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Article

Valorization of Amazonian Fruit Biomass for Biosurfactant Production and Nutritional Applications

by
Alan Moura Feio
1,2,
Giulian César da Silva Sá
1,2,
Alexandre Orsato
3,
Karoline Leite
3,
Lucas Mariano Siqueira Pimentel
1,
Joane de Almeida Alves
1,
Glenda Soares Gomes
1,2,
Evelly Oliveira Ramos
1,
Cristina M. Quintella
4,5,
Sinara Pereira Fragoso
6,
José Augusto Pires Bitencourt
7,
Emilly Cruz da Silva
1,8 and
Sidnei Cerqueira dos Santos
1,2,*
1
Bioassays and Bioprocesses Laboratory, Institute of Health and Biological Studies, Federal University of South and Southeast Pará (Unifesspa), Marabá 68500-000, PA, Brazil
2
Graduate Program in Chemistry, Institute of Exact Sciences, Federal University of South and Southeast Pará (Unifesspa), Marabá 68500-000, PA, Brazil
3
Medicinal Molecules Synthesis Laboratory, Department of Chemistry, State University of Londrina (UEL), Londrina 86057-970, PR, Brazil
4
Molecular Kinetics and Dynamics Laboratory, Department of Inorganic and General Chemistry, Federal University of Bahia (UFBA), Salvador 40170-115, BA, Brazil
5
Center for Energy and Environment, Federal University of Bahia (UFBA), Campus Ondina, Salvador 40170-115, BA, Brazil
6
Food Technology Laboratory, Federal University of Paraíba (UFPB), Technology Center, João Pessoa 58051-900, PB, Brazil
7
Vale Technological Institute (ITV), Belém 66055-090, PA, Brazil
8
Materials for Adsorption and Sensing Studies Laboratory, Federal University of Roraima (UFRR), Paricarana Campus, Boa Vista 69310-000, RR, Brazil
*
Author to whom correspondence should be addressed.
Biomass 2025, 5(4), 60; https://doi.org/10.3390/biomass5040060
Submission received: 17 July 2025 / Revised: 29 August 2025 / Accepted: 18 September 2025 / Published: 2 October 2025
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Abstract

Processing economically and socio-culturally significant Amazonian fruits—andiroba (Carapa guianensis Aubl.), açai (Euterpe oleracea Mart.), and babassu (Attalea speciosa Mart. ex Spreng.)—generates substantial biomass waste, posing critical environmental and waste management challenges. This study explored the valorization of these abundant residual biomasses as sustainable feedstocks for biosurfactant production by bacterium Pseudomonas aeruginosa P23G-02, while simultaneously profiling their nutritional value and broader implications for a circular bioeconomy. Through liquid fermentation, biosurfactants were produced at an approximate yield of 6 mg/mL. The isolated biosurfactants exhibited favorable properties, including emulsification indices of around 60% and surface tension reduction to below 30 mN/m, with the andiroba-derived biosurfactant identified as a rhamnolipid type. Nutritional profiling of the residues revealed significant energy values, reaching up to 656 kcal/100 g, with açai and babassu residues being carbohydrate-rich (exceeding 80%), and andiroba residues exhibiting a high lipid profile (up to 57%). These distinct compositions critically influenced biosurfactant yield. These findings underscore the viability of Amazonian fruit biomass as valuable resources for developing eco-friendly bioproducts and innovative waste management solutions. While highlighting a promising pathway for circular bioeconomy development, future research should address biosafety and explore alternative microbial hosts for applications in sensitive sectors such as food and nutrition.

1. Introduction

The Amazon rainforest, spanning approximately 6 million km2, represents a vast reservoir of biodiversity, with over 300 billion trees and more than one million documented species [1]. Among the prominent Amazonian palms (Arecaceae), Euterpe oleracea Mart. (açai) and Attalea speciosa Mart. ex Spreng. (babassu) stand out, yielding more than 90 tons annually. These species play essential roles in sustaining local economies and providing nutritional resources to surrounding communities [2,3]. In addition, Carapa guianensis Aubl. (andiroba)—from the Meliaceae family—is widely exploited for its oil, which has multiple applications in the pharmaceutical, cosmetic, and food industries [4]. Recent market projections indicate that the Amazonian fruit sector will expand from USD 227 million in 2022 to USD 347 million by 2032, emphasizing the global significance of these resources [5].
Despite its extraordinary biodiversity, the Amazon is increasingly threatened by environmental degradation, jeopardizing both its biological diversity and the livelihoods of local communities [6]. The processing of Amazonian fruits generates substantial amounts of waste, often exceeding current utilization capacities, thereby causing economic losses and posing waste management challenges [7]. Converting these residues into diverse applications—including dietary supplements for humans [8] and animals [9], as well as biochar [10,11] and biofertilizers [12]—represents a promising strategy to enhance sustainability and economic resilience. It is hypothesized that the varying nutritional compositions of these fruit residues influence biosurfactant yield and structure, in alignment with the United Nations Sustainable Development Goals on responsible consumption and production, and climate action.
Current trends indicate a growing interest in utilizing residual biomass from fruit processing as sustainable substrates for fermentative processes [13]. Biosurfactant production has attracted considerable attention due to its broad applications and environmental benefits [14,15]. These amphiphilic molecules, primarily synthesized by microorganisms such as Pseudomonas aeruginosa, are characterized by low toxicity and high biodegradability [16]. The use of Amazonian bioresources for biosurfactant production is particularly relevant for biodiversity conservation and local economic development [17]. Valorization of by-products through their conversion into biosurfactants can create new employment opportunities and income streams, thereby fostering a circular bioeconomy and promoting sustainable practices within traditional communities [18].
The economic potential of Amazonian natural resources extends beyond Brazil, offering opportunities to strengthen local economies and support the establishment of sustainable business models. The responsible utilization of Amazonian by-products for biosurfactant production underscores their relevance across socioeconomic and ecological dimensions of global development. In addition, characterizing the nutritional profiles of these substrates may reveal innovative applications in food science while advancing knowledge of their biotechnological potential.
This study assessed the potential of residual biomass obtained from the processing of açai, babassu, and andiroba fruits for biosurfactant production, while also examining the nutritional benefits and environmental implications of these by-products. By integrating these findings into broader sustainability frameworks, the research aims to inform policies and practices that reconcile development needs with the conservation of Amazonian biodiversity.

2. Material and Methods

2.1. Amazon Fruit Residues

Residual biomass from açai (Euterpe oleracea Mart.) was sourced from the Guajanaíra Indigenous Village within the Jurunas Settlement Project in Itupiranga, Pará, Brazil. Babassu (Attalea speciosa Mart. ex Spreng.) biomass was obtained from the Associação do Movimento Interestadual das Quebradeiras de Coco Babaçu (Association of the Interstate Movement of Babassu Coconut Breakers), located in São Domingos do Araguaia, Pará, Brazil. The biomasses of both açai and babassu consisted mainly of fruit peels and residual pulp remaining after oil extraction. The collected biomasses were dried in a forced-air oven (SP-102/64, SP Labor, Presidente Prudente-SP, Brazil) at 60 ± 1 °C until constant weight and subsequently ground to a particle size of 0.07 mm using a knife mill (SL-31, Solab, São Paulo-SP, Brazil). Unrefined andiroba (Carapa guianensis Aubl.) biomass was obtained from the Agroextractivist Settlement Project of Praialta and Piranheira in Nova Ipixuna, Pará, Brazil, consisting primarily of oil extraction bagasse (defatted fraction). The use of açai and babassu is registered with the Brazilian System for Management of Genetic Heritage and Associated Traditional Knowledge (SisGen) under registration number AB70C5B, while andiroba is registered under SisGen number A27D6F4.

2.2. Nutritional Composition of Fruit Residues

Proximate analysis of non-delipidated fruit biomass was conducted in triplicate, following the methods established by the Association of Official Analytical Chemists (AOAC). Moisture content (%) was determined (AOAC method 950.46) by heating empty capsules to 105 °C, followed by desiccation until constant weight was achieved, with moisture calculated as the weight difference before and after heating [19]. Ash content (%) (AOAC method 923.03) was determined from the weight loss after incineration in a muffle furnace at 550 °C, followed by desiccation to constant weight [20]. Crude protein (%) was quantified using the Kjeldahl method (AOAC method 981.10), encompassing digestion, distillation, and titration to assess total nitrogen, with a conversion factor of 5.75 applied to estimate vegetable protein content [21]. Total lipid content (%) was determined by chloroform–methanol extraction according to the method of Bligh and Dyer [22]. Carbohydrate content (%) was calculated by difference [23] using Equation (1).
Carbohydrates = 100 − (Moisture + Ash + Lipids + Crude Protein)
The pH was measured using a digital pH meter (Mapa 200, Marconi, Piracicaba-SP, Brazil). The total energy value (kcal per 100 g of biomass) was calculated as the sum of the products of carbohydrate, lipid, and protein contents and their respective caloric coefficients [23], using values of 4 kcal/g for carbohydrates, 9 kcal/g for lipids, and 4 kcal/g for proteins.

2.3. Isolation of Biosurfactant-Producing Bacteria

Gasoline samples were collected using a drainage pump from a fuel storage tank at a depth of four meters in Marabá, Pará, Brazil. Gasoline was selected as an enrichment medium to isolate robust hydrocarbon-degrading biosurfactant-producing bacteria. Samples were stored in 250 mL Erlenmeyer flasks containing mineral salt medium 1 (MSM-1) with the following composition (g/L): K2HPO4 (4.0), Na2HPO4 (1.5), NaNO3 (1.0), MgSO47H2O (0.2), CaCl22H2O (0.02), and FeCl36H2O (0.02) [24], supplemented with 1% (v/v) gasoline. Incubation was performed in an orbital shaker (SL-22, Solab, Piracicaba-SP, Brazil) at 150 rpm and 30 ± 1 °C for 7 days, repeated three times for a total incubation period of 21 days. Subsequent serial dilutions of the enriched culture were prepared, and 100 μL aliquots were plated onto tryptic soy agar (TSA; Kasvi, Pinhais-PR, Brazil) using the Drigalski loop streaking technique, followed by incubation at 30 ± 1 °C for 48 h [17]. Morphologically distinct colonies were isolated and characterized by morphotintorial analysis [16].
Biosurfactant production was assessed using an emulsification assay with cell-free broth, as detailed in Section 2.7. The strain identified as P23G-02 is preserved in a glycerol solution at −20 °C in the Bioassays and Bioprocessing Laboratory (L@βio), Unifesspa, Marabá, Pará, Brazil. Although −80 °C is standard for long-term storage of Pseudomonas strains to minimize genetic alterations, storage at −20 °C in glycerol is adequate for the experimental procedures in this study. It is important to note that prolonged storage at −20 °C may induce genetic drift, potentially causing phenotypic variations that could affect reproducibility in future applications. This strain has been registered with SisGen under registration number A7DF022.

2.4. Molecular Identification of Biosurfactant-Producing Bacteria

This study focused exclusively on the molecular identification of a single isolated strain, P23G-02. Total genomic DNA from strain P23G-02 was extracted using the ZR Bacterial DNA MiniPrep™ Kit according to the manufacturer’s instructions. The extracted DNA was stored at −20 °C until further use. Amplification of the 16S rRNA gene, specifically targeting the V1–V2 regions, was performed using universal primers 8F (5′–AGA GTT TGA TCC TGG CTC AG–3′) and 1492R (5′–GGT TAC CTT GTT ACG ACT T–3′). These primers were selected based on GC content and minimal potential for primer-dimer or secondary structure formation, ensuring efficient PCR amplification.
PCR reactions were carried out in a total volume of 20 μL, containing 10 μL of GoTaq® Colorless Master Mix 2× (Promega, Madison, WI, USA), 0.6 μM of each primer, 40 ng of genomic DNA, and sterile ultrapure water to reach the desired volume. The amplification protocol consisted of an initial denaturation at 95 °C for 5 min, followed by 30 cycles of denaturation at 95 °C for 30 s, annealing at 56 °C for 40 s, and extension at 72 °C for 1 min, with a final extension at 72 °C for 5 min. Reactions were performed using a Veriti™ Thermal Cycler (Applied Biosystems, Waltham, MA, USA).
PCR products were resolved on a 2% agarose gel and visualized under UV light using UniSafe Dye (Qiagen®, Venlo, The Netherlands). Verified products were purified with Agencourt AMPure XP beads (Beckman Coulter, Brea, CA, USA) and quantified with a Qubit fluorometer (Applied Biosystems, Waltham, MA, USA). Sequencing was performed via capillary electrophoresis on an ABI3730xl DNA Analyzer (Applied Biosystems, Waltham, MA, USA) using the BigDye® Terminator v3.1 Cycle Sequencing Kit (Beckman Coulter, Inc., Brea, USA). PCR products were purified through EDTA/ethanol precipitation and resuspended in 10 μL of Hi-Di™ Formamide (Applied Biosystems, Waltham, MA, USA). Electropherograms were quality-checked with Chromas Lite 2.01, and consensus sequences were assembled in Geneious 4.8.3. Primer regions were trimmed, and the resulting sequence was queried against the GenBank database using BLAST 2.16.0.
For phylogenetic analysis, the eight closest NCBI matches were retrieved and aligned using Clustal Omega v2.1 [25]. Maximum likelihood inference was conducted with RAxML-HPC2 on XSEDE via the CIPRES Science Gateway, employing the GTRGAMMA model with 1000 bootstrap replicates. Alignments were refined using MAFFT v7 with the G-INS-i algorithm and subsequently trimmed with TrimAl v1.3 (Gappyout option) [26,27]. The phylogenetic tree was visualized using FigTree v1.4.4 and rooted with Pseudomonas aeruginosa DSM 50071 (NR 117678.1), identified as the closest reference strain to P23G-02. The 16S rRNA sequence of P23G-02 has been deposited in NCBI (GenBank: PV656461).

2.5. Biosynthesis and Extraction of Biosurfactants

Biosurfactant production was carried out in Erlenmeyer flasks containing 50 mL of sterilized MSM-2. Each flask was supplemented with 3% (w/v) biomass residues from açai, babassu, or andiroba, serving as the sole nutrient source. The pH was adjusted to 7.0. A 5% bacterial inoculum, standardized to an optical density of 0.6–0.8 at 600 nm using a Bel V-M5 Visible Spectrophotometer (Biovera, Wildberg, BW, Germany), was transferred into each culture medium. Colony-forming unit (CFU) counts were performed on inoculum batches to validate correlations with optical density measurements. Flasks were incubated on an orbital shaker at 180 rpm and 30 ± 1 °C for eight days. Biomass concentration and incubation duration was determined by preliminary screening experiments and optimized for biosurfactant production by Pseudomonas sp. [28].
After incubation, cultures were centrifuged at 4500 rpm for 15 min (SL-700, Solab, Aberdeen, UK) to obtain cell-free broth. The supernatant was acidified to pH 2.0 using hydrochloric acid and stored at 4 °C overnight. The precipitate was recovered by centrifugation (4500 rpm, 4 °C, 15 min), resuspended in a chloroform–methanol (3:1, v/v) solvent system in a separatory funnel, manually agitated at 25 ± 1 °C, and allowed to settle for 24 h for phase separation. The hydroalcoholic phase was discarded, and the organic phase was collected; the extraction was repeated twice to maximize biosurfactant recovery. The organic solvent was removed via rotary evaporation (LGI-52CS-1, Scientific, São Paulo, SP, Brazil) and the residue dried in a circulating air oven until a constant weight was achieved [29]. The isolated biosurfactants were subsequently stored for further analysis.

2.6. Characterization of Biosurfactants

Characterization of the biosurfactants included multiple assessments to determine their functional properties. Emulsifying activity was quantified using the emulsification index, following Santos et al. [17]. An emulsification index ≥ 50% is commonly considered indicative of effective emulsifying capability, which is essential for evaluating industrial applicability and correlates strongly with biosurfactant concentration. Non-inoculated MSM and 1% sodium dodecyl sulfate (SDS; Dinâmica Química Contemporânea LTDA, Indaiatuba, SP, Brazil) served as negative and positive controls, respectively. Surface tension was measured using a tensiometer (OCA15 Plus, Data Physics, Filderstadt, Germany) via the hanging drop method at 25 °C, supported by an integrated video imaging system and OCA 10/OCA 20 software [30].
Structural characterization of the biosurfactants was performed using Fourier-Transform Infrared Spectroscopy (FT-IR) with a Vertex 70 model (Bruker, Billerica, MA, USA) equipped with a Platinum ATR reflectance accessory. Spectra were recorded over 4000–400 cm−1 and analyzed using OriginPro 8.0 software. Additionally, Electrospray Ionization Mass Spectrometry (ESI-MS) was employed for detailed structural analysis using an ultra-high-resolution QTOF mass spectrometer (COMPACT model, Bruker Daltonics, Billerica, MA, USA) via direct injection. Mass spectra were acquired in negative ion mode over a mass range of 0–1000 m/z, with a capillary voltage of 3.8 kV, nitrogen nebulizing gas at 4.0 L min−1, gas temperature maintained at 200 °C, and ion energy set at 5.0 eV. Internal calibration was performed using sodium format (m/z 90–1500). A mass tolerance of ±5 ppm was applied for annotation and database searches, with isotopic fit (mSigma) acceptance set at ≤50. Rhamnolipid identification was based on the detection of [M-H]-adducts and confirmed by the error computed between the calculated exact mass and the experimental high-resolution m/z values.

2.7. Statistical Analysis

Statistical analysis was conducted using Analysis of Variance (ANOVA) in GraphPad Prism® version 6.01 (GraphPad Software, Boston, MA, USA) to compare means between biosurfactant and control groups, as well as among the different fruit biomasses. When significant differences were detected (p < 0.05), the Tukey HSD post hoc test was applied to identify specific pairwise differences. All analyses were performed in triplicate (n = 3) per group, with a significance level of 95% (α = 0.05).

3. Results

3.1. Characterization of the Biosurfactant-Producing Bacterium

Alignment analyses performed using Clustal Omega demonstrated 100% identity of the P23G-02 sequence with 16S rRNA sequences of the genus Pseudomonas, particularly P. aeruginosa (Table 1), indicating that the isolated strain P23G-02 belongs to this species. Additionally, alignment results revealed that sequences from the NCBI database shared over 96% similarity with other Pseudomonas species, supporting the taxonomic classification and suggesting a close evolutionary relationship, particularly with P. aeruginosa DSM 50071 (Figure 1). This similarity implies a shared ecological and functional niche among these strains.
Morphologically, strain P23G-02 formed colonies with white pigmentation and greenish hues, exhibiting wavy and convex surfaces. Gram staining confirmed the strain as Gram-negative bacilli, consistent with the morphological characteristics of this species. Initial biosurfactant activity assays demonstrated that the cell-free supernatant from strain P23G-02 achieved an emulsification index of 80% after 24 h at 25 ± 1 °C, indicating substantial biosurfactant production potential.

3.2. Production and Physicochemical Characterization of Biosurfactants

As detailed in Table 2, strain P23G-02 demonstrated effective biosurfactant production using three distinct substrates derived from the residual biomass of Amazonian fruits, achieving emulsification indices of approximately 60%, comparable to the positive control. Importantly, the biosurfactants maintained stable foam formation for over 24 h, indicating potential for industrial applications. Extracted biosurfactant yields varied across the substrates, with peak concentrations reaching approximately 6 mg/mL. The biosurfactants exhibited favorable surface-active properties, reducing surface tension to values as low as 30 mN/m.
In the evaluation of biosurfactant properties (Table 2), ANOVA revealed statistically significant differences among groups for emulsification index (EI), surface tension (ST), and yield (YD). Subsequent Tukey’s post hoc tests indicated that all pairwise comparisons for EI, ST, and YD were statistically significant (p < 0.05).

3.3. Nutritional Profiles of Fruit Residues

Table 3 summarizes the nutritional profiles of açai, babassu, and andiroba biomass, revealing distinct compositional traits, with energy values ranging from 359 to 656 kcal/100 g. Açai and babassu residues were carbohydrate-rich, exceeding 80% of their composition, whereas andiroba residue contained over 55% lipids. The pH of the residues ranged from acidic to nearly neutral.
Regarding the nutritional composition analysis of biomass residues (Table 3), ANOVA indicated significant differences across all evaluated parameters (p < 0.001). Tukey’s post hoc tests confirmed significant pairwise differences among the three biomass types for moisture, lipids, proteins, carbohydrates, pH, and energy value (p < 0.05). Importantly, andiroba biomass differed significantly from both açai and babassu in ash content (p < 0.05), whereas no significant difference was observed between açai and babassu (p > 0.05).

3.4. Biosurfactant Structural Characterization

The chemical characterization of the biosurfactants was performed using FT-IR spectroscopy (Figure 2). The FT-IR spectrum of the andiroba-derived biosurfactant (black spectrum) displayed a prominent band at 3380 cm−1, corresponding to O–H stretching vibrations, commonly associated with hydroxyl groups in fatty acids and alcohols. A peak at 2917 cm−1 indicated C–H stretching vibrations characteristic of methylene (–CH2–) and methyl (–CH3) groups, representing the aliphatic chain structures of the biosurfactants. A distinct peak at 1714 cm−1 was attributed to C=O stretching, associated with ester functionalities and potentially carboxylic acids. The most intense peak at 1031 cm−1 suggested C–O vibrations, typical of cyclic carbohydrate moieties.
The babassu biosurfactant (blue spectrum) displayed a broad band at 3232 cm−1 corresponding to O–H stretching, indicative of hydroxyl functional groups, similar to those observed in andiroba and açai (red spectrum) biosurfactants. C–H stretching vibrations appeared at 2923 cm−1, and C–O stretching was evident at 1075 cm−1.
Complementary ESI-MS analysis further revealed the structural complexity of the biosurfactants (Figure 3). The andiroba biosurfactant exhibited ions with higher m/z ratios, consistent with rhamnolipid-like structures. In contrast, the açai and babassu biosurfactants showed ions with lower m/z ratios, suggesting structures less characteristic of rhamnolipids.
The peak at 503 m/z in the ESI-MS spectrum of the andiroba biosurfactant likely corresponds to the principal compound identified as the mono-rhamnolipid Rha-C10-C10 (one rhamnose unit). The high-resolution m/z value for this peak (m/z 503.3151) results in a 14.9 ppm error when compared to the calculated exact mass of this congener (m/z 503.3226), which reinforces the characterization reported in this work. Additionally, the peak at 333 m/z may represent a possible fragmentation product of Rha-C10-C10, while the peak at 649 m/z corresponds to the molecular ion of the di-rhamnolipid Rha-C10-C10 (two rhamnose units). These results indicate that the rhamnolipid variants in the andiroba biosurfactant likely comprise both mono- and di-rhamnolipids containing two saturated C10 fatty acids. In contrast, ESI-MS analysis of the açai and babassu biosurfactants revealed peaks corresponding to the C10-C10 fragment, lacking rhamnose.

4. Discussion

Natural resources, including bacterial species, serve as crucial reservoirs of bioactive compounds with broad relevance in biological and biotechnological applications [31]. Pseudomonas aeruginosa is widely distributed in soil, water, air, and in both animals and humans [32]. Molecular characterization of the bacterial isolate P23G-02 confirmed its classification as P. aeruginosa, with its 16S rRNA sequence showing complete alignment with reference sequences from this species (Figure 1 and Table 1). Morphological observations further corroborated phenotypic traits typical of Pseudomonas. This phylogenetic and phenotypic congruence supports previous reports [33] on the prevalence of P. aeruginosa in hydrocarbon-contaminated environments influenced by anthropogenic activities.
The strain P. aeruginosa P23G-02 exemplifies the intersection of ecological dynamics and biotechnological innovation. Its ability to metabolize complex organic compounds derived from petroleum underlines its potential in microbial hydrocarbon degradation, consistent with observations by Soberón-Chávez et al. [34]. This metabolic capability positions P23G-02 as a promising biological model for synthesizing industrially relevant biomolecules, particularly biosurfactants. The strain demonstrated biosurfactant production using residual biomass from Amazonian fruits, achieving yields of approximately 6 mg/mL and an emulsification index of 80% (Table 2), surpassing results reported by Santos et al. [17] for P. aeruginosa grown on glycerol.
The selection of residual biomasses as substrates significantly influences biosurfactant yield, as noted by George and Jayachandran [35], who reported variability in production and emulsifying properties depending on the plant residue. For example, orange peel-derived biosurfactants reached 9 mg/mL with an emulsification index of 73%, whereas residues from carrots, lemons, and bananas yielded comparatively lower outputs. These findings highlight the competitive advantage of P23G-02 in fostering a sustainable bioeconomy through valorization of biomass residues, emphasizing the strategic importance of local substrate selection to optimize biosurfactant yield and functional performance.
Furthermore, the produced biosurfactants exhibited remarkable surface-active properties, maintaining stable foam formation for over 24 h and reducing the surface tension of non-inoculated MSM from 69 mN/m to below 30 mN/m (Table 2). Similar surfactant performance has been reported for biosurfactants produced by Candida bombicola [36] and P. aeruginosa [16]. The ability to form stable emulsions is a critical parameter for evaluating industrial surfactant efficacy and correlates strongly with biosurfactant concentration [37]. Achieving surface tension values below 30 mN/m indicates strong potential for the development of commercially viable surfactants [38].
Market analyses by Zhu et al. [39] highlight shifting trends favoring natural over synthetic surfactants, emphasizing the relevance of these findings. Projections from Global Market Insights [40] indicate growing consumer demand for eco-friendly products in sectors such as personal care, pharmaceuticals, therapeutics, and food. This trend has contributed to the expansion of the global biosurfactant market, valued at over $8 billion in 2022 and projected to reach $14.3 billion by 2032. Integrating locally sourced biomass can enhance biosurfactant yield and functionality, strategically positioning these products within a sustainability-driven market landscape while aligning with consumer preferences and regulatory requirements.
The residual biomass from Amazonian fruits constitutes complex nutritional matrices that are critical for fermentative processes. Compositional analysis revealed distinct nutritional profiles (Table 3), with açai and babassu biomasses predominantly carbohydrate-rich (exceeding 80% of their composition), whereas andiroba biomass displayed a high lipid content (exceeding 55%). These nutritional differences, together with the variable biosurfactant yields observed (Table 2), suggest that carbohydrate-rich residues may primarily support biomass growth of strain P23G-02, with carbon flux directed toward cellular maintenance and proliferation, potentially limiting product formation.
The diversity of nutrients in the substrates significantly influenced the metabolic activity of strain P23G-02, thereby affecting biosurfactant production. These findings align with previous studies emphasizing the importance of organic matter availability in biosurfactant biosynthesis [41,42]. Al-Marri et al. [43] further demonstrated the potential of biodegradation processes utilizing complex substrates, such as plant residues, highlighting the link between substrate composition and microbial metabolism. Accordingly, it can be inferred that P. aeruginosa P23G-02 metabolizes residual biomass from Amazonian fruits differently, resulting in variable yields and distinct physicochemical properties of the produced biosurfactants. This observation is consistent with Raaijmakers et al. [44], who showed that a single Pseudomonas strain can produce multiple variants of the same biosurfactant. For a more comprehensive understanding of these mechanisms and to optimize biosurfactant production, future studies should focus on comparative fermentation kinetics, microbial growth curves, and detailed substrate consumption analyses in response to different Amazonian fruit biomasses, thereby refining fermentation parameters and exploring underlying metabolic pathways.
In the analyzed substrates, moisture content remained below the Brazilian regulatory maximum of 15% for plant products [45], a parameter critical for microbial activity and nutrient bioavailability [46]. In naturally low-moisture environments, bacterial metabolic functions may be constrained [47]; however, under the liquid fermentation conditions employed in this study, the low moisture content of the biomasses did not limit the growth or activity of P23G-02. Moreover, the relatively low moisture content is advantageous for biotechnological processing and storage, enhancing biomass stability and reducing the risk of microbial contamination prior to fermentation [48].
The ash content of açai and babassu biomass residues complied with the maximum limit of 1.35% established by Brazilian regulations [45]. While ash content serves as an indicator of mineral composition, many micronutrients within plant matrices display low bioavailability due to complexation with indigestible components [49]. During biosurfactant synthesis, P. aeruginosa effectively disrupted these matrices, potentially increasing mineral accessibility [50]. The mineral content of the substrates can also influence nutrient cycling in soil ecosystems [51]. Nevertheless, essential minerals involved in enzymatic processes critical for biosurfactant synthesis were not quantified in this study. A more comprehensive assessment of these micronutrients could provide insights into optimizing biosurfactant yields and support potential applications in food supplementation.
The lipid content of andiroba biomass is substantially higher than that of açai and babassu residues, reflecting its origin from oilseed processing. Andiroba oil is traditionally recognized for its insect-repellent properties and therapeutic effects against a range of ailments, including arthritis, diarrhea, diabetes, otitis, and throat inflammation [52,53,54]. In the context of biosurfactant production by P. aeruginosa, this elevated lipid content is critical, as lipid-rich substrates provide fatty acids that function as essential precursors for energy metabolism and biosurfactant synthesis [41,55]. The higher biosurfactant yield obtained from andiroba biomass compared to açai and babassu (Table 2) is likely attributable to this abundant lipid availability.
Efficient assimilation and metabolism of complex lipids by P. aeruginosa require initial enzymatic hydrolysis into simpler fatty acids [56]. This strain demonstrates a remarkable capacity to catalyze metabolic transformations, as evidenced by its proficiency in metabolizing complex organic compounds derived from petroleum, which underlies its hydrocarbon degradation efficiency [57]. Although lipid assimilation is generally slower than carbohydrate metabolism, the eight-day fermentation experiment under controlled conditions enabled effective utilization of andiroba lipids by P. aeruginosa P23G-02. Recent work from our group [16] further confirms the energy efficiency of andiroba biomass in supporting biosurfactant production by P. aeruginosa BM02, highlighting the critical role of lipid substrates in the biosynthesis process. Beyond biosurfactant production, andiroba biomass presents additional opportunities in biocombustibles, enzyme production, vitamin supplementation, antioxidant generation, livestock feed, antibiotic formulation, biofertilizers, and metabolite production via solid-state fermentation [58].
Proteins and their derivatives are essential for biosurfactant synthesis [14]. The elevated protein and lipid content in andiroba biomass residues likely supply P. aeruginosa P23G-02 with critical nutrients, enhancing metabolic activity and upregulating genes involved in biosurfactant biosynthesis, thereby explaining the higher yields obtained from this substrate. The protein matrix of these biomasses exhibits notable nutritional potential, consistent with previous reports on the viability of plant processing residues as protein sources, including essential amino acids [59,60]. The protein levels observed in this study align with existing recommendations, which vary according to age as per Brazilian legislation [61]. Furthermore, with each kilogram of wasted food protein contributing between 15 and 750 kg of CO2 emissions, the environmental impact on atmospheric quality is considerable [62]. Consequently, repurposing Amazonian fruit biomasses is expected not only to enhance biosurfactant production and mitigate food insecurity but also to contribute to environmental sustainability.
Carbohydrates were identified as the predominant component in both açai and babassu biomass, exceeding 80% of their composition. Plant carbohydrates include glucose, starch, cellulose, fructose, and hemicellulose, all of which serve as substrates for microbial fermentation [63]. Although extensive research has focused on conventional substrates such as glucose, glycerol, acetates, and organic acids, associated production costs and environmental sustainability concerns often present challenges [64]. Therefore, the use of renewable substrates, such as the carbohydrate-rich residual biomasses evaluated here, offers ecological benefits and cost efficiencies by valorizing these residues.
From a nutritional standpoint, carbohydrates are often scrutinized for potential health effects; however, they account for approximately 45–65% of human caloric intake [65]. Beyond their role in biosurfactant biosynthesis, carbohydrates derived from industrial biomass display distinctive physicochemical properties that are advantageous for diverse nutritional applications [66]. For instance, jackfruit residues, with carbohydrate and protein contents comparable to those observed in this study, are currently being explored for the production of nutritionally enhanced flour [67]. Likewise, avocado biomass residues, rich in carbohydrates and lipids, show potential as thickening and gelling agents in food products, with additional prospects for pharmaceutical applications and sustainable packaging solutions [68]. These observations support the notion that carbohydrates from the investigated Amazonian biomasses could serve as valuable substrates for developing a range of bioproducts with biotechnological and nutritional relevance.
The energy content of the Amazonian fruit biomass residues further underscores their intrinsic nutritional value, highlighting their role as nutrient-dense substrates capable of enhancing bacterial metabolism in various applications. The near-neutral pH levels observed are particularly significant, aligning with prior reports that demonstrate optimal biosurfactant production by P. aeruginosa under neutral pH conditions [17,69].
A critical implication of these findings for nutritional innovation lies in the redefinition of Amazonian fruit by-products: rather than being treated as waste, they can be considered valuable energy- and nutrient-rich resources. The high energy density and favorable macronutrient composition of these residues—including carbohydrate-rich açai and babassu, as well as lipid- and protein-rich andiroba (Table 3)—support diverse valorization pathways beyond biosurfactant production. These include direct nutritional applications, such as animal feed supplements or value-added ingredients for human consumption [8].
Within the broader framework of waste management and circular bioeconomy principles, the use of these residues for biosurfactant production represents a high-value application, particularly for materials that would otherwise contribute to environmental degradation and economic loss [14,15]. Ensuring safety is paramount for any direct nutritional application. This requires the use of non-pathogenic, food-grade microbial strains for fermentation or, when employing strains such as P. aeruginosa P23G-02, the implementation of rigorous purification protocols to remove microbial cells and potential toxins, ensuring the final product is safe for human or animal consumption.
Furthermore, evaluating the repurposing of Amazonian fruit biomass residues from a multi-stakeholder perspective—including local communities, industry participants, and policymakers—enhances the societal relevance of these findings. Such an approach fosters circular bioeconomies and bioproduct innovation while delivering environmental benefits, including reduced carbon footprints and improved soil health, aligning with global sustainability goals. Utilizing Generally Recognized as Safe (GRAS) microbial strains can mitigate inherent safety risks, simplify purification processes, lower production costs, and facilitate regulatory approval, thereby supporting the development of a sustainable, safe, and economically viable circular bioeconomy.
During chemical characterization of the fermentation products, FT-IR analysis (Figure 2) revealed key functional groups in biosurfactants produced by P. aeruginosa P23G-02. The O–H and C=O stretching signals indicate the presence of polar functional groups, contributing to enhanced emulsifying capacity and solubility in aqueous systems. C–H stretching signals confirm the presence of long hydrocarbon chains, while C–O stretching bands suggest incorporation of sugar moieties and ester groups, consistent with previous reports [17,28,70,71].
A distinct absorption band at 1714 cm−1, corresponding to C=O stretching of ester functionalities and potentially carboxylic acids, was exclusively observed in the biosurfactant derived from andiroba residues. This spectral feature serves as a structural marker for rhamnolipid configurations [72]. In contrast, biosurfactants produced from açai and babassu residues lacked this band. The presence of an additional rhamnose unit in di-rhamnolipids increases molecular hydrophilicity, positively influencing surface tension reduction and micelle stability [73], thereby explaining the superior performance indices observed for the andiroba biosurfactant.
The structural diversity of the biosurfactants was further confirmed through ESI-MS analysis (Figure 3), highlighting the range of functionalities and application potential of compounds derived from Amazonian biomasses. The combination of polar groups (hydrophilic components such as hydroxyls and carboxyls) and nonpolar alkyl chains (hydrophobic components) allows these biosurfactants to effectively reduce surface tension and stabilize emulsions under industrial conditions [73]. This structural variability facilitates the selection of optimal substrates for producing compounds with tailored functionalities suitable for specific applications.
In the context of nutritional innovations and food industry applications, these functional properties are highly relevant, providing both direct and indirect uses. Direct applications include incorporation into baking formulations, as demonstrated in patents by Haesendonck and Vanzeveren [74], enhancing the quality of fresh and frozen products. Indirect applications include biosurfactant-based formulations for cleaning and disinfecting food contact surfaces, as well as deodorizing and eradicating pathogenic microorganisms, as patented by DeSanto [75].
Additionally, the ability of biosurfactants to function within lipid nanocarrier systems enables efficient encapsulation and delivery of nutrients or bioactive compounds in fortified foods or dietary supplements, representing a significant advance in functional nutrition. Yang et al. [76] showed that biosurfactants used as gelling agents in cookies improved appearance, texture, mouthfeel, and storage stability. Studies by Azevedo et al. [77] and Cheng et al. [78] demonstrated the effectiveness of biosurfactants as surfactants in lipid-based nanocarrier systems across various food applications.
Moreover, the versatility of biosurfactants produced by P. aeruginosa extends to multiple industrial sectors, with a comprehensive summary of applications provided in Table 4. This broad applicability offers substantial opportunities for future research, including optimizing functional properties for existing applications, exploring novel uses in emerging fields, and expanding their integration into a circular bioeconomy. Such efforts could reinforce their role in sustainable development and maximize the valorization of these valuable Amazonian resources.
An interdisciplinary approach is essential for a comprehensive assessment of the potential of residual Amazonian fruit biomass in biosurfactant production and its implications for the food industry. Understanding the intricate interconnections among social, economic, and environmental factors is fundamental for discussions regarding biodiversity and sustainable development [95]. A critical issue in this domain is food waste, which remains largely unexploited. The Food and Agriculture Organization of the United Nations reported that, in 2018, approximately 1.3 billion tons of food were wasted globally, resulting in an estimated economic loss of roughly one trillion dollars. The United Arab Emirates exhibits the highest per capita food waste at 986 kg annually, whereas South Africa and Brazil report lower values of 172 kg and 71 kg per capita, respectively [96]. The relatively lower rates in Brazil can be attributed to successful campaigns and educational initiatives that have strengthened food security [97].
The United Kingdom has been recognized by Impact Economist [98] as a benchmark for food loss and waste mitigation, achieving a 27% reduction in 2018 relative to a baseline established in 2007. This success has been linked to the implementation of three strategic interventions: a voluntary scheme assisting businesses in reducing supply chain waste, the public awareness campaign “Love Food Hate Waste” providing practical guidance on recipes and food storage, and proactive measures promoting improved usability of food packaging among retailers and manufacturers.
Promoting a values-based approach to the utilization of plant-derived materials, particularly frequently overlooked biomass residues, may incentivize agricultural practices that optimize natural resource efficiency. This paradigm has the potential to reduce waste, improve systemic efficiency within the food chain, and generate new economic opportunities. Sustainable exploitation of Amazonian fruit biomass streams can provide critical insights and actionable solutions to address environmental degradation and food security challenges. Future research should further investigate the potential applications of Amazonian fruit residues, with particular emphasis on their nutritional and biotechnological properties.

5. Conclusions

This study demonstrates the substantial potential of Amazonian fruit biomass residues, including andiroba, açai, and babassu, as sustainable feedstocks for the production of biosurfactants by Pseudomonas aeruginosa P23G-02. Structural characterization of the resulting biosurfactants revealed a diverse array of configurations dependent on the biomass source, leading to variations in functionality and surface-active properties. These findings hold significant implications for the nutritional innovation sector and the food industry, particularly in regions with limited access to nutrient-rich foods. By promoting the incorporation of agricultural by-products into nutritional frameworks, this research supports sustainable practices that enhance socioeconomic resilience and stimulate bioprocess innovation. Engagement with local agricultural sectors through pilot projects may facilitate the translation of these findings into practical applications, aligning research outcomes with environmental and economic sustainability. The sustainable management principles and circular bioeconomy concepts highlighted in this study extend beyond the Brazilian Amazon, offering global relevance and the potential to generate considerable economic and ecological benefits. Furthermore, these insights may inform waste management policies at multiple governance levels, emphasizing the importance of integrating such strategies into broader sustainable development initiatives.
Critical limitations must be addressed to enable effective implementation. Future research should explore non-pathogenic biosurfactant-producing microorganisms to develop safe bioproducts suitable for direct application in the food and pharmaceutical sectors. Addressing scalability challenges will require studies employing bioreactor systems to optimize fermentation conditions and substrate compositions, thereby improving biosurfactant yields and economic feasibility. Collaboration with industry partners on pilot projects is essential to evaluate commercial viability and support the transition toward eco-friendly bioproducts and nutritional innovations, reinforcing circular bioeconomy principles. Additionally, the comparatively lower biosurfactant yields obtained from açai and babassu residues relative to andiroba underscore the need for optimization strategies tailored to these substrates. Future investigations should aim to overcome these limitations to fully exploit the potential of all biomass residues examined.

Author Contributions

A.M.F.: Formal analysis, Investigation, Methodology. L.M.S.P., J.d.A.A., G.S.G., E.O.R., K.L., J.A.P.B. and S.P.F.: Investigation. C.M.Q., A.O. and E.C.d.S.: Data curation, Formal analysis, Supervision, Validation, Writing—review and editing. G.C.d.S.S.: Conceptualization, Data curation, Formal analysis, Methodology, Validation, Writing—original draft. S.C.d.S.: Conceptualization, Data curation, Funding acquisition, Methodology, Project administration, Resources, Supervision, Validation, Writing—original draft, review and editing. All authors have read and agreed to the published version of the manuscript.

Funding

The authors acknowledge the financial support provided by the Coordination for the Improvement of Higher Education Personnel (CAPES, Financial Code 001), the National Council for Scientific and Technological Development (CNPq, Universal Project 2016), the Amazon Foundation for the Support of Studies and Research (Fapespa, Amazon+10 Project CONFAP 102/2023), and Instituto Vale. This support was instrumental in advancing the present research. CMQ acknowledges a productivity scholarship in technological development research from the National Council for Scientific and Technological Development of Brazil (CNPq).

Data Availability Statements

All data from this research are included in the article. Further inquiries can be directed to the corresponding author.

Conflicts of Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Figure 1. Phylogenetic tree based on 16S rRNA sequences from strain P23G-02 and Pseudomonas sp.
Figure 1. Phylogenetic tree based on 16S rRNA sequences from strain P23G-02 and Pseudomonas sp.
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Figure 2. FT-IR spectra of the biosurfactants produced by Pseudomonas aeruginosa P23G-02 using babassu, açai, and andiroba biomass residues. The dashed line highlights the absorption band at 1714 cm−1, a key structural marker indicative of di-rhamnolipid configurations in the biosurfactants.
Figure 2. FT-IR spectra of the biosurfactants produced by Pseudomonas aeruginosa P23G-02 using babassu, açai, and andiroba biomass residues. The dashed line highlights the absorption band at 1714 cm−1, a key structural marker indicative of di-rhamnolipid configurations in the biosurfactants.
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Figure 3. ESI-MS spectrum (negative mode) of the biosurfactants produced by Pseudomonas aeruginosa P23G-02 from andiroba (black tone), açai (red tone), and babassu (blue tone) biomass residues.
Figure 3. ESI-MS spectrum (negative mode) of the biosurfactants produced by Pseudomonas aeruginosa P23G-02 from andiroba (black tone), açai (red tone), and babassu (blue tone) biomass residues.
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Table 1. Identity of 16S rRNA sequences of strain P23G-02 and Pseudomonas sp. retrieved from NCBI.
Table 1. Identity of 16S rRNA sequences of strain P23G-02 and Pseudomonas sp. retrieved from NCBI.
NCBI CodeBacterium123456789
1NR_041702.1Pseudomonas knackmussii B1310095.9295.0497.9596.0496.1196.4496.4795.96
2NR_179771.1Pseudomonas lalkuanensis PE0895.9210098.6095.9897.5997.9398.5298.4798.39
3NR_181770.1Pseudomonas boanensis DB195.0498.6010096.1897.9097.5197.9897.9298.03
4NR_180597.1Pseudomonas nicosulfuronedens LAM 190297.9595.9896.1810097.9097.1196.7196.6196.78
5NR_179382.1Pseudomonas tohonis TUM 1899996.0497.5997.9097.9010098.5098.0597.9998.10
6NR_043289.1Pseudomonas otitidis MCC 1033096.1197.9397.5197.1198.5010098.5298.6298.62
7NR_114471.1Pseudomonas aeruginosa ATCC 1014596.4498.5297.9896.7198.0598.5210099.9399.87
8-P23G-0296.4798.4797.9296.6197.9998.6299.93100100
9NR_117678.1Pseudomonas aeruginosa DSM 5007195.9698.3998.0396.7898.1098.6299.87100100
(-): Not applied.
Table 2. Production and physicochemical properties of the biosurfactants from Pseudomonas aeruginosa P23G-02.
Table 2. Production and physicochemical properties of the biosurfactants from Pseudomonas aeruginosa P23G-02.
Biosurfactant/ControlEI (%)ST (mN/m)YD (mg/mL)
Açai biosurfactant56.0 ± 2.0035.7 ± 0.310.8 ± 0.22
Babassu biosurfactant54.1 ± 1.0031.6 ± 0.281.6 ± 0.36
Andiroba biosurfactant60.8 ± 2.0129.0 ± 0.085.6 ± 0.43
Non-inoculated mineral saline medium0.0 ± 0.0069.9 ± 0.740.0 ± 0.00
1% sodium dodecyl sulfate69.9 ± 1.10NtNt
(EI): emulsification index, expressed as a percentage (%); (ST): surface tension, expressed in mN/m; (YD): yield, expressed in mg/mL; (Nt): not tested. Data were obtained in triplicate (n = 3) and are presented as arithmetic mean ± standard deviation.
Table 3. Proximate composition of Amazonian fruit biomass residues used for biosurfactant production.
Table 3. Proximate composition of Amazonian fruit biomass residues used for biosurfactant production.
ParametersAçaiBabassuAndiroba
Moisture (%)8.81 ± 0.0510.05 ± 0.032.75 ± 0.03
Ash (%)1.31 ± 0.221.25 ± 0.184.42 ± 0.08
Total lipids (%)3.72 ± 0.480.98 ± 0.1857.02 ± 0.58
Total proteins (%)3.73 ± 0.072.51 ± 0.0910.76 ± 0.23
Total carbohydrates (%)82.43 ± 0.4685.21 ± 0.2525.06 ± 0.46
pH5.00 ± 0.075.91 ± 0.116.74 ± 0.14
Energetic value (Kcal/100 g) 378.12359.70656.46
Data are presented as mean ± standard deviation, with measurements performed in triplicate (n = 3). All compositional parameters, except pH and energy value (kcal/100 g), are expressed as percentages (%).
Table 4. Industrial applications of rhamnolipid biosurfactants produced by different strains of Pseudomonas aeruginosa.
Table 4. Industrial applications of rhamnolipid biosurfactants produced by different strains of Pseudomonas aeruginosa.
Industry/ActivityP. aeruginosa StrainReference
Pharmaceutical Industry
AntitumorPAO1[79]
BM02[17]
ImmunomodulationMc210[80]
AntifungalA4[81]
ZJU211[82]
AntibiofilmJS29[83]
UKMP14T[84]
Nanoparticles for drug deliverySP4[85]
BS01[86]
Wound healingC2[87]
JS29[83]
Skin treatmentATCC 27853[88]
Automotive Industry
Mechanical response on automotive, railway and aeronautical materialsUnidentified strain[89]
Steel Industry
Steel corrosion inhibitionATCC 9027, LFM634, and ATCC 9027[90]
Petrochemical Industry
Bioremediation of soil contaminatedBM02[91]
LBI[92]
PTCC 1340[93]
Timber Industry
Wood adhesivesATCC 9027[94]
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Feio, A.M.; Sá, G.C.d.S.; Orsato, A.; Leite, K.; Pimentel, L.M.S.; Alves, J.d.A.; Gomes, G.S.; Ramos, E.O.; Quintella, C.M.; Fragoso, S.P.; et al. Valorization of Amazonian Fruit Biomass for Biosurfactant Production and Nutritional Applications. Biomass 2025, 5, 60. https://doi.org/10.3390/biomass5040060

AMA Style

Feio AM, Sá GCdS, Orsato A, Leite K, Pimentel LMS, Alves JdA, Gomes GS, Ramos EO, Quintella CM, Fragoso SP, et al. Valorization of Amazonian Fruit Biomass for Biosurfactant Production and Nutritional Applications. Biomass. 2025; 5(4):60. https://doi.org/10.3390/biomass5040060

Chicago/Turabian Style

Feio, Alan Moura, Giulian César da Silva Sá, Alexandre Orsato, Karoline Leite, Lucas Mariano Siqueira Pimentel, Joane de Almeida Alves, Glenda Soares Gomes, Evelly Oliveira Ramos, Cristina M. Quintella, Sinara Pereira Fragoso, and et al. 2025. "Valorization of Amazonian Fruit Biomass for Biosurfactant Production and Nutritional Applications" Biomass 5, no. 4: 60. https://doi.org/10.3390/biomass5040060

APA Style

Feio, A. M., Sá, G. C. d. S., Orsato, A., Leite, K., Pimentel, L. M. S., Alves, J. d. A., Gomes, G. S., Ramos, E. O., Quintella, C. M., Fragoso, S. P., Bitencourt, J. A. P., da Silva, E. C., & Santos, S. C. d. (2025). Valorization of Amazonian Fruit Biomass for Biosurfactant Production and Nutritional Applications. Biomass, 5(4), 60. https://doi.org/10.3390/biomass5040060

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