Essential oils from Eucalyptus species: a review of their activities, applications, and the Brazilian market

Template: Editora Letra1 | www.editoraletra1.com.br This is an open-access article distributed under the terms of the Creative Commons Attribution License. | 1 Essential oils from Eucalyptus species: a review of their activities, applications, and the Brazilian market Mayara Rodrigues1 , Paulo Mazzafera1,2*  1Universidade de São Paulo, Escola Superior de Agricultura Luiz de Queiroz (ESALQ), Departamento de Produção Vegetal, Piracicaba, SP, Brazil. 2Universidade Estadual de Campinas, Instituto de Biologia, Departamento de Biologia Vegetal, Campinas, SP, Brazil. *Corresponding author: [email protected] ABSTRACT Essential oils are volatile, lipophilic substances mainly composed of terpenes, derived from secondary metabolism. Widely used in pharmaceutical, agricultural, and cosmetic industries, essential oils from eucalyptus species are particularly notable for their economic and biological significance. This review explores the antimicrobial, anti-inflammatory, antioxidant, antiviral, and insecticidal activities of essential oils from various eucalyptus species and their isolated constituents, such as eucalyptol and α-terpineol. Because eucalyptus represents 96.7% of the reforested area in Brazil, economic analyses focused on the Brazilian market, highlighting a 25-year trajectory of production and trade, the dominance of eucalyptolrich oils, and the growing demand for sustainable and high-quality products. Future market prospects include expanding the use of underutilized species, improving oil formulations for enhanced stability and bioavailability, and meeting stringent international quality standards. Addressing these challenges and opportunities underscores the essential role of ongoing research in leveraging eucalyptus essential oils as versatile and sustainable bioresources. Keywords: composition, essential oils, Myrtaceae, sustainability, terpenes Introduction The Eucalyptus genus is the most widely cultivated forest genus worldwide (Mateus et al., 2021), and the area of plantation has exceeded 22.57 million ha (Hua et al., 2022). According to the Brazilian Institute of Geography and Statistics (IBGE, 2024), the planted forests in Brazil reached 9.7 million hectares in 2023, with eucalyptus and pine together accounting for covering 96.3% of this area. Eucalyptus alone covers 7.6 million hectares, representing 78.4% of the total area. Eucalyptus is used mainly for the pulp and paper industry and Brazil is the world’s largest exporter of pulp, with the top destinations being China, the United States, the Netherlands, and Italy. Eucalyptus forests serve various purposes, from wood use per se to the extraction of essential oils for use in the pharmaceutical and chemical industries, but the primary use of eucalyptus is as raw material for paper and pulp production. More recently, there has been increased interest in eucalyptus as an Review doi: https://doi.org/10.1590/1677-941X-ABB-2024-0111 Acta Botanica Brasilica , 2025, 39: e20240111 Received May 02, 2024; Accepted April 06, 2025 Editor-in-Chief: Thaís Elias Almeida; Associate Editor: Moemy Moraes How to cite: Rodrigues M, Mazzafera P. 2025. Essential oils from Eucalyptus species: a review of their activities, applications, and the Brazilian market. Acta Botanica Brasilica 39: e20240111. doi: 10.1590/1677-941X-ABB-2024-0111 Template: Editora Letra1 | www.editoraletra1.com.br Rodrigues M, Mazzafera P 2 | Acta Botanica Brasilica, 2025, 39: e20240111 energy crop due to its rapid growth and biomass accumulation (Pérez-Cruzado et al., 2011). Tab. 1 shows the uses of several eucalyptus species. The superior silvicultural characteristics of eucalyptus compared to other species are what distinguish the genus. These include rapid growth rate, upright stem, ability to adapt to a wide range of climates and soils, and good wood quality for pulp production (Grattapaglia & Kirst, 2008; Moura et al., 2010). The term Eucalypt includes species from the genera Eucalyptus L’Hérit., Corymbia Hill and Johnson, and Angophora Cav., with the latter two no longer being subgenera of the first, but rather referred to as genera based on molecular studies (Grattapaglia & Kirst, 2008). The centre of origin for the Eucalyptus genus is Australia and nearby islands such as Papua New Guinea, Timor, Indonesia, Tasmania, and the Philippines, providing different species of these genera with adaptation to various soil and climatic conditions, with each species having specific characteristics (Ladiges & Wasshausen, 1996). The term “Eucalyptus” refers to a group of more than 700 species and varieties that belong to the genera Eucalyptus and Corymbia (Barbosa et al., 2021). Table 1. Primary uses of the studied Eucalyptus species Species Purpose Eucalyptus camaldulensis Flowers for honey production Sawmill, sleepers, charcoal, posts, stakes, and firewood Eucalyptus dunnii Firewood, civil construction (shores), sawmilling/lamination, posts/stakes/posts, cellulose and paper Eucalyptus exserta Heavy, hard, and durable wood, useful in constructions Eucalyptus globulus Essential oil Eucalyptus grandis Firewood, civil construction (shores), sawmilling/lamination, posts/stakes/posts, cellulose and paper Corymbia henryi Flowers for honey production, wood for constructions and tool handles Eucalyptus pellita Flowers for honey production Wood for constructions and structures Eucalyptus resinifera Sawmill, mobile constructions, crates, sleepers, posts, and stakes Eucalyptus saligna Firewood, civil construction (shores), sawmilling/lamination, posts/stakes/posts, cellulose and paper, furniture, structures, crates, charcoal Eucalyptus urophylla Firewood, charcoal, civil construction (shores), sawmilling/lamination, posts/stakes/posts, Flowers for honey production Source: (Ferreira, 1979; CABI, 2019a; b; Embrapa, 2022) There is significant interest in natural products for the development of new products for medical, agrochemical, and cosmetics applications. This demand has increased due to the need for safer raw materials. Additionally, the demand also responds to the rise in cases of microbial resistance and the emergence of pests and diseases without known control measures. Because of these challenges, the essential oils (EOs) market has grown considerably in recent years (Bizzo & Rezende, 2022), particularly essential oils from eucalyptus. According to ISO 9235 of the International Organization for Standardization (ISO 9235, 2021), “essential oil is a product obtained from natural raw plant material by steam distillation, by mechanical processes from the epicarp of citrus fruits, or by dry distillation after the separation of the aqueous phase - if present - by physical processes.” It is important to complement this definition by stating that aromatic products obtained by other extraction techniques, such as solvents, waxes, fats, supercritical fluids, headspace techniques, or any other means are not considered essential oils and have other designations as described in the same technical standard (Bizzo & Rezende, 2022). The extraction of Eucalyptus oil is carried out by hydrodistillation, thus making it an essential oil. Essential oils are volatile, lipophilic substances originating from the secondary metabolism, composed of bioactive volatiles, in which terpenes are the largest group of components, classified according to their number of carbon atoms as monoterpenes, sesquiterpenes and diterpenes (Dhakad et al., 2018; Koyama & Heinbockel, Essential oils from Eucalyptus Template: Editora Letra1 | www.editoraletra1.com.br Acta Botanica Brasilica, 2025, 39: e20240111 | 3 2020). In plants, volatile terpene roles are not fully understood, although they have several functions highlighted. EOs contribute to attracting pollinators and seed dispersers, serve as a defence mechanism against phytopathogenic organisms, and exhibit allelopathic effects, influencing the growth and development of neighbouring plants. These multifaceted roles underscore their ecological and industrial significance (Zeroual et al., 2021). The constituents of Eucalypt oil are predominantly volatile terpenes and aromatic compounds (Aldoghaim et al., 2018). Paul et al. (2020) reported the presence of terpenes, mainly monoterpenes (camphene, isocamphene, p-cymene, fenchone, fenchol, camphor, isoborneol, and borneol), preserved in amber from the late Oligocene, showing that the synthesis mechanism of these constituents evolved in the Cenozoic Era. Terpenes represent the largest group of secondary compounds. They are classified by the number of isoprene units they possess, synthesized in the cytoplasm and/or plastids of plant cells from the mevalonic acid (MEV) pathway in the cytosol and the methylerythritol phosphate (MEP) pathway in the chloroplast (Taiz et al., 2017). Depending on the species, essential oils can be produced in different parts of plants, such as flowers, leaves, stems, roots, fruits, and secretory structures, where they are also stored (Barbosa et al., 2016). Eucalypt essential oils are mainly produced and stored in leaves (Filomeno et al., 2016). Oil cavities are characteristic structures in Myrtaceae genera (Al-Edany & Al-Saadi, 2012), with variable distribution and size (Döll-Boscardin et al., 2010). The chemical composition, yield, and effectiveness of essential oils vary between and within species (Mugao, 2024). They are influenced by both endogenous and exogenous factors. Endogenous factors include plant characteristics such as age, density, plant part used, and genotype, which directly affect the synthesis and chemical variation of essential oils. Exogenous factors include environmental variables like light intensity, temperature, water availability, altitude, latitude, soil composition, extraction method, and storage (Mugao, 2024). The combination of these factors determines the final quality and effectiveness of the oil. Eucalypt essential oils (EOs) are classified into medicinal, industrial, and perfumery categories based on specific constituents. They are used in the pharmaceutical and food industries, for therapeutic activities, and serve as raw materials for producing cosmetics, perfumes, cleaning products, insecticides, and herbicides. They are easily biodegradable, allowing their use in public places with extensive exposure, such as schools and hospitals (Dhakad et al., 2018). EO production also enables greater profitability for eucalyptus, primarily used for wood and cellulose exploration, where the leaves are discarded (Filomeno et al., 2016). Some factors hinder the use of essential oils in commercial products, such as their standardization. Differences in the composition of oils from plants of the same species are observed depending on biotic and abiotic factors such as climatic conditions, light, soil type and conditions, agronomic management, plant phenological stage, in addition to the drying methods, extraction, and storage conditions used (Warnke et al., 2009; Barbosa et al., 2016; Bett et al., 2016; Salem et al., 2018). These varying conditions, combined with the genetic background of plants, can influence the activation or inhibition of metabolic pathways in aromatic plants (Knezevic et al., 2016). Consequently, biological activities may also vary as they depend on chemical composition (Barbosa et al., 2016). All these factors should be considered when seeking specific constituents. Two other factors that complicate using EOs in commercial products are the relatively rapid evaporation or degradation, which reduces effectiveness (Pavela, 2015). Terpene biosynthesis in plants Terpenes are present in almost all plants and they represent the largest and most diverse class of plant secondary metabolites. Terpenes volatiles are found in two terpene classes, monoterpenes and sesquiterpenes (Aqeel et al., 2023), which can have Template: Editora Letra1 | www.editoraletra1.com.br Rodrigues M, Mazzafera P 4 | Acta Botanica Brasilica, 2025, 39: e20240111 several important physiological roles in plants, like defence against herbivory, attraction of mutualists such as pollinators, disease resistance, plant-plant communication, antioxidants, etc. (Butler et al., 2018; Boncan et al., 2020; Rosenkranz et al., 2021). In general, stresses may change terpene plant composition and emission rates (Copolovici & Niinemets, 2016) and their profile can even vary within species and among individual plants (Kopaczyka et al., 2020). Terpenes composition in Eucalyptus camaldulensis Dehnh., which has the largest range of distribution worldwide, both latitudinally and longitudinally, is influenced by environmental variables, mainly favouring the 1,8-cineole chemotype in arid locations (BustosSegura et al., 2017). Significant changes in essential oil relative composition were also found in E. camaldulensis (Leicach et al., 2010) and E. globulus (Queiroz et al., 2017) submitted to drought. Terpenes composition in eucalyptus may also change with plant age (Shiferaw et al., 2019). Terpene synthesis is controlled by the terpene synthase gene family, which is highly diversified throughout the plant kingdom (Butler et al., 2018). The biosynthesis of volatile organic compounds (VOCs) such as terpenes is limited by the enzymes involved in the process and the amount of substrate available for their functioning. Some enzymes require specific substrates, while others can synthesize various products from a single substrate (Vattekkatte et al., 2018). Analyzing the genomes of two eucalyptus species (E. globulus and E. grandis), Külheim et al. (2015) confirmed that this diversity of terpenes is partially attributable to the largest family of terpene synthase (TPS) genes ever described. Terpene synthase enzymes synthesize terpenes from isopentenyl diphosphate (IPP) and dimethylallyl diphosphate (DMAPP), which originate from the mevalonic acid (MEV) pathway in the cytosol and the methylerythritol phosphate (MEP) pathway in the plastids (Butler et al., 2018). The synthesis of monoterpenes occurs in plastids (via methylerythritol 4-phosphate - MEP) from D-glyceraldehyde-3-phosphate pyruvate, while sesquiterpenes are synthesized in the cytosol (via mevalonate - MVA) from acetyl-CoA (Rajčević et al., 2019), as illustrated in Fig. 1. In several plants, the two enzymes involved in the initial steps of the MEP pathway have been shown to influence the foliar yield of terpenes (Xu et al., 2019). However, according to results obtained by Külheim et al. (2015), the foliar concentration of monoterpenes in eucalyptus appears to be influenced by the two final steps of the MEP pathway. For sesquiterpenes in the MVA pathway, the metabolite flow seems influenced by 3-hydroxy-3-methylglutaryl-CoA synthase – HMGS (Külheim et al., 2015). Several factors influence the ratio of monoand sesquiterpenes in leaves, such as substrate availability for geranyl pyrophosphate synthase (GPPS) in the chloroplast and for farnesyl pyrophosphate synthase (FPPS) in the cytosol. Both enzymes use IPP (isopentenyl pyrophosphate) and DMAPP (dimethylallyl pyrophosphate) in different proportions (Bouvier et al., 2000; Külheim et al., 2015; Abdoul-Latif et al., 2023). IPP and DMAPP react to produce geranyl pyrophosphate (GPP), serving as a precursor to monoterpenes. When GPP interacts with an additional IPP molecule, it forms farnesyl pyrophosphate (FPP), a sesquiterpene precursor. Similarly, the reaction of FPP with a third IPP molecule results in geranylgeranyl pyrophosphate (GGPP), which acts as a precursor for diterpenes (Abdoul-Latif et al., 2023). Typically, oxygenated monoterpenes predominate in the composition of essential oils (Dhifi et al., 2016), however, there are plants in which non-oxygenated terpenes are more abundant. Biological activities of eucalyptus essential oils Essential oils are increasingly recognized and utilized for numerous purposes across different sectors. Their lipophilic nature allows interactions with the cell plasma membrane, making it permeable. Moreover, they can depolarize the mitochondrial membrane, rendering it permeable and potentially leading to toxicity (Gautam et al., 2014; Abdoul-Latif et al., 2023). Another relevant point is that terpenes can easily traverse cell Essential oils from Eucalyptus Template: Editora Letra1 | www.editoraletra1.com.br Acta Botanica Brasilica, 2025, 39: e20240111 | 5 Figure 1. Synthesis of volatile terpenes (highlighted in yellow) through the mevalonate (MVA) pathway in the cytosol and the methylerythritol 4-phosphate (MEP) pathway in the plastid. The MVA pathway gives rise to sesquiterpenes, while the MEP pathway gives rise to monoterpenes. There is the export of isopentenyl pyrophosphate (IPP) from the plastid to the cytosol, with solid arrows indicating well-established steps and the dashed arrow representing hypothetical reactions. Enzymes in blue are located in the peroxisome, and those in red are in the endoplasmic reticulum. Abbreviations: AACT, acetyl-CoA acetyltransferase; HMGS, HMG-CoA synthase; HMG-CoA, hydroxymethylglutaryl-CoA; HMGR, HMG-CoA reductase; MVK, mevalonate kinase; MVP, mevalonate 5-phosphate; PMK, phosphomevalonate kinase; MVPP, mevalonate 5-pyrophosphate; MPDC, mevalonate diphosphate decarboxylase; IPP, isopentenyl pyrophosphate; IDI, isopentenyl pyrophosphate isomerase; DMAPP, dimethylallyl pyrophosphate; FPPS, FPP synthase; FPP, farnesyl pyrophosphate; TPS, terpene synthase; G3P, glyceraldehyde 3-phosphate; DXS, DXP synthase; DXP, 1-deoxy-D-xylulose 5-phosphate; DXR, 1-deoxy-D-xylulose 5-phosphate reductoisomerase; MCT, 2-C-methyl-D-erythritol 4-phosphate cytidylyltransferase; CDP-ME, 4-diphosphocytidyl-2-C-methyl-D-erythritol; CMK, 4-(cytidine 5’-diphospho)-2-C-methyl-D-erythritol kinase; CDP-MEP, CDPME 2-phosphate; MECPS, MECPD synthase; MECDP, 2-C-methyl-D-erythritol 2,4-cyclodiphosphate; HDS, 4-hydroxy-3-methylbut-2en-1-yl diphosphate synthase; HMBPP, (E)-4-hydroxy-3-methylbut-2-en-1-yl diphosphate; IDS, isopentenyl diphosphate synthase; GGPPS, GGPP synthase; GGPP, geranylgeranyl pyrophosphate; GPPS, GPPS synthase; GPP, geranyl pyrophosphate. Source: Adapted from Dudareva et al. (2013). membranes due to their low molecular weight, thereby inducing biological activities such as antimicrobial and antioxidant effects (Al-Radadi, 2022). Defining the modes and mechanisms of action determining the efficacy of essential oils is challenging due to their complex and variable composition, requiring further studies. Many studies focus on the use of EOs as potential antimicrobials because conventional chemicals typically lose their efficacy due to misuse, the development of microbial resistance, or the emergence of microorganisms that require new means of control. In addition to the resistance mechanisms developed by microorganisms, there Template: Editora Letra1 | www.editoraletra1.com.br Rodrigues M, Mazzafera P 6 | Acta Botanica Brasilica, 2025, 39: e20240111 is the risk of the resistance factor being transmitted to pathogenic descendants (Salem et al., 2018). The increased use of EOs in agricultural pest management is also justified by changes in pesticide market regulations in the European Union (EU) (Directive 2009/128/EC), which mandate the implementation of Integrated Pest Management (IPM) and the sustainable use of pesticides, especially non-synthetic ones, for weed control (Verdeguer et al., 2020b). A comprehensive review of the bioherbicidal potential of EOs from various weed species was conducted by Verdeguer et al. (2020b). These authors highlighted two challenges for evaluating EOs in vivo: they must be mixed with emulsifiers and not with water, making the type of emulsifier another factor to be tested, and rapid volatilization requires appropriate formulations, with microencapsulation and nanoemulsion being the most commonly used for EOs (Vinceković et al., 2017). Tab. 2 shows the biological activities of eucalyptus EOs published in the literature. The eucalyptus oils have a large range of bioactivity probably because of their diverse composition. The bioactivities cover from agriculture applications to the control of human pathogens. Other eucalyptus species from which essential oil is extracted for medicinal use include E. maidenii F. Muell, E. bicostata Maiden, Blakely & Simmond, E. sideroxylon A. Cunn. ex Woolls, E. cinerea F. Muell. ex Benth., E. leucoxylon F. Muell., and E. tereticornis Sm. (Mieres-Castro et al., 2021). Formulations that yield satisfactory results have to meet environmental and safety concerns. Pant et al. (2014) prepared a nanoemulsion using aqueous filtrates of Pongamia glabra Vent. and Jatropha curcas L. left after biodiesel oil extraction to enhance the activity of Eucalyptus globulus Labill. oil in controlling Tribolium castaneum (Herbst), a pest of stored grains. The product was effective in control, with karanjin, phorbol esters, and eucalyptol being the main constituents responsible for the observed insecticidal effect. It is worth noting that such products improve the performance of EOs and can be used on food grains, not polluting the environment and expanding the range of pesticide products, reducing resistance risks. Table 2. Biological effects of the essential oils of different Eucalyptus species, described in the literature. Effect Reference Eucalyptus camaldulensis Dehnh Activity against gram-positive and gram-negative bacteria (Salem et al., 2015) Inhibition of the growth of fungi Alternaria alternata and Chaetomium globosum in commercial wood (Salem et al., 2016) Antibacterial against Bacillus subtilis, Listeria monocytogenes, Staphylococcus aureus, Yersinia enterocolitica, Salmonella Paratyphi, Salmonella typhi, Salmonella typhimurium, Vibrio vulnificus, Escherichia coli, and Pseudomonas aeruginosa (Debbarma et al., 2013) Repellent against Sitophilus zeamais (Karemu et al., 2013) Antibacterial against Staphylococcus aureus, Bacillus subtilis and Escherichia coli Antifungal against Aspergillus niger and Rhizopus solani Antioxidant for linoleic acid (Ghaffar et al., 2015) Insecticide against Ectomielois ceratoniae (Jemâa et al., 2013) Insecticide against Callosobruchus maculatus, Sitophilus oryzae and Tribolium castaneum (Negahban & Moharramipour, 2007) Antibacterial against Acinetobacter baumannii isolates (Knezevic et al., 2016) Inhibition of germination and growth of invasive species Amaranthus hybridus L. and Portulaca oleracea L. (Verdeguer et al., 2009) Antibacterial Against Escherichia coli ATCC 25922, Staphylococcus aureus ATCC 29213 and Enterococcus faecalis ATCC 29212 Moderate antibacterial effect against Pseudomonas aeruginosa ATCC 27853 (Tine et al., 2022) Essential oils from Eucalyptus Template: Editora Letra1 | www.editoraletra1.com.br Acta Botanica Brasilica, 2025, 39: e20240111 | 7 Effect Reference Antibacterial against Bacillus cereus, Citrobacter diversus, Klebsiella oxytoca, Klebsiella pneumoniae, Proteus vulgaris, Shigella flexneri Antifungal against Aspergillus clavatus, Candida albicans, Cladosporium cladosporioides, Lenzites sulphureus, Myrothecium verrucaria, Penicillium citrinum, Phanerochaete chrysosporium, Phaeolus schweintizii, Trametes versicolor, Trichoderma viride Acaricidal against Varroa destructor Insecticidal against Aedes aegypti, Aedes albopictus, Atta sexdens rubropilosa, Ephestia cautela, Ephestia kuehniella, Pediculus humanus capitis, Thyrinteina arnobia (Barbosa et al., 2016) Herbicide against Erigeron bonariensis L. (Verdeguer et al., 2020a) Eucalyptus dunnii Maiden Repellent and insecticide against Sitophilus zeamais (Mossi et al., 2011) Insecticide against Aedes aegypti, Blattella germanica, Pediculus humanus capitis (Barbosa et al., 2016) Eucalyptus exserta F.Muell Inhibition of gram-negative bacteria Salmonella enteritidis (Ambrosio et al., 2017) Fumigant against Tribolium castaneum (Li & Xu, 2012) Fungicide against Aspergillus flavus and Aspergillus niger (Oanh & Giang, 2017) Eucalyptus globulus Labill Activity against larvae and pupae of Musca domestica (Kumar et al., 2012) Termiticidal activity against termites Odontotermes assamensis (Pandey et al., 2012) Repellent against Sitophilus zeamais (Karemu et al., 2013) Potential control of dental plaque by Streptococcus mutans (Goldbeck et al., 2014) Antibacterial against Staphylococcus aureus, Bacillus subtilis and Escherichia coli Antifungal against Aspergillus niger and Rhizopus solani Antioxidant for linoleic acid (Ghaffar et al., 2015) Insecticide and ovicide against Pediculus humanus capitis (Yones et al., 2016) Antibacterial against Staphylococcus aureus, Streptococcus pyogenes, Staphylococcus epidermidis, Salmonella typhi, Escherichia coli, Shigella spp. And Pseudomonas aeruginosa Antifungal against Trichophyton spp1 and Aspergillus spp (Mekonnen et al., 2016) Antioxidant – radical-scavenging properties and ability to inhibit lipid peroxidation Antibacterial – synergistic effect between the oil and conventional antibiotics against Acinetobacter baumannii (Luís et al., 2016) Antibacterial against Staphylococcus aureus, Escherichia coli and Pseudomonas aeruginosa (Pombal et al., 2014) Antibacterial against Staphylococcus aureus Wound healing in rats (Sugumar et al., 2014) Repellent and insecticide against Sitophilus zeamais (Mossi et al., 2011) Antimicrobial against multidrug-resistant bacteria (Mulyaningsih et al., 2011) Strong antimicrobial, especially against Streptococcus pyogenes, Escherichia coli, Candida albicans, Staphylococcus aureus, Acinetobacter baumannii and Klebsiella pneumoniae (Damjanović-Vratnica et al., 2011) Antibacterial against gram-negative periodontopathogenic bacteria, mainly Fusobacterium nucleatum ATCC25586 and Porphyromonas gingivalis ATCC33277 (Harkat-Madouri et al., 2015) Insecticide against Triboliu confusum (oil extracted from young leaves had a better effect compared to oil from mature leaves) (Russo et al., 2015) Inhibition of germination and development of invasive plants Amaranthus blitoides and Cynodon dactylon (Rassaeifar et al., 2013) Antioxidant through the reduction of metal ions that perpetuate the formation of free radicals, induce lipid peroxidation, and consequently disrupt the homeostatic balance of human cells (Salem et al., 2018) Table 2. Cont. Template: Editora Letra1 | www.editoraletra1.com.br Rodrigues M, Mazzafera P 8 | Acta Botanica Brasilica, 2025, 39: e20240111 Effect Reference Antibacterial against Bacillus cereus, Citrobacter diversus, Enterococcus faecalis, Fusobacterium nucleatum, Klebsiella oxytoca, Klebsiella pneumoniae, Porphyromonas gingivalis, Pseudomonas fluorescens, Salmonella paratyphi, Salmonella typhimurium, Staphylococcus intermedius, Staphylococcus sciuri, Staphylococcus warneri Antifungal against Aspergillus flavus, Aspergillus parasiticus, Aspergillus spp., Candida albicans, Fusarium oxysporum, Mucor spp., Penicillium digitatum, Rhizopus nigricans, Saccharomyces cerevisiae Acaricidal against Boophilus microplus Insecticidal against Aedes aegypti, Lutzomyia longipalpis, Sitophilus oryzae, Tribolium castaneum (Barbosa et al., 2016) Inhibition of growth of gram-positive against Sthaphylococcus aureus, gram-negative Proteus vulgaris and Escherichia coli, and the fungus Candida albicans.(Mota et al., 2015) Anti-influenza A activity (Vimalanathan & Hudson, 2014) Acaricidal and repellent activity against Rhipicephalus bursa (Madreseh-Ghahfarokhi et al., 2019) Activates Complement Receptor-Mediated Phagocytosis and Stimulates Podosome Formation in Human Monocyte-Derived Macrophages Zonfrillo et al., 2022 Eucalyptus grandis W.Hill ex Maiden Antibacterial against e Escherichia coli (ATCC 8739), Psuedomonas aeruginosa (ATCC 19582), Staphylococcus aureus (ATCC 6538), Streptococcus faecalis (ATCC 29212), Bacillus cereus (ATCC 10702), Bacillus Pumilus (ATCC 14884), Psuedomonas aeruginosa (ATCC 7700), Enterobacter cloacae (ATCC 13047), Klebsiella pneumonia (ATCC 10031), Bacillus subtilis (KZN), Shigella flexineri (KZN), Salmonella spp. (KZN), Staphylococcus epidermidis (KZN) and Enterococcus faecalis (KZN) Antibacterial against antibiotic-resistant species: Staphylococcus aureus (P12702/ P12763/ P12724/ B10808), Streptococcus viridans (S17141), Klebsiella spp. (S17302) and Klebsiella pneumonia (S17298) (Sewanu et al., 2012) Antibacterial against Staphylococcus aureus ATCC6538, Escherichia coli ATCC8739, Escherichia coli 0:158 and Salmonella choleraeseus ATCC10708 (Estanislau et al., 2001) Larvicidal against Aedes aegypti (Gallon et al., 2020) Antifungal against Aspergillus clavatus, Aspergillus niger, Chaetomium globosum, Cladosporium cladosporioides, Lenzites sulphureus, Myrothecium verrucaria, Penicillium citrinum, Phaeolus schweintizii, Phanerochaete chrysosporium, Trametes versicolor, Trichoderma viride Insecticidal gainst Aedes aegypti, Atta sexdens rubropilosa, Blattella gainsta, Pediculus humanus capitis, Thyrinteina arnobia (Barbosa et al., 2016) Eucalyptus pellita F.Muell Antibacterial against strains of Pseudomonas aeruginosa, Escherichia coli, Staphylococcus aureus and Bacillus subtilis (Proenza et al., 2013) Antibacterial against Streptococcus sobrinus and Streptococcus mutans, causative agents of dental caries (Kartiko et al., 2021) Eucalyptus resinifera Sm Fumigant and larvicidal against Aedes aegypti (Lucia et al., 2012) Insecticidal against Rhyzopertha dominica (more toxic to insects than the commercial insecticide pirimiphos-methyl) (Filomeno et al., 2020) Fumigant and repellent against Hypothenemus hampei (Reyes et al., 2019) Insecticidal against Haematobia irritans (Barbosa et al., 2016) Eucalyptus saligna Sm Repellent against Sitophilus zeamais (Karemu et al., 2013) Repellent and insecticidal against Sitophilus zeamais (Mossi et al., 2011) Antimicrobial against bacteria Staphylococcus aureus and Escherichia coli and fungi Candida albicans and Phytophthora cactorium (Sartorelli et al., 2007) Insecticidal and repellent against Sitophilus zeamais and Tribolium confusum in grain storage (Tapondjou et al., 2005) Antibacterial against Bacillus cereus, Bacillus subtilis, Citrobacter diversus, Klebsiella oxytoca, Klebsiella pneumoniae, Pseudomonas aeruginosa, Salmonella choleraesuis Insecticidal against Aedes aegypti, Atta sexdens rubropilosa, Pediculus humanus capitis, Thyrinteina arnobia (Barbosa et al., 2016) Table 2. Cont. Essential oils from Eucalyptus Template: Editora Letra1 | www.editoraletra1.com.br Acta Botanica Brasilica, 2025, 39: e20240111 | 9 Eucalyptus essential oil was trapped within the micro-nanostructures of a polydimethylsiloxane (PDMS) substrate, which increased significantly the hydrophobicity (Hidouri et al., 2024). The authors demonstrated the antibacterial ability of these superhydrophobic surfaces to inhibit the growth of Escherichia coli (Migula) Castellani & Chalmers and Bacillus cereus Frankland & Frankland bacterial colonies and biofilms. Due to the complexity of EOs composition, relating observed effects to only one constituent is still a challenging task, as additive, synergistic, or antagonistic effects may occur with their combinations (Yap et al., 2014; Filomeno et al., 2020; Verdeguer et al., 2020b). While some researchers associate the fumigant effect of eucalyptus EOs against grain pests with its eucalyptol content (Aref et al., 2015; Hamdi et al., 2015), Filomeno et al. (2020) found that eucalyptus EOs with approximately 70% eucalyptol showed intermediate mortality, suggesting that other constituents may promote antagonistic effects. When evaluating the antibacterial efficacy of eucalyptus EOs compared to eucalyptol, its dominant constituent, Hendry et al. (2009) found better results with the oil, suggesting that other minor constituents also contribute to the final objective. A similar situation was observed in other studies when testing the effect of Eucalyptus resinifera EO and its main active ingredients against the pests Rhyzopertha dominica Frabricius (Filomeno et al., 2020) and Hypothenemus hampei Ferrari (Reyes et al., 2019). It has been suggested a possible synergistic effect between eucalyptol and p-cymene in increasing the antibacterial activity of EOs, additive and synergistic effects between eucalyptol and aromadendrene against methicillin-resistant Staphylococcus aureus (Salem et al., 2018), and additive effects between eucalyptol and p-cymene against Rhyzopertha dominica (Filomeno et al., 2020). A wide range of binary mixtures of components present in EOs was tested to assess the effect (antagonism, synergism, or no effect) against larvae of Spodoptera littoralis (Pavela, 2014). EOs are used in the composition of commercial products, such as some herbicides available in the North American market, such as Matratec (50% clove oil), WeedZap (45% clove oil + 45% cinnamon oil), and GreenMatch EX (50% lemongrass oil) (Verdeguer et al., 2020b). They can also be combined to develop new products, as demonstrated by Golestani et al. (2015), who formulated two phytomedicines from different concentrations of four EOs (Eucalyptus globulus, Dianthus caryophyllus L., Mentha piperita L., and Thymus vulgaris L.) against Escherichia Effect Reference Potential antimicrobial against gram-positive bacterium Staphylococcus aureus and yeast Candida albicans Cytotoxicity against tumor cell lines Calu-3 and 3T3 (Saulle, 2018) Antibacterial against Staphylococcus aureus ATCC6538, Escherichia coli ATCC8739, Escherichia coli 0:158 and Salmonella choleraeseus ATCC10708 (Estanislau et al., 2001) Fumigant and repellent against pest insects Callosobruchus chinensis and Sitophilus zeamais (Bett et al., 2013) Insecticidal and repellent against stored grain pests (Tribolium castaneum, Acanthoscelides obtectus, Sitotroga cerealella and Sitophilus zeamais)(Bett et al., 2016) Eucalyptus urophylla S.T.Blake Larvicide and repellent against Culex quinquefasciatus, the filariasis vector mosquito (Pujiarti & Kasmudjo, 2016) Antibacterial against Bacillus subtilis, Escherichia coli, Klebsiella oxytoca, Klebsiella pneumoniae Antifungal against Aspergillus clavatus, Aspergillus niger, Chaetomium globosum, Cladosporium cladosporioides, Lenzites sulphureus, Myrothecium verrucaria, Penicillium citrinum, Phanerochaete chrysosporium, Phaeolus schweintizii, Trametes versicolor, Trichoderma viride Insecticidal against Atta sexdens rubropilosa, Thyrinteina arnobia Herbicidal against Lactuca sativa (Barbosa et al., 2016) Table 2. Cont. Template: Editora Letra1 | www.editoraletra1.com.br Rodrigues M, Mazzafera P 16 | Acta Botanica Brasilica, 2025, 39: e20240111 It is essential to emphasize that in addition to the comprehensive number of factors to be controlled, there are also numerous eucalypt species and possibilities for the application of EOs and their actives. The lack of study of some species because they are already considered for a particular purpose may prevent the discovery of effective products for other purposes. All these points generate an infinity of possibilities for studies, which should always document all plant characteristics, location, collection, extraction, and oil analysis whenever possible. Although many studies present benefits and the responsible and safe use of EOs, possible negative or undesired impacts can also be obtained; this type of response is found less frequently in the literature but should be equally documented when obtained. With more consolidated data related to raw materials and efficacy, another challenge is to make viable the use of these products through formulations that provide EO stability and are suitable for each application and objective, since one of their main characteristics is high volatility. Economic aspects should also be addressed to stimulate the entire previously mentioned process. The production and marketing of EOs have the potential for expansion and must be organized to ensure the quality and reliability of the products, as it is a market that supplies other countries with specific requirements and diverse uses. Even for internal use, oils need to meet certain standards to be used, and considering this economic context, in addition to specific constituents, another factor to be considered is the oil yield of the selected eucalyptus species. A quick way to assess interest in EOs is by surveying published works on the topic. Searching for the terms “essential oil” and “Eucalyptus” as topics in the Web of Science yielded 1945 works (publications between 1900 and 2023). When limiting this search to the last ten years, 1265 publications are found, showing that it is a topic that has been more studied in recent years. Solutions to problems faced today may be present in elements available in nature, such as EOs, which can be an alternative source for the development of natural medicines and more sustainable bio-inputs. Companies are increasingly interested in these products that can act through different mechanisms against microorganisms already resistant to traditional agents or generate new solutions. Thus, the more information and solutions obtained, the more viable this market will be, including financially. Acknowledgements The authors thank Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for doctorate and research fellowships, respectively. Authors’ Contributions Conceptualization: Rodrigues M., Mazzafera P. Writingoriginal draft: Rodrigues M. Writing review & editing: Rodrigues M, Mazzafera P. Conflict of Interest The authors declare no conflicts of interest (personal, scientific, commercial, political, or financial) in the submitted manuscript. References Abbass HS. 2020. Eucalyptus essential oil; an off-label use to protect the world from COVID-19 pandemic: Review-based hypotheses. Universal Journal of Pharmaceutical Research 5: 61-64. doi: 10.22270/ujpr. v5i4.440 Abdoul-Latif F M, Ainane A, Aboubaker IH, Mohamed J, Ainane T. 2023. Exploring the Potent Anticancer Activity of Essential Oils and Their Bioactive Compounds: Mechanisms and Prospects for Future Cancer Therapy. 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