The soil carbon stock is not closely related to root density or root carbon stock. The data presented in this study do not provide a definitive conclusion. Keywords: Grevillea robusta; Coffea arabica; root density; root diameter; organic matter. R E S U M O O objetivo deste trabalho foi avaliar a distribuição de raízes finas e sua influência no estoque de carbono orgânico total do solo, em 20 cm de profundidade, em um sistema agroflorestal de Grevillea robusta e Coffea arabica. O estudo foi realizado em um sistema agroflorestal estabelecido há 15 anos em uma área de transição dos biomas Caatinga e Mata Atlântica no Brasil. Árvores de Grevillea robusta mais representativas da classe de diâmetro de maior frequência foram selecionadas e definidas três distâncias de coleta destas árvores (0, 0,75 e 1,50 m) e duas profundidades do solo (0–10 e 10–20 cm). As raízes presentes nas amostras foram digitalizadas e quantificadas com auxílio de um software. Na menor distância da camada superficial houve predomínio de raízes com diâmetro de 0,6 mm, enquanto, em todas as distâncias da camada 10–20 cm, houve dominância de raízes com diâmetro de 0,4 mm. Na distância de 0,75 m, o estoque de carbono das raízes foi superior na profundidade de 0–10 cm (0,60 Mg ha-1). O estoque de carbono orgânico do solo também apresentou maior resultado na camada 0-10 cm em relação à camada 10–20 cm, embora com variação significativa apenas na distância de 1,5 m. Na camada superficial, ocorreu maior concentração de raízes finas, provavelmente influenciada por uma maior disponibilidade de água e nutrientes provenientes dos resíduos vegetais. O estoque de carbono do solo não está intimamente relacionado com a densidade de raízes e estoque de carbono das raízes. Os dados apresentados neste estudo não fornecem uma conclusão definitiva. Palavras-chave: Grevillea robusta; Coffea arabica; densidade de raízes; diâmetro de raízes; matéria orgânica. Fine root contribution to the soil carbon stock of an agroforestry system in a Caatinga-Atlantic Forest transition zone Contribuição de raízes finas no estoque de carbono do solo de um sistema agroflorestal em zona de transição caatinga-mata atlântica Paulo Henrique Marques Monroe1 , Patrícia Anjos Bittencourt Barreto-Garcia1 , Maida Cynthia Duca Lima1 , Rayka Kristian Alves Santos1 , Elismar Pereira Oliveira1 , Sarah Rabelo Silva , Dráuzio Correa Gama1 1Universidade Estadual do Sudoeste da Bahia – Vitória da Conquista (BA), Brazil. Correspondence address: Paulo Henrique Marques Monroe – Rua Maria Viana Leal, 412, ap. 202 – Alto Maron – CEP: 45005-008 – Vitória da Conquista (BA), Brazil – E-mail: paulomonroes@gmail.com Conflicts of interest: the authors declare that there are no conflicts of interest. Funding: Improvement of Higher Education Personnel (CAPES). Received on: 03/20/2020. Accepted on: 09/10/2020 https://doi.org/10.5327/Z21769478736 Revista Brasileira de Ciências Ambientais Brazilian Journal of Environmental Sciences Revista Brasileira de Ciências Ambientais Brazilian Journal of Environmental Sciences ISSN 2176-9478 Volume 56, Number 1, March 2021 https://orcid.org/0000-0002-0000-3426 https://orcid.org/0000-0002-8559-2927 https://orcid.org/0000-0002-9946-1258 https://orcid.org/0000-0003-2232-8288 https://orcid.org/0000-0002-9299-4633 https://orcid.org/0000-0002-2608-9643 http://orcid.org/0000-0002-6357-0698 mailto:paulomonroes@gmail.com https://doi.org/10.5327/Z21769478736 http://www.rbciamb.com.br http://abes-dn.org.br/ Fine root contribution to the soil carbon stock of an agroforestry system in a Caatinga-Atlantic Forest transition zone 129 RBCIAMB | v.56 | n.1 | Mar 2021 | 128-136 - ISSN 2176-9478 Introduction Agroforestry systems (AFS) are defined as any land use system that implies the deliberate introduction or maintenance of two or more plant species, where at least one of these is arboreal or other perennial species and is associated with agricultural crops, pasture and/or live- stock to exploit the ecological and economic interactions of the differ- ent components (NAIR; NAIR, 2014). AFS are known for contributing to mitigating greenhouse gases (STOUT; LAL; MONGER, 2016; VI- CENTE; GAMA-RODRIGUES; GAMA-RODRIGUES, 2016), as they provide environmentally beneficial services and economic advantages to farmers (RODRIGUES et al., 2007). Interspecific competition for water and nutrients between inter- cropped crops is a variable that interferes in the productivity of AFS. Tree roots can extend into the crop lines in the active crop competition zone, which occurs up to just over 1 m apart, but mainly in the surface layers (THEVATHASAN; GORDON, 2004). Thus, the choice of deeper-rooted tree species in AFS may reduce vertical competition due to their expansion by layers not exploited by crops, promoting better use of the soil profile (PADOVAN et al., 2015; BORDEN; THOMAS; ISAAC, 2017). On the other hand, even for deep-rooted species, it is observed that a large vol- ume of roots is present in the surface layers, absorbing nutrients from the mineralization of plant residues. According to Defrenet et al. (2016), even though species in AFS exploit different soil niches, coffee roots often dom- inate the fine root population of the system in surface layers. It is due to higher root biomass in coffee AFS (MEIRELES et al., 2019). The accumulation of organic carbon in the soil in AFS results from added and decomposing plant residues (from shoots and roots) com- ing from the different species that compose the system (CHATTERJEE et al., 2018). It is estimated that the soil organic carbon (SOC) stock in AFS differs across regions of the world and at different soil depths, ranging from 30-300 Mg C ha-1 (AGEVI et al., 2017). Roots in general, and in particular fine roots, constitute an import- ant carbon input into the soil (LIAO et al., 2014). The degree of fine root carbon (FRC) contribution to SOC depends on land use and manage- ment system, which determines the root architecture, cycling rate, root exudates and colonization by mycorrhizae (HERTEL; HARTEVELD; LEUSCHNER, 2009; POLLIERER et al., 2012). In addition, root distri- bution is influenced by edaphic characteristics. For example, Addo-Dan- so et al. (2020) reported a positive relationship between base saturation and specific root length and specific root area. Fine roots also show greater length in tropical sites where available phosphorus in the soil is low. Le Bissonnais et al. (2018) evaluated the effect of land use gradients (monocultures to AFS and forest) on soil aggregate stability, which was higher in surface layers than deeper layers. Aggregate stability was the main driver of SOC, cation exchange capacity and root traits. The quantification of root biomass and its distribution in the soil may help to understand the relationship between root dynamics and SOC stock. Although many studies on this topic have already been per- formed with tree species, most of them are based on quantifying the total root system biomass, and the number of studies measuring the density of fine roots in different diameter classes is limited. In this work, fine roots were found in the surface layer and directly related to the higher con- centration of organic matter and nutrients (WITSCHORECK; SCHUM- ACHER; CALDEIRA, 2003). Chatterjee et al. (2020) also reported that the SOC stock in coffee AFS was increased only to a depth of 10 cm after 17 years of establishment due to shade-pruned Erytrina spp. Given the above, the present study aimed to evaluate the distribu- tion of fine roots and its influence on the SOC stock, at a depth of 20 cm, in a Grevillea robusta and Coffea arabica agroforestry system. Our study is based on the fact that fine root turnover is the dominant form of below-ground carbon input (UPSON; BURGESS, 2013), with a cycle of less than one year (FREITAS; BARROSO; CARNEIRO, 2008). Materials and methods Study site characterization The study was carried out in an AFS formed by the G. robusta A. Cunn. ex. R. Br and C. arabica plants planted 15 years ago, spaced 3.5 m (between trees) × 1.5 m (between trees and coffee plants) × 2.5 m (between coffee plants). The area is located in Lucaia District, Planalto municipality, Bahia State (coordinates UTM X: 334277 and Y: 8368812). The AFS was located in an area with pasture naturally formed with predominance of genus Brachiaria grass. Other species were not present in the AFS. The region has an average altitude of 943 m and a tropical alti- tude climate (Cwb type according to the Köppen classification), with an average temperature of 19.2ºC and rainfall of 641 mm year-1 (CLI- MATE-DATA, 2012). The study area is located in a transition section between the Caatinga and Atlantic Forest biomes and has soil classified as dystrophic yellow latosol (EMBRAPA, 2013). The chemical charac- teristics are presented in Table 1. Fine root and soil collection We selected six G robusta trees in the most frequent diameter mea- surements at a height of 1.3 m (DBH) (class center = 27.95 cm) to per- form the root and soil collection. The DBH measurement distribution for trees considering an amplitude of 6.14 cm is shown in Figure 1. The selection was carried out in a total area of 1.5 ha with 132 G. robusta tree/ha and 3530 C. arabica plants/ha. Soil sampling was performed on the G. robusta planting line at 0, 0.75 and 1.50 m from the trunk of each selected tree at depths of 0–10 Table 1 – Chemical attributes of soil under a Grevillea robusta and Coffea arabica agroforestry system. pH P K Ca Mg H + Al H2O mg/dm3 --------------cmolc/dm3 ------------- 6.2 41.5 0.5 3.8 2.6 2.7 Monroe, P.H.M. et al. 130 RBCIAMB | v.56 | n.1 | Mar 2021 | 128-136 - ISSN 2176-9478 and 10–20 cm (Figure 2). Two undisturbed soil samples to evaluate fine roots and soil density and one disturbed sample for SOC determination were taken at each distance and depth, making a total of 108 samples. The disturbed samples were taken using a Dutch auger and the undis- turbed samples using a cylindrical ring auger. It was not possible to iden- tify the origin of the roots as to whether they came from trees or coffee. Soil density Soil density was determined by the volumetric ring method (EM- BRAPA, 2017) in samples with preserved structure and known volume (7 cm in height and 7 cm in diameter, totaling 269.3 cm3 volume). Fine root mass and diameter The soil samples were placed in plastic containers and then washed with running water and collected on a 0.25 mm sieve to remove the soil mass to determine the mass and diameter of fine roots (KUMAR; UDAWATTA; ANDERSON, 2010). After washing, all roots were man- ually clamped and arranged on white-bottomed acrylic slides. The roots of each sample were scanned (Figure 3) and were dis- tributed in ten root diameter classes (0.45, 0.63 0.81 1.0 1.18, 1.34, 1.52, 1.70, 1.87, 2.05 mm) with the aid of SAFIRA® software (JORGE; RODRIGUES, 2008) for the different studied distances and depths, ac- cording to the method described by Costa et al. (2014). After scanning, the root samples were placed in aluminum con- tainers, which were then put in a forced-air oven at 65ºC for 72 hours. The samples were subsequently weighed on an analytical balance accu- rate to 0.001g to determine dry mass. Fine root density Root dry mass was used to determine soil root density by means of Equation 1: D = m/v (1) in which: D = density in g cm-3; m = root mass in g; v = ring volume (269.4 cm3); Fine root and soil organic carbon stock Root (after oven drying) and soil (after air drying and 2.0 mm siev- ing) samples were macerated in a mortar. Next, 0.02 g of roots and 0.2 g of soil subsamples were removed and submitted to chemical analy- sis to determine carbon content, using the wet oxidation method with K2Cr2O7 in acid medium and titration with ammonium ferrous sulfate (EMBRAPA, 2017). SOC was calculated on the basis of carbon content and soil density according to Equation 2: SOC = TSOC (g 100 g -1) × Ds × Slt (2) Figure 2 – Representation of soil and fine root sampling points in a Grevillea robusta and Coffea arabica agroforestry system. Figure 1 – Frequency histogram of the diameter classes at a height of 1.3 m from the soil of Grevillea robusta in an agroforestry system with Coffea arabica. Figure 3 – Scanned images of samples utilized to determine root diameter of Grevillea robusta and Coffea arabica. Fine root contribution to the soil carbon stock of an agroforestry system in a Caatinga-Atlantic Forest transition zone 131 RBCIAMB | v.56 | n.1 | Mar 2021 | 128-136 - ISSN 2176-9478 in which: SOC = soil organic carbon stock in Mg ha-1; TSOC = total soil organic carbon content; Ds = soil density (g cm-3); Slt = soil layer thickness (cm). FRC was calculated according to Equation 3: FRC = CFRC (g 100g -1) × Dr × Slt (3) in which: FRC = fine root carbon stock in Mg ha-1; CFRC = fine root organic carbon content; Dr = root density (g cm-3); Slt = soil layer thickness (cm). Statistical analysis The root density, SOC stock and FRC values met the parametric criteria and were then submitted to analysis of variance (ANOVA) according to a 3 × 2 factorial scheme with 6 replications (3 dis- tances and 2 depths). Student’s t-test at 5% significance was adopt- ed to compare means between distances and depths. The analyses were performed using STATISTICA® v.10.0 software (StatSoft Inc., 1984–2011). A descriptive frequency analysis was performed for mean root di- ameter values using the SIGMAPLOT® v.12.0 software program (Systat Software inc.) and the contour maps were produced using the Surfer® v.8.0 program, considering a vertical Cartesian plane formed by the spa- tial distribution of the soil layers and distances of the G. robusta trees. Results Fine root diameter Root diameters ranged from 0.4 to 1.6 mm at the depth of 0–10 cm and from 0.45 to 1.34 mm at the depth of 10–20 cm (Figure 4). Roots in classes of 0.63 mm were more frequent at distances of 0 and 0.75 m and 0.45 mm at 1.5 m. Overall, there was a predominance of roots with a diameter of 0.6 mm in the distances near the trees at the 0–10 cm depth, while there was a predominance of roots with a diameter of 0.4 mm at the distance of 1.5 m at the 0–10 cm depth, and in all distances of the 10–20 cm depth, which represented 60 to 80% of the total roots. Root density and fine root and soil organic carbon stocks The interaction between distance and depth produced a significant effect for the variables of root density and SOC and FRC stocks (Ta- ble  2). Significance was only observed for depth when evaluating the isolated effect of the considered factors. The results of the distance × depth interaction are presented in Table 3 and Figure 5. The FRC and SOC stocks did not vary between the different dis- tances studied. In the case of FRC, differences between depths were only found at a distance of 0.75 m, which showed the highest value in the 0–10 cm layer (Table 3). The SOC stock only showed variation between depths at a distance of 1.5 m, with higher results in the sur- face layer. Higher root density was observed in the first soil layer (0–10 cm). However, there was only variation between distances at a depth of 10–20 cm (Figure 5). Higher values were observed in the distance 0 m, although only with a significant difference at the distance of 0.75 m. Figure 4 – Root distribution frequencies in diameter classes in the (A) 0–10 cm and (B) 10–20 cm soil layer at different distances of Grevillea robusta in an agroforestry system with Coffea arabica. Monroe, P.H.M. et al. 132 RBCIAMB | v.56 | n.1 | Mar 2021 | 128-136 - ISSN 2176-9478 Spatial root distribution influencing soil organic carbon stock The contours formed by the distribution of root density data and FRC and SOC stocks showed a decrease with increasing depth (Figure 6A, 6B and 6C). This means that higher values are present in the surface layer, with a slight displacement to positions closer to the G. robusta tree line. It was noted that the distance influenced the root density contours and consequently the FRC stock. Higher root concentration occurred in the 0-10 cm layer and specifically at a distance of 0 and 0.75 m, decreas- ing vertically and horizontally moving away from this region. SOC distribution ranged from 30 to 20 Mg ha-1 in depth, and showed higher values according to distance (Figure 6C) in the surface layer at a distance of 1.5 m. On the other hand, lower SOC stock values were found at a distance of 0.75 m (25 Mg ha-1), increasing about 2 Mg ha-1 close to the G. robusta trees. It was also possible to notice a similar distribution pattern at greater depth for all evaluated indicators. Discussion Fine root diameter The predominance of roots with larger diameter in the 0–10 cm layer (Figure 4) suggests that roots close to G. robusta trees are associated with plant support, while the roots at increasing distance from the tree are more associated with absorption and therefore have smaller diameters. Greater amounts of subsurface roots are also associated with nutrient exploitation from plant waste mineralization (ISAAC; BORDEN, 2019). According to Mora-Garcés (2018), the distribution of fine roots gener- ally decreases with increasing soil depth. The roots can be found in a non-standardized grouping concentrated in cracks or animal pits, show- ing a large amount of short branches (ZONTA et al., 2006). Fine roots in AFS can also be concentrated in locations with a large agglomeration of plant residues, absorbing nutrients directly from the litter after mineral- ization (THAKUR; KUMAR; KUNHAMU, 2015). Root density and fine root carbon and soil organic carbon stocks The highest FRC values observed in the surface soil layers (Table 3) were expected, as they showed higher root concentration (PADOVAN Table 3 – Fine root carbon (FRC) and soil organic carbon (SOC) stocks as a function of different distances and depths in a Coffea arabica and Grevillea robusta agroforestry system*. Depth (cm) FRC (Mg ha-¹) SOC (Mg ha-1) Distance (m) 0 0.75 1.5 0 0.75 1.5 0–10 0.61 A (0.11) 0.60 A (0.09) 0.47 A (0.08) 26.56 A (3.03) 24.83 A (2.25) 28.36 A (1.99) 10–20 0.35 A (0.08) 0.17 B (0.03) 0.16 A (0.03) 22.36 A (4.22) 20.81 A (2.02) 20.36 B (1.00) *The same letters in the column indicate no significant difference between values by the t-test at 5% probability. Values in parentheses represent the standard error of the mean, n = 6. Table 2 – Summary of variance analysis for fine root carbon (FRC) and soil organic carbon (SOC) stocks and root density in Grevillea robusta with Coffea arabica agroforestry system. SV DF Mean squares FRC SOC Root density Distance 2 0.08NS 10.13NS 1.99E-5NS Depth 1 0.96* 263.16* 8.05 E-7* Dist × Dep 2 0.03* 15.08* 8.58 E-7* Error 30 4.13 41.09 0.85 SV: source of variance; DF: degrees of freedom; Dist: distance; Dep: depth; *significant (p < 0.05) by analysis of variance; NSnot significant. Figure 5 – Root density at three distances of Grevillea robusta trees at two soil depths. The same letters, which compare distances at the same depth, indicate no significant difference between values by the t-test at 5% probability*. ns: not significant; *bars linked to histogram correspond to standard error. Fine root contribution to the soil carbon stock of an agroforestry system in a Caatinga-Atlantic Forest transition zone 133 RBCIAMB | v.56 | n.1 | Mar 2021 | 128-136 - ISSN 2176-9478 et al., 2015; ALBUQUERQUE et al., 2015). This results from the greater contact of the litter (leaves, branches and bark) with the soil, which promotes greater nutrient flow in the surface layer, stimulates the de- velopment of proteoid roots and the accumulation of carbon in the soil after its turnover (PULROLNIK et al., 2009). Morais et al. (2017) found that carbon stored in fine roots is more concentrated in the topsoil (0–10 cm). The authors also observed that fine roots store 40% more carbon than thick and medium roots in this layer. The absence of a difference in FRC stock at distances of 0 and 1.5 m (Table 3) suggested unevenness in G. robusta root development. Thus, it is likely that horizontal variability in fine root distribution is being more influenced by soil resource availability than by root archi- tecture of the species in question. In studying the distribution of G. robusta roots in AFS, Smith et al. (1999) observed great unevenness in the distribution of fine roots and argued that the root distribution complementarity of the different components in the AFS may be com- promised by restrictions in the availability of water and nutrients for the tree component, which results in increased competition (BALJIT; PARAMPARDEEP; GILL, 2016). The reduction in root concentration with increasing depth (Fig- ure 5) can be attributed to the large amount of litter found in AFSs, which contributes to the development of fine roots in the topsoil and in the organic layer itself. Similar results were obtained by Defrenet et al. (2016) in evaluating the biomass and root dynamics of AFSs based on coffee planted in Costa Rica, finding higher amounts of fine roots in surface soil (12% of total roots). Fine root biomass was also twofold higher in the row compared with between rows. The litter acts as a mulch, protecting the surface soil and provid- ing nutrients. Freitas, Barroso and Carneiro (2008) point out that the growth of fine roots (≤ 2 mm) has a strong correlation with the avail- ability of organic matter and soil moisture, being closely associated with litter, since it is a carbon source and favors water retention. Figure 6 – Spatial distribution (A) of root density and (B) fine root carbon (FRC) and (C) soil organic carbon (SOC) stocks at different distances of Grevillea robusta trees associated with Coffea arabica. Monroe, P.H.M. et al. 134 RBCIAMB | v.56 | n.1 | Mar 2021 | 128-136 - ISSN 2176-9478 Spatial root distribution influencing soil organic carbon stock The results obtained did not allow us to determine the origin of the evaluated roots (of C. arabica or G. robusta), which would indicate which species would predominantly be contributing to the carbon ac- cumulation in the soil, since the roots of C. arabica can develop in the middle of the G. robusta lines and vice versa. Some mechanisms (still little known) may alter the root growth of the intercropped crop in an AFS. For example, Livsley, Gregory and Buresh (2000) reported that corn crops showed a greater amount of fine roots and root length in an AFS with Grevillea sp. when compared to a monoculture. A similar pattern was observed by Duan et al. (2019) for oat roots, which were influenced by the presence of walnut, which caused increased root length and decreased root diameter. Regardless of the soil carbon origin, considering the components present in the AFS, it was observed that the FRC stock had little influ- ence on the SOC stock. On the one hand, the SOC stock did not follow a similar distribution as FRC stock in the 0–10 cm layer, on the other hand there was a high correlation between SOC stock, root density and FRC stock in the 10–20 cm layer (Figure 6). This indicated that the carbon accumulation in the 0–10 cm layer depended on more litter contribution than only the carbon originating from the fine root turn- over, as in natural systems of Brazilian biomes (OZÓRIO et al., 2019). The influence of the FRC stock in the 10–20 cm layer was high due to a decreasing carbon incorporation rate from the surface to the deeper layer (CHATTERJEE et al., 2020). AFS are known to have complex re- lations between species which results in a heterogeneous environment, especially in the soil-plant transition. AFSs also help to maintain the natural physical properties of the soil, especially because soil tillage is usually only done in pre-plant- ing (FALCÃO et al., 2020). The conservationist character of AFS assists in natural root turnover, without harming the soil carbon accumulation. Conclusion There is a higher concentration of fine roots in the topsoil which decreases with increasing depth. The root density shows a homoge- neous horizontal distribution from the base of G. robusta, probably being more influenced by litter and edaphic characteristics than by the fine roots. The SOC stock is not closely related to root density or root carbon stock. The data presented in this study do not provide a de- finitive conclusion. Thus, more investigations focusing on identifying the fine root origin of the different species in the AFS are necessary, including deeper layers (up to 100 cm) and evaluating other edaphic characteristics. Acknowledgements We thank CAPES (Improvement of Higher Education Personnel) for funding this project and for granting a postdoctoral scholarship to the first author. We thank the anonymous reviewers for their sugges- tions regarding the manuscript. Contribution of authors: Monroe, P.H.M.: Writing – original draft, Formal analysis, Investigation, Methodology. Barreto-Garcia, P.A.B.: Supervision, Methodology, Writing – review & editing. Lima, M.C.D.: Formal analysis, Writing – review & editing. Santos, R.K.A.: Formal analysis, Writing – review & editing. Oliveira, E.P.: Formal analysis, Writing – review & editing. Silva, S.R.: Formal analysis, Writing – review & editing. Gama, D.C.: Formal analysis, Writing – review. References ADDO-DANSO, S. D.; DEFRENNE, C. E.; MCCORMACK, M. 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