LAND USE AND LOCAL CONDITIONS EFFECTS ON EXTRACELLULAR ENZYME ACTIVITY OF ACTINOBACTERIA STRAINS

Purpose: Evaluate how land use and local conditions affects extracellular enzyme activity of actinobacterial strains isolated from soils in the Brazilian Semi-arid Northeast region. Theoretical framework: The study is framed within soil microbial ecology and environmental microbiology, examining how environmental factors shape the diversity and function of soil bacteria. Method: Soil samples were collected along transects from 3 conservation units into surrounding areas with different land uses. Actinobacteria were isolated and identified morphologically. Extracellular enzyme activities were measured and compared between land uses and locations. Results: The study found considerable actinobacterial diversity, with Streptomyces as the most abundant genus. Amylase, cellulase and xylanase activities varied between locations and land uses. Multivariate analysis revealed heterogeneity in the functional diversity of actinobacterial communities related to land use. Conclusions: Both land use and minor local distinctions can strongly influence the metabolic potential and ecosystem functions of actinobacterial populations in semi-arid environments. Originality/value: Provides new insights into actinobacterial ecology in Brazilian semi-arid soils, showing land use and local conditions strongly influence bacterial community distribution.


INTRODUCTION
Actinobacteria make up a diverse bacterial phylum present in varied terrestrial and aquatic environments.These filamentous Gram-positive bacteria perform essential functions in soils, such as nutrient cycling and regulating ecosystem productivity (Xie;Pathom-Aree, 2021).They do this by secreting a wide range of extracellular enzymes, including proteases, cellulases, lipases and chitinases (Cavalcante et al., 2023).
These enzymes promote the degradation of complex organic polymers like cellulose, chitin and proteins, releasing simpler compounds that other soil organisms can assimilate (Silva et al., 2015).Also, many actinobacterial strains fix atmospheric nitrogen and solubilize unavailable forms of phosphorus, incorporating these essential nutrients into the soil food chain (Gtari et al., 2012).
Land use change involves installing new vegetation and soil management, which frequently removes plant biomass, adds mineral and organic fertilizers, and uses pesticides.These cultivated areas are also a dynamic microbial habitat, due to the periodic destruction of microhabitats by various agricultural practices (Kumar;Karthika, 2020).
In semi-arid climates, like Brazil's Northeast region, characterized by high temperatures, prolonged water deficit and variable annual rainfall, abiotic factors strongly shape actinobacterial community structure (De Barros Corrêa et al., 2019).Furthermore, different land uses and vegetation covers in these semi-arid environments also influence the ecology and functional diversity of these bacterial populations (Bandeira, 2020;Da Cunha et al., 2023).3 Conservation units (CUs) are protected areas mainly used for preserving biodiversity and ecological processes.They also represent privileged locations for ecological and environmental microbiology research (Cavalcante et al., 2023).This is because CUs harbor a variety of ecosystems in a relatively pristine state.Therefore, they are ideal environments to investigate microbial diversity, distribution and function under natural conditions (Lima et al., 2014).
Additionally, establishing CUs helps maintain ecologically distinct habitats and niches, allowing assessment of climate, vegetation and soil management effects on microbiota structure (Cavalcante et al., 2023).
Thus, CUs promote soil microbiology research that provides knowledge about the ecology, diversity and function of these organisms essential for ecosystem functioning.The unique environmental conditions of the semi-arid Northeast shelter yet unexplored actinobacterial diversity, including strains that may produce new bioactive compounds.
In this context, studies investigating the ecology of this microbial group in Northeastern soils contribute to understanding Brazilian diversity and its importance for regional sustainability.Given the above, this study aimed to evaluate how land use affects the extracellular enzyme production activity of actinobacterial strains isolated from soils in the Brazilian Semi-arid Northeast region.

Research Area
The study was developed at the Estação Ecológica de Aiuaba (ESECA), Parque Nacional de Sete Cidades (PNSC), Parque Nacional de Ubajara (PNU) and their respective surrounding áreas, all located in the Northeast region of Brazil.Due to its peculiar characteristics, the Parque Nacional de Ubajara was divided into two areas: Ubajara 1 (dry area) and Ubajara 2 (humid area).The CUs areas are presented in Figure 1.

Sampling
The selection of sampling areas was based on high resolution satellite images provided by the Geo Catalog of the Ministério do Meio Ambiente (MMA), which allowed the delimitation and prior mapping of plots representing different land use classes.These classes were defined in geoprocessing software as protected, conserved, secondary and agricultural, as detailed in Table 1.

Conserved
Collection areas outside the conservation unit, but with the same characteristics as preserved areas.

Preserved
Collection areas located within the conservation unit (PHC).

Intermediate succession
Collection areas that were previously occupied by other ecological communities and are at an intermediate stage of succession.Source: Elaborated by the authors (2023).
The sampling design consisted of four transects starting from each conservation unit studied towards external areas up to 1 km from the edge, with a minimum distance of 3 km between transects.Land use classes were distributed along these gradients, including protected area as control and two external zones labeled as Zones 1 and 2. In each Zone, vegetation fragments and agricultural areas were randomly selected for sampling (Figure 2).Soil collections up to 10 cm deep were carried out between April and July 2018, totaling 98 samples distributed as follows: 28 in Aiuaba, 28 in Sete Cidades and 42 in Ubajara.

Microorganisms
Actinobacteria strains were isolated from soil using the spread plate technique and the casein dextrose agar (CDA) medium (Clark, 1965).The 459 strains of actinobacteria were added to the Culture Collection of the Laboratório de Microbiologia Ambiental (LAMAB) of the Department of Biology of the Universidade Federal do Ceará (UFC).

Soil Chemical Analysis
The collected soil samples were subjected to chemical analyses following methodologies described by Teixeira et al. (2017).The following edaphic parameters were evaluated: pH, electrical conductivity, total organic carbon and total nitrogen contents, available phosphorus, cationic macro and micronutrients and exchangeable sodium.

Micromorphological Characterization (Genus)
The micromorphology of the isolated actinobacteria was analyzed following a methodology adapted from Kern and Blevins (1999) for the inference of the bacterial genus.CDA medium discs (pellets) were obtained by aseptically puncturing the medium in Petri dishes.For each isolate, two pellets were transferred to previously flamed and identified microscopy slides.With the aid of a loop, part of the colony was inoculated around each pellet.A sterile coverslip was added over the pellets and the slides were incubated for 6-7 days at 28 ± 2°C in plates containing moistened cotton to prevent drying.
After incubation, the coverslips were transferred to clean slides containing Amann's Lactophenol cotton blue stain.The preparations were sealed with glycerol and analyzed under an optical microscope (1000x) for photographic documentation.The identification of the bacterial genera was based on the comparison of the observed micromorphological characteristics with reference images contained in specialized atlases and manuals (Miyadoh, 1997;Amaresan et al., 2020).

Enzymatic Activities
The ability of the actinobacteria to secrete extracellular enzymes was evaluated by the spot technique on media containing specific substrates (Gopinath et al., 2017;Balagurunathan et al., 2020).Each strain was inoculated in the form of four spots on plates containing the appropriate enzymatic medium.After the incubation period required for the assay, enzymatic production was evidenced by the formation of a clear halo around the colony, detected directly or after addition of revealing reagents.
The diameter of the degradation halo (Dh) and the colony diameter (Dc) were measured in millimeters using a caliper.The enzymatic index (EI) was calculated according to the formula: EI = Dh/Dc.The assays were performed in quadruplicate and repeated at least twice for each strain, aiming to minimize sampling error.In cases of discrepancy or contamination, a third repetition was required.EI values allowed inference of the enzymatic potential of each studied isolate.

Amylolytic activity
For the amylolytic assay, the strains were inoculated on 1% Starch medium which is composed by peptone 10g, meat extract 3g, NaCl 5g, starch 10g, agar 15g in 1000 mL of distilled water.The strains were incubated for 10 days at 28 ± 2°C (Balagurunathan et al., 2020;Silva et al., 2019).After incubation, starch hydrolysis was revealed by the addition of lugol, which highlights clear halos around colonies producing the enzyme.The degradation halos were measured and related to colony diameters to obtain enzymatic indices, following a standardized methodology.
Similarly, xylanolytic activity was evidenced on medium containing wood xylan, which is composed by xylan 1g, MgSO4.7H2O0.5g, Yeast Extract 0.1g, NaNO3 0.5g, FeSO4.7H2O0.01g.K2HPO4 1g, Agar 15g as substrate, incubating for 7-10 days.Congo red was also used to reveal.In both assays, the halos were measured and related to colony diameters to obtain the enzymatic index.The procedures followed standardized methodology for detecting these activities.

Statistical Analysis
The qualitative data were plotted in pie charts.The quantitative data obtained by the enzymatic indices were subjected to normality and homogeneity of variance tests, followed by ANOVA and Tukey's test (p<0.05),and plotted in Box-Plot graphs comparing land use classes and different locations.
For graphical visualization of the communities, NMDS (non-metric multidimensional scaling) ordination analysis was used.For this analysis, the amylase, cellulase and xylanase activities of the strains and soil chemical analysis data were used.The parameter used to verify the adequacy of the statistical model was stress, and the distance metric used was Euclidean distance (Legendre;Legendre, 2012).The graphs and NMDS analysis were performed using the PAST statistical software version 4.13.

Genus
Micromorphological analysis allowed the identification of 174 bacterial strains, with considerable diversity among the genera (Figure 3).The most abundant genus was Streptomyces, with 103 identified strains, corresponding to 59% of the total.This genus is known for its rich morphological diversity and is one of the main producers of new antibiotics, with a wide range of biological activities (Donald et al., 2022).
The second most abundant genus was Actinomadura, with 25 strains, representing 14% of the total.Actinomadura is an actinobacterial genus that has been associated with the production of bioactive compounds and has shown significant diversity in terms of biological activities (YU; LI; LOU, 2022).The rarest genera identified were Actinoallumurus, Actinobacter, Arthrobacter, Catellatospora, Frankia and Mycobacterium, each with only one strain identified.Arthrobacter is a genus known for its ability to survive in adverse conditions and has been associated with a variety of ecological functions (Krishnan et al., 2016).These results highlight the diversity and biotechnological potential of soil bacteria, especially with regard to the production of bioactive compounds.However, more testing is needed to fully explore the potential of these microorganisms.

Chromogenic Characterization
The results of the chromogenic characterization indicate a diversity in the production of pigments by the strains studied.In the aerial mycelium (Figure 4A), the strains were capable of producing 13 pigments of different colors, with a predominance of white, gray and brown colors, which correspond to 27%, 23% and 19% of the total, respectively.This suggests that these colors may be more common in environmental conditions where aerial mycelium is formed.The pink, blue and orange colors were the rarest, which may indicate a more specific production or particular environmental conditions for their formation (Vandelook et al., 2021).
In relation to the reverse mycelium (Figure 4B), the strains showed 11 distinct colors.The colors white, gray and yellow were predominant, corresponding to 27%, 32% and 12% of the total, respectively2.This may indicate a preference of these strains to produce these pigments under reverse growth conditions.Pink and orange colors were the rarest (Yang et al., 2021).
This diversity of pigments may have important implications for the ecology of these organisms.Pigmentation can play several functional roles in actinobacteria, including protection against UV radiation, interactions with other organisms and adaptation to different environmental conditions (Akanuma et al., 2011;Gmoser et al., 2017).Understanding

Soil Characteristics
Table 2 presents the results of soil analyzes for preserved and conserved land use.In preserved land use, the pH ranges from acidic (4.17 in P-UBJ 1) to slightly acidic (5.67 in P-AIU), which is common in soils from tropical regions due to the leaching of exchangeable bases (Xu et al., 2012).The electrical conductivity is much higher in P-UBJ 1 (2230 μS/cm), indicating a higher concentration of dissolved ions, possibly due to the influence of saline water in this humid location (Othaman et al., 2020).The organic carbon is quite high in P-UBJ 2 (43.33 g/kg), typical of preserved environments with high contribution of organic matter (Gomes et al., 2019).
Nitrogen follows the trend of organic carbon, with higher levels in P-UBJ 2 due to mineralization of O.M. (Marzi, et al., 2020).Potassium ranges from medium (0.54 cmolc/kg in P-AIU) to high (0.63 cmolc/kg in P-UBJ 1), reflecting the availability of this nutrient in the soils of these regions (Setu et al., 2022).Phosphorus is higher in P-UBJ 2 (15.07 g/kg), probably related to nutrient cycling in the preserved environment (Hertzberger, et al., 2020).Calcium and magnesium are higher in P-AIU and P-UBJ 2, expected results in non-leached soils (Sparks et al., 2020).
In conserved land use, pH ranges from acidic (4.72 in C-7Ci) to slightly acidic (5.66 in C-Aiu), typical values in tropical region soils (Xu et al., 2012).The electrical conductivity is higher in C-7Ci and C-UBJ 1, possibly reflecting the presence of dissolved salts in these environments (Othaman et al., 2020).The organic carbon is higher in C-UB 2 (27.46 g/kg), indicating high contribution of organic matter in this area (Gomes et al., 2019).
Nitrogen follows organic carbon levels, especially in C-Aiu (11.49g/kg) due to O.M. mineralization (Marzi, et al., 2020).Potassium ranges from medium (0.3 cmolc/kg in C-UB 2) to high (0.56 cmolc/kg in C-Aiu), important for nutrient cycling (Setu et al., 2022).Phosphorus is higher in C-UB 1 (10.94 g/kg), possibly related to biological activity in the environment.Calcium and magnesium are higher in C-Aiu, typical of well conserved soils (Sparks et al., 2020).Table 3 presents the results of soil analyzes for intermediate succession and agriculture land use.Regarding the intermediate succession land use, pH ranged from acidic (4,76 in I-7Ci) to slightly acidic (5.87 in I-Aiu), which can be explained due to previous land use.Electrical conductivity was higher in I -7ci (818 μS/cm), possibly reflecting the presence of salts in this environment.Organic carbon was high in I-UB 2 (22.94 g/kg) possibly due to O.M. contribution in this site (Gomes et al., 2019).
Regarding the agriculture land use, pH ranged from acidic (5.18 in A-UB 1) to slightly acidic (5.72 in A-Aiu) due to agricultural practices (Park et al., 2010).Electrical conductivity was high in A-UB 1 (1650 μS/cm), possibly due to the cultivation methods used in the area.Organic carbon was higher in A-UB 2 (24.38 g/kg), indicating O.M. contribution (Mcgill;Cole, 1981) Nitrogen followed organic carbon, especially A-UB 2 (10.83 g/kg) by O.M. mineralization.Potassium ranged from medium (0.11 cmolc/kg in A-7Ci) to high (1.03 cmolc/kg in A-UB 1) due to fertilization.Phosphorus was high in all areas, reflecting fertilizer additions.Calcium and magnesium were higher in A-Aiu, resulting from soil correction.10 Comparing all land uses, the preserved areas showed the highest levels of organic carbon, nitrogen and nutrients like calcium, magnesium and potassium, reflecting nutrient cycling in these sites.The conserved areas followed, then the intermediate succession areas and agricultural areas, which showed impacts from previous land use.Fertilizer additions can increase phosphorus in the agricultural areas, which shows in the results we found.

Effect of Agricultural Land Use on the Enzymatic Activity of Soil Actinobacteria
This section compared the amylase, cellulase and xylanase activities of actinobacterial strains isolated from agricultural areas in the three sampled locations (Aiuaba, Sete Cidades and Ubajara).
Amylase activity differed significantly between the agricultural areas of the three localities (ANOVA, p<0.05).Strains isolated from the Sete Cidades agricultural area exhibited the highest amylolytic activity, reaching 17.78.Strains from the Aiuaba and Ubajara agricultural areas showed lower amylolytic activity, with maximum values of 7.82 and 6.9, respectively (Tukey, p<0.05) (Figure 5A).This pattern suggests crops and management practices in the Sete Cidades agricultural area can promote greater starch release to the soil through residue deposition or root exudation (Zakalyukina et al., 2021).Consequently, higher substrate availability appears to have selected for bacterial populations with greater starch degradation and assimilation capacity in this location.Therefore, local variations in agricultural activities seem to shape the functional diversity of soil microbial communities (Helal et al., 2022).
Cellulase activity secreted by cultivable actinobacteria also differed between the sampled agricultural areas (ANOVA, p<0.05).Strains isolated from the Aiuaba and Ubajara agricultural areas exhibited lower cellulolytic activity, with maximum values of 6.71 and 15.97 (Figure 5B), respectively.Strains from the Sete Cidades agricultural area showed greater cellulose degradation capacity, reaching 19.67 (Tukey, p<0.05).This pattern indicates crops and management in Sete Cidades provide higher cellulose inputs to the soil through plant residue deposition (Khosravi et al., 2022).Greater substrate availability appears to have selected for bacterial lineages metabolically more apt at degrading cellulose in this location.Again, local variations in agricultural activities seem to distinctly shape the enzymatic potential of soil microorganisms.
Xylanase activity also differed significantly between the agricultural areas of the three locations (ANOVA, p<0.05).Strains isolated from the Ubajara agricultural area exhibited the greatest xylanolytic capacity, with a maximum value of 11.86 (Figure 5C).Strains from the Aiuaba and Sete Cidades agricultural areas showed lower xylanolytic activity, with maximum values of 5.24 and 6.78, respectively (Tukey, p<0.05).This regional pattern suggests crops and management in Ubajara promote higher xylan inputs to the soil, possibly through deposition of woody crop residues (Dhiman & Mukherjee, 2018).Greater substrate availability appears to have selected for bacterial populations metabolically more capable of degrading xylan in this location.Again, local agricultural differences seem to shape soil microbial functional diversity.

Effect of Conserved Land Use on the Enzymatic Activity of Soil Actinobacteria
This section compared the amylase, cellulase and xylanase enzymatic activities of actinobacterial strains isolated from conserved areas in Aiuaba and Sete Cidades.
Amylolytic activity differed significantly between the conserved areas (ANOVA, p<0.05).Strains from Sete Cidades exhibited higher activity, reaching 5.36.Strains from Aiuaba displayed significantly lower values, from 1.002 to 2.77 (Tukey, p<0.05) (Figure 6A).This regional contrast suggests that edaphoclimatic characteristics and native vegetation conserved in the Sete Cidades area promote greater starch inputs to the soil.Consequently, greater substrate availability appears to have selected for bacterial populations more capable of assimilating and degrading starch in this location (Paul et al., 2021).This result supports the hypothesis that local variations, even among conserved areas, can distinctly shape the enzymatic potential of culturable actinobacteria.
Cellulolytic activity also differed between the conserved areas (ANOVA, p<0.05).Strains from Aiuaba exhibited the lowest indices, ranging from 1.02 to 5.85.Strains from Sete Cidades displayed greater cellulolytic activity, reaching 8.72 (Tukey, p<0.05) (Figure 6B).This spatial pattern suggests that native vegetation characteristics and edaphoclimatic conditions in the Sete Cidades conserved area promote greater cellulose inputs to the soil via litter and root exudation (Zakalyukina et al., 2021).Consequently, greater substrate availability appears to have selected for more efficient cellulose-degrading bacterial lineages.Thus, subtle local variations between these conserved areas were sufficient to distinctly shape the cellulolytic capacity of culturable actinobacterial populations in these locations (Helal et al., 2022).
Xylanolytic activity also differed significantly between Aiuaba and Sete Cidades (ANOVA, p<0.05).Strains from the Aiuaba conserved area displayed the lowest xylanolytic indices, ranging from 1.0 to 10.81.Strains from the Sete Cidades conserved area demonstrated greater xylan degradation capacity, with activity values reaching 11.13 (Tukey, p<0.05) (Figure 6C).Again, this regional contrast suggests local variations in native vegetation and edaphoclimatic conditions between these conserved areas.The plant community and soils in the Sete Cidades conserved area appear to have greater xylan availability (Rachmania et al., 2020).Consequently, these conditions seem to have promoted selection of bacterial lineages with greater capacity to degrade and assimilate this polysaccharide.These results reinforce the hypothesis that subtle local variations, even in conserved areas, can significantly differentiate the enzymatic potential of microbial communities (Goyal et al., 2008).

Effect of Preserved Land Use on the Enzymatic Activity of Soil Actinobacteria
This section compared the amylase, cellulase and xylanase enzymatic activities of culturable actinobacteria isolated from preserved areas in Aiuaba, Sete Cidades and Ubajara.
Amylolytic activity differed significantly between locations (ANOVA, p<0.05).Strains from Sete Cidades displayed the highest activity, differing from Aiuaba and Ubajara (Tukey, p<0.05) (Figure 7A).This result suggests local variations in vegetation composition and edaphoclimatic conditions, generating differences in amylolytic activity.Vegetation formations in Sete Cidades possibly release more starch, supporting bacterial populations more capable of assimilating it (Paul et al., 2021).The variations appear to shape the amylolytic capacity of actinobacteria in each location (Farooq et al., 2021).
Cellulolytic activity also differed between locations (ANOVA, p<0.05).Strains from Aiuaba exhibited the highest indices, differing from Sete Cidades and Ubajara (Tukey, p<0.05) (Figure 7B).One explanation is that the native vegetation in Aiuaba has a composition favorable to cellulose, enriching the soils and promoting more efficient cellulose-degrading bacterial lineages (Domingues et al., 2022).

Effect of Intermediate Succession Land Use on the Enzymatic Activity of Soil Actinobacteria
This section compared the amylase, cellulase and xylanase enzymatic activities of culturable actinobacteria isolated from secondary areas in Aiuaba, Sete Cidades and Ubajara.
These regional contrasts suggest local variations in successional stages, colonizing species, and edaphoclimatic conditions of these areas (Dhiman & Mukherjee, 2018).Consequently, the differential availability of substrates from regenerating vegetation appears to have distinctly shaped the enzymatic capacities of culturable bacterial populations among areas.

NMDS Analysis (Aiuaba)
Nonmetric multidimensional scaling analyses (NMDS) revealed a high heterogeneity among the enzymatic activities of the strains considering the different land uses in Aiuaba (Figure 9).Greater heterogeneity was observed in the intermediate succession area since the points are more widely scattered throughout the graph.Soil calcium content and organic matter are the factors that most influence the distribution of the data along the two-dimensional plane, while sodium content has the least influence.However, it is important to note that NMDS analysis is not a hypothesis test.The model stress remained within an acceptable limit (0.15) for a robust representation of similarity distances between samples (stress < 0.2).

NMDS Analysis (Sete Cidades)
In Sete Cidades, NMDS analysis revealed heterogeneity in the enzymatic activities of the strains, although it is lower than in Aiuaba, as the points are closer together along the twodimensional plane (Figure 10).Unlike Aiuaba, the secondary area in Sete Cidades was the most homogeneous among the land uses, which can be observed by the proximity of the points.Magnesium, sodium, and soil organic matter content are the factors that most influence the distribution of the data along the two-dimensional plane, with nitrogen content being the least influential factor.The model stress remained within an acceptable limit (0.11) for a robust representation of similarity distances between samples (stress < 0.2).

CONCLUSIONS
The study demonstrated that land use significantly affects the extracellular enzyme production activities of cultivable actinobacterial strains isolated from semi-arid soils.Enzyme production capacities varied between agricultural areas, reflecting differences in crop types, residue inputs and management practices.Even conserved and protected areas exhibited spatial heterogeneity in enzyme activities due to subtle local environmental variations.The results show that both land use and minor local distinctions strongly influence the metabolic potential and ecosystem functions of actinobacterial populations in these semi-arid environments.

Figure 1 :
Figure 1: Location Map of Sampled Areas Source: Elaborated by the authors (2023).

Figure 2 :
Figure 2: Soil collection pattern PHCarea within the Conservation Unit; Zone -Areas out of the Conservation Unit; aagriculture bconserved c -intermediate succession Source: Elaborated by the authors (2023).

Figure 5 :
Figure 5: Comparison of the amylolytic (A), cellulolytic (B) and xylanolytic (C) activity of actinobacteria strains in agricultural land use between the sampled areas Source: Elaborated by the authors (2023).

Figure 6 :
Figure 6: Comparison of the amylolytic (A), cellulolytic (B) and xylanolytic (C) activity of actinobacteria strains in conserved land use between the sampled areas Source: Elaborated by the authors (2023).

Figure 7 :
Figure 7: Comparison of the amylolytic (A), cellulolytic (B) and xylanolytic (C) activity of actinobacteria strains in cpreserved land use between the sampled areas Source: Elaborated by the authors (2023).

Figure 8 :
Figure 8: Comparison of the amylolytic (A), cellulolytic (B) and xylanolytic (C) activity of actinobacteria strains in intermediate succession land use between the sampled areas Source: Elaborated by the authors (2023).

Figure 9 :
Figure 9: Multidimensional scaling analysis (NMDS) among the enzymatic activities of the strains considering the different land uses in Aiuaba.Source: Elaborated by the authors (2023).

Figure 10 :
Figure 10: Multidimensional scaling analysis (NMDS) among the enzymatic activities of the strains considering the different land uses in Sete Cidades.Source: Elaborated by the authors (2023).

Table 1 -
Specifications of criteria used for different land uses

Table 3 -
Result of soil chemical analysis in the sampled areas