Selection of beneficial fungi for plants with the potential to metabolize lunar and Martian regolith

Abstract

The establishment of sustainable agriculture on extraterrestrial bodies (the Moon, Mars, etc.) depends on transforming regolith, an unconsolidated, microbiota-free, mineral substrate, into a functional soil-like material with numerous beneficial properties including as a growth substrate for plants. Lunar and Martian regolith present physicochemical challenges, including alkaline pH, high concentrations of toxic elements (e.g., aluminum, manganese, perchlorates), and limited bioavailability of essential nutrients such as nitrogen, phosphorus, and potassium. This review explores the potential of beneficial fungi to support regolith-based agriculture (RBA) through biomineralization, nutrient mobilization, and bioremediation. We present fungal species capable of solubilizing phosphates, chelating metals via siderophores, and metabolizing iron and aluminum oxides, thereby enhancing nutrient accessibility. While some highlighted genera, such as Trichoderma, Penicillium, and Aspergillus, include well-known pathogenic species, they also encompass strains with potential applications for promoting plant growth under abiotic stress. Extremophilic fungi like Cryomyces antarcticus are also noted for their resilience. Furthermore, we discuss the potential role of arbuscular mycorrhizal fungi (AMF) from the phylum Glomeromycota in promoting plant growth under harsh conditions. On Earth, these fungi are known to enhance iron uptake, mitigate oxidative stress, and improve soil structure via glomalin-mediated aggregation, mechanisms that may be applicable to regolith systems. We also examine fungal strains isolated from the International Space Station which may possess pre-adapted traits suitable for these off-world environments. While knowledge gaps remain, particularly regarding biosafety, strain selection, and performance validation under simulated extraterrestrial conditions, including radiation exposure, we contend that fungi are essential biotechnological allies for ISRU (In Situ Resource Utilization), contributing to sustainable agriculture in both extraterrestrial habitats and degraded ecosystems on Earth. We emphasize the need for interdisciplinary research integrating astrobiology, microbial ecology, and space agriculture in preparation for future crewed missions to the Moon and Mars.

1 Introduction

Loss of flavor and nutritional content in pre-packaged foods as well as the need to ensure food security has led most space agencies to conclude that off-world settlements will require a plant growth habitat (‘greenhouse’) which leverages in-situ (on-site) resources (Johnson et al., 2021; Poulet et al., 2022). Such in situ resource utilization (ISRU) efforts include the use of the Martian or Lunar surface material (i.e., regolith) as a substrate for plant growth (Eichler et al., 2021). Regolith is a broad classification for unconsolidated materials on rock that can include dust, broken rocks, and other related materials (Fackrell et al., 2021; Duri et al., 2022) and is generally used in reference to surface layers. Regolith from these sites is particularly distinguished from even poor terrestrial soil by the absence of biological processes at off-world locations (Duri et al., 2022). However, lunar regolith is available only in limited amounts and no supply of Martian regolith is available from which to conduct plant studies. As a result, most space agriculture research relies on the production of regolith simulants based on existing literature on the composition of these sites (Fackrell et al., 2024).

Regolith has been widely regarded as a substrate for plant growth as it is a physical substrate readily available on site as well as a potential source of nutrients for plants, especially iron, magnesium, potassium and phosphorus (Fackrell et al., 2024). For example, calcium, aluminum, magnesium, iron, potassium, phosphorus, and titanium are present in lunar regolith and could potentially help plant growth, as these elements play roles in nutrient delivery, growth, and photosynthesis (Taiz et al., 2017; Oze et al., 2021) but may lead to plant toxicity when in excess (Chrysargyris et al., 2022). Micronutrients, so-called because they are only required in small amounts, may quickly become toxic when found in excess. Fackrell et al. (2024) compared the composition of lunar (sea and highlands) as well as Martian regolith to those required by plants, showing that some micronutrients are in quantities much higher than those required by plants. For example, about 0.01 w/v % of iron is needed to nourish plants, but there are 6, 15 and 12.4 w/v %, respectively, in highland lunar regolith, lunar sea and Martian regolith, that is 600, 1,500 and 1,240 times more than what is generally tolerated by most crops of interest.

However, in many cases, the mineral forms in which these macro and micronutrients are contained are not readily available to plants and may require additional effort, such as the use of microorganisms, to bioweather/transform them and make them more bioavailable (Fackrell et al., 2024). Yet the regolith at these sites will have no associated microbiota and only a limited amount of essential macronutrients such as nitrogen and potassium, and therefore, these substrates cannot adequately support plant growth in isolation. In addition, both Lunar and Martian regolith have characteristics that are harmful to plant health, such as alkaline pH, high sodium availability, low cohesion of mineral components, predominance of macropores over micropores, and low water retention capacity, as well as containing toxic perchlorates (Misra et al., 2021; Fackrell et al., 2024). On the other hand, despite these challenges, studies have shown success in some plants in the regolith system (Ming and Henninger, 1989; Berni et al., 2023). In this review we will consider the challenges associated with regolith-based agriculture systems, the composition of these materials, and how fungi may provide new tools for improving plant growth and/or modifying regolith in these challenging environments. While regolith simulants for other sources are available, we will concern ourselves with efforts related to the two major off-world targets for settlement of Earth’s moon and Mars.

1.1 The potential of regolith-based agriculture (RBA)

Wamelink et al. (2014) demonstrated the successful cultivation in a lunar regolith simulant (JSC-MARS-1A) of plants such as: tomato (Lycopersicon esculentum), rye (Secale cereale), carrot (Daucus carota subsp. sativus) and four nitrogen-fixing species. Rye, watercress (Nasturtium officinale) and field mustard (Sinapis arvensis) even flowered, and the latter two also formed seeds, indicating that it is possible to obtain productivity in RBA. Meanwhile, Paul et al. (2022) showed that Arabidopsis thaliana could be grown directly in real lunar regolith with the support of exogenous nutrient solutions. However, their growth was slower, and the plants showed a severe stress phenotype, which was subsequently confirmed by transcriptomic data. Collectively, these results indicate that it is necessary to improve the physical and chemical properties of lunar regolith including increasing the availability of essential nutrients to plants before establishing an extraterrestrial higher plant cultivation system that could effectively provide life support and edible crop production on a lunar station.

Cicer arietinum (chickpea), Lactuca sativa (lettuce), Pisum sativum (pea) and Capsicum annuum (pepper) seeds grew in lunar and Martian regolith simulants following seed priming with the green algae Ulva lactuca L. (Oliveira et al., 2024). These cultivars had a good percentage of germination and seedling establishment when the algae solution was applied in small amounts (0.2 g⋅L⁻¹). For example, chickpeas (C. arietinum) cultivated in Martian regolith simulant had a 97.5% ± 5% germination in relation to control (70% ± 8.2%) and a similar percentage of seedling emergence as well. Pea (P. sativum) germination was higher when applied 0.2 g⋅L⁻¹ (97.5% ± 5%) of the extract was applied and seedling emergence was equally significant. In the case of lettuce (Lactuca sativa) grown in lunar regolith simulant germination reached 92.5% ± 5% and seedling emergence was up to 100%, both higher than the untreated control (85%). These results suggest that algal secondary metabolites which are precursors of plant phytohormones could help mitigate stress (Oliveira et al., 2024). This technique consists of exposing the plant to a little stress, preparing it to deal with more intense stresses in the future. In this preparation, phytohormones act as triggers, reinforcing the plant’s defense (El-Aziz Kasim et al., 2016; García et al., 2020; Reis et al., 2020). Additionally, both algae and cyanobacteria are relevant sources of organic matter, through the release of organic carbon during photosynthesis highlighting their central role in bioregenerative systems.

Sasi et al. (2025) evaluated whether Brassica oleracea and Medicago sativa show improved performance in the Martian regolith simulant MGS-1 when treated with plasma-activated water (PAW) or mineral-rich Aquapulse® water (APW). In B. oleracea, PAW seed soaking followed by APW irrigation sharply increased germination (≈566.7%), biomass (≈412.4%), and chlorophyll content (≈17.7%). In M. sativa, APW used for both soaking and irrigation raised germination by 41.7% and biomass by 45.2%, while PAW alone produced the strongest improvement in germination (41.7%) compared with the control.

Supplementation with spirulina, Arthrospira platensis, blue-green cyanobacteria recently reclassified as Limnospira platensis, also shows potential to enhance the growth of radish microgreens (Raphanus sativus) in lunar and Martian regolith simulants (MGS-1S and LMS-1) under both elevated CO₂ and ambient conditions. The Martian regolith simulant with 0.6% spirulina under elevated CO₂ produced the best results. The type of regolith simulant influenced biomass more strongly than plant height, with overall better growth observed in the Martian simulant compared with the lunar one. The application of spirulina aqueous solution promoted healthier plant development in both simulants across CO₂ levels, improving traits such as height, volume, leaf number, and coloration throughout the growth cycle, with seed formation occurring only in the high-CO₂ tents (Miller et al., 2025). All these studies demonstrate the potential of regolith-based agriculture (RBA), but they consistently require some techniques involving microorganisms that promote plant growth. These studies highlight a number of alternate strategies that could be leveraged for conditioning regolith simulants.

1.2 Microbiota as a solution for regolith conditioning

Making lunar and Martian soil suitable for cultivation is a complex task, but the addition of organic matter combined with the activity of plant growth-promoting microorganisms represents a promising strategy. However, transporting terrestrial substrates to space remains economically prohibitive: under NASA’s interim commercial marketing pricing policy first released in June 2019, up mass costs for delivering cargo to the International Space Station were estimated at approximately US $20,000 per kilogram, illustrating the high logistical burden of sending materials from Earth. Although NASA has since updated its pricing policy to reflect full reimbursement for the value of station resources as part of broader commercial activities, this shift underscores that up mass continues to be a major cost driver for low-Earth orbit operations and beyond. Consequently, research on regolith-based systems has focused on optimizing in situ resource utilization (ISRU) to adapt the lunar and Martian environment (Eichler et al., 2021), yet relatively few studies have investigated the interaction between microbial communities and regolith systems (Fackrell et al., 2024).

Zaets et al. (2011) inoculated marigolds (Tagetes patula L.) in a lunar regolith simulant (terrestrial anorthosite) with a consortium of bacteria that which led to a reduction of Ni, Cr, Zn, and Na levels in the regolith and altered the Ca/Mg and Fe/Mn ratios in the biomass of marigold cultivated in anorthosite. More recently, Misra et al. (2021) discussed known microbial remediation processes using rhizosphere microorganisms and extremophiles, which are the most likely candidates for successful microbial remediation of Martian regolith, considering the perchlorate-rich environment. For example, four strains of extremophilic bacteria (Peribacillus frigoritolerans, Bacillus atrophaeus, Cellulomonas sp., and Pseudarthrobacter sp.) were isolated from desert soils of the Qaidam Basin in the northern Tibetan Plateau (China), where perchlorates (ClO₄⁻) are present in both surface and subsurface soils, highlighting the ability of these microorganisms to tolerate this compound (Li et al., 2024).

Even less attention has been paid to fungus-regolith system interactions (Duri et al., 2022). In laboratory experiments without the use of regolith, Heinz et al. (2020) reported the perchlorate tolerance of the halotolerant yeast Debaryomyces hansenii was 2.4 M NaClO₄ (23 wt.%). Perchlorates which have been found in the Martian soil at the Phoenix landing site at concentrations of 0.4–0.6 wt.% (Hecht et al., 2009). According to the authors, the perchlorate tolerance data available in the literature to date indicates that some fungal species may be more efficient at metabolizing or managing perchlorates than many bacteria and archaea.

Proteobacteria such as Sphingomonas and fungi such as Penicillium spp. can be used or genetically modified to remove perchlorates (converting them into molecular chloride and oxygen) from regolith, as well as other hazardous substances such as heavy metals, radioactive species, acids, and salts (Misra et al., 2021; Koehle et al., 2023). These microorganisms can catalyze very specific reactions, resulting in efficient resource use (Koehle et al., 2023). Given that both lunar and Martian regolith are devoid of organic matter, recent evidence demonstrating the central role of saprotrophic fungi in organic matter recycling and nutrient cycling within closed biological systems (Qiao et al., 2025) reinforces the relevance of integrating these fungi into bioregenerative life support systems as a means to sustain ecosystem functionality in regolith-based habitats. Therefore, for this review we consider the composition of lunar regolith and Martian regolith and the potential roles a variety of fungal species could play in missions beyond Earth.

2 Characteristics of regolith

2.1 Chemical composition

It is important to understand the chemical properties of regolith, such as their mineral composition to improve their adaptation for plant cultivation (Ding et al., 2024). In the absence of abundant regolith samples returned from space, we are dependent on terrestrially derived proxies (simulants) for the majority of the biological, chemical, and mechanical research for these off-world sites at present. While a number of Martian and lunar regolith simulants have been produced by a variety of organizations/researchers it is those produced by Space Resource Technologies (formerly Exolith) which are arguably the most prevalent in research efforts at present. Of the eleven simulants developed by Space Resource Technologies (Long-Fox and Britt et al., 2023), we present here the data on relative abundances of oxides in the lunar simulants LHS-1 (Highlands Simulant) and Martian MGS-1 (High-Fidelity Martian Regolith Simulant) (Table 1). It is worth noting that these simulants are not the most accurate, however we consider them to be the most relevant for plant growth (Fackrell et al., 2024).

The pH of Martian regolith is often reported to be above 6, and in some cases, it has alkaline properties (Zaets et al., 2011; Wamelink et al., 2014; Caporale et al., 2020; Eichler et al., 2021). Alkaline pH reduces the solubility of essential nutrients like phosphorus and potassium. This suggests low rates of mineral weathering and cation release (Duri et al., 2022). The pH of lunar regolith is also alkaline, but it can vary depending on its location on the Moon and the specific mineral composition of the sample (Papike et al., 1982; Noble, 2009).

Lunar and Martian regolith differ in physico-chemical properties that could support different fungal adaptations. Lunar regolith is extremely dry, highly fractured, and dominated by fine, glassy and angular particles formed by impact processes, with very low volatile and organic content, whereas Martian regolith contains hydrated minerals and water-related phases and is characterized by the presence of toxic salts such as perchlorates and a broader range of iron oxides and weathered minerals (Duri et al., 2022). These contrasts suggest that fungi interacting with lunar substrates would need to cope with extreme desiccation and abrasive mineral surfaces, while those in Martian regolith would additionally face ionic stress and potential interactions with hydrated phases.

2.2 Components of lunar and Martian regolith and the impact on plant health

The oxides present in regolith impact plant health in several physiological processes, involving not only the absorption of nutrients, but also causing osmotic imbalance, cell damage or negatively affecting the photosynthesis process, when in toxic concentrations. It is important to note that the toxicity of each component is related not only to its concentration itself, but to the interaction with different plant species or genotypes, and environmental factors. Some examples of impacts of each oxide component of the regolith and their respective affected plant species are listed in Table 2.

The limited concentrations of macronutrients on the Moon or Mars such as nitrogen, phosphorus, and potassium are of particular concern. According to Fackrell et al. (2024), nitrogen has been detected on the Moon and Mars, but only in trace amounts. On the Moon, potassium and phosphorus are limited, and plant growth would require supplementation. On Mars, potassium has been detected, but no large reserves capable of sustaining in situ agricultural supply have been identified, while phosphorus is relatively abundant and is present, at least partially, in forms extractable by plants.

3 Fungi with regolith metabolization potential

Fungi possess the ability to produce a substantial amount of secondary metabolites and are distinguished by their broad metabolic capacities (Muhammad et al., 2024). The enzymes present in their extracellular contents play a crucial role in the degradation of pollutants, which can then be digested and absorbed by the cell wall (Shelke et al., 2023). These fungi and their exoenzymes in conjunction with other microorganisms present in the soil, enrich the diversity and overall functionality of soil ecosystems. Under conditions of environmental stress in a space environment, fungi can be essential candidates to support plant growth, especially in challenging substrates such as lunar and Martian regolith type (Vezzola et al., 2023). Fungi-based products can be applied to plants to promote improved nutrient utilization and uptake, stress tolerance, and increased crop yields (du Jardin, 2015). Thus, fungi-derived plant growth biostimulants could prove to be useful alternatives considering the costs and risks associated with transporting fertilizers to a lunar or Martian colony (Cannon and Britt, 2019).

3.1 Species with potential application as biostimulants in RBA

Species of Trichoderma spp. (Ascomycota) are known for their biostimulant properties and promoting plant health, as well as being antagonists to soil pathogens (Konappa et al., 2022). They can improve soil structure, break down organic matter, and release essential nutrients to plants (Konappa et al., 2022). Fungus spores of Trichoderma longibrachiatum were exposed to low-Earth orbit for almost 2 years aboard the EXPOSE-R facility outside the International Space Station ('Spores in artificial meteorites' SPORES). About 30% of the spores in the vacuum survived space travel when protected from radiation. These data opened up discussions about the likelihood of lithopanspermia, the potential natural transfer of microorganisms between planets (Neuberger et al., 2015). Recently, a space tomato inoculation experiment associated with Trichoderma: TASTIE, for research on tomato growth on the ISS grown on the VEGGIE equipment. During the experiment, a group of samples is being grown in petri dishes along with a potentially beneficial fungus of the genus Trichoderma (Barker et al., 2020).

Konings-Dudin et al. (2014) evaluated how endophytic fungi affect seed germination in the lunar regolith simulant JSC-1. The fungus Trichoderma viride, present in the roots of seedlings of Opuntia ficus-indica, significantly improved seed germination rates compared to other fungi isolated from the rhizosphere of the Opuntia. This work also suggested that a symbiotic association between T. viride and a species of the mycorrhizal fungus of Glomus, may be crucial for the germination and establishment of plants in nutritionally poor soils, such as the lunar regolith. Based on this evidence, we propose that these species have the potential to improve regolith structure and viability as an agricultural substrate in extraterrestrial environments.

3.2 Mechanisms of fungal bioremediation

Fungi are heterotrophic organisms and commonly used in biomining, a process that uses living organisms to extract heavy metals such as Co, Cu, Mn, and Fe from ores and waste (Li et al., 2024). These microorganisms can be useful in regolith-based agriculture as they accelerate the dissolution of metal-containing minerals (Vezzola et al., 2023). The process uses fungi-derived organic acids to extract metals from rocks with oxides, elements in abundance in both lunar regolith and Martian regolith (Vezzola et al., 2023). Iron present in Martian regolith is found in largely insoluble states, inaccessible to plants, such as oxides (Duri et al., 2022). Fungal organisms have the biochemical and biological capacity to degrade ecologically natural synthetic compounds and to diminish the dangers related to metals, metalloids, and radionuclides, either by blend modification or by affecting substance bioavailability (Govindan et al., 2021). These fungal characteristics can help plants to be able to adapt to environments of metal rich (Table 3).

A study analyzing biomining by the fungus Penicillium simplicissimum in the lunar regolith simulator EAC-1A highlighted the potential of the fungus to extract metals such as aluminum, iron, magnesium and calcium under reduced gravity conditions. The bioleaching process was conducted over 2 weeks, resulting in an average recovery of 10 ± 3 g/L of metal dust from the leachate (Figueira et al., 2025). This highlights a promising ISRU approach in future lunar missions.

The biocorrosion of aluminum and aluminum-based alloys by 13 micromycete species belonging to the genera Alternaria, Aspergillus, Chaetomium, Fusarium, Paecilomyces, Penicillium, and Trichoderma was evaluated and the species Alternaria alternata, Aspergillus terreus, Chaetomium globosum, Fusarium moniliforme, Paecilomyces variotii, Penicillium funiculosum, P. martensii, P. ochro-chloron evidenced showed surface with obvious corrosion damage after 60 days. Alternaria alternata was identified as the most active biodegrader (Smirnov et al., 2008). Given this metabolic versatility, we hypothesize that the application of some species, such as A. alternata, in lunar and Martian regolith may favor aluminum extraction and could contribute to the development of innovative technologies.

Boriová et al. (2019) demonstrated that A. niger was also able to remove aluminum from contaminated aqueous solutions. In the case of lunar regolith where aluminum is in high concentrations (26.6%) and in Martian regolith (12.8%), we believe that Aspergillus may be a remedial when implemented in crops associated with plants in this type of environment, rich in aluminum, contributing to the safety of the cultivation environment. Oxalate-aluminum complexes contribute to microbial regulation and nutrient dynamics, mitigating aluminum toxicity through the formation of stable complexes that reduce aluminum bioavailability without the need for its removal from the substrate, as demonstrated in the fungus Tricholoma matsutake (Nishino et al., 2017). Aspergillus tubingensis was able to extract metals from an alternate Martian regolith simulant, Mars Mojave Simulant (or MMS-1) via biomining (Vezzola et al., 2023). The results indicated that the fungal strain produced organic acids, especially oxalic acid, in the first 5 days, leading to a rapid reduction in the pH of the culture medium. Moreover, understanding and exploring the diversity of metal-metabolizing fungi is essential for regolith-based applications, as this knowledge can inform genetic engineering strategies to enhance metal tolerance, detoxification, or mineral-transforming traits in well-characterized fungal species suited for extraterrestrial substrates.

4 Phosphate solubilization through fungi and biotransformation

Fungi can contribute to phosphate mobilization in nutrient-poor regolith through organic acid production, metal chelation, phosphatase activity, and biologically mediated mineral weathering, thereby supporting early soil-forming processes. Fungi such as Penicillium and Aspergillus, are also known for their ability to solubilize phosphates, making phosphorus available to plants. Usually, the solubilization process is associated with a decrease in pH, which is beneficial in making phosphorus more available to plants (Barate and Gawai, 2024). Although specific research on their application in lunar or Martian regolith is limited, their function in terrestrial soils suggests that they may play a similar role in improving regolith. The ability of these fungi to solubilize phosphates can be crucial in environments where phosphorus is present in forms that are inaccessible to plants, or in small quantities, as in the case of both regolith, as they help in the mobilization and sequestration of metals. For example, Islas-Valdez et al. (2024) indicated four of 25 fungal isolates as being the most efficient Phosphorus Solubilizing Microorganisms (PSMs) and Zinc Solubilizing Microorganisms (ZnSMs): Aspergillus awamori, Fusarium circinatum, Fusarium longifundum and Mucor circinelloides. The authors evaluated a variety of environments including different land uses, crop rotation systems, and soil amendments, while quantitatively and qualitatively tracking PSMs and ZnSMs capable of solubilizing different sources of poorly soluble P (i.e., tricalcium phosphate, rock phosphate, and calcium phytate).

5 Siderophore-producing fungi

Many fungi naturally produce high-affinity siderophores to acquire iron under low-iron conditions in diverse terrestrial soil environments such as forest soils, grassland soils, and rhizosphere environments, demonstrating their ecological adaptation to nutrient-limited substrates; this adaptive capacity suggests that similar siderophore-mediated iron mobilization could be relevant when such organisms are introduced into mineral-rich, nutrient-limited regolith. Fungi utilize at least two of the four iron uptake mechanisms: (i) direct Fe²⁺ permeation, (ii) coupled ferroxidase/permease uptake, (iii) heme/hemin uptake, and (iv) siderophore-mediated iron uptake. Direct permeation of Fe²⁺ is well characterized in the yeast Saccharomyces cerevisiae and some opportunistic fungi express receptor genes for the heme/hemin pathway. Among the fungi that produce siderophores, secondary metabolites with a high affinity for ferric iron (Choi et al., 2024), is Beauveria bassiana, whose siderophore molecules chelate iron, converting it into a soluble and more bioavailable state, in the form of ferritin (David and Easterbrook, 1971) for plants (Benite et al., 2002; Sun et al., 2024). The metal sequestration properties of siderophore-producing fungi can play a crucial role in processes such as iron biotransformation (Tafazzoli et al., 2024), in addition to favoring the acclimatization of plants in Martian regolith, characterized by a high iron content (11.2 wt%). The mutualistic interactions between siderophore-producing fungi and plants is a viable strategy to increase plant growth in environments rich in iron oxides and with a high concentration of aluminum. This is primarily due to their ability to increase the production of siderophores to chelate and detoxify these metals when present in high proportions (Desai and Archana, 2011). Studies by Illmer and Buttinger. (2006) based on the microbial biomass and production of siderophores by three fungi (Aspergillus nidulans, Neurospora crassa and Hymenoscyphus ericae demonstrated that the quantity but not quality of fungal siderophores are distinctly influenced by Al-availability and that there are strong interactions between Al-toxicity and concentrations of Fe and iron-chelating substances (ICS). Whereas both Fe and Al inhibit ICS-synthesis, the toxic effects of Al on microbial biomass are reduced by Fe and enhanced by ICS. According to the authors, the production of siderophores varied significantly between species, even under similar conditions. These results suggest that aluminum affects the synthesis of siderophores through mechanisms that can be both defensive and adaptive, modulated by environmental conditions and the specific physiological characteristics of each fungus. Such variability reinforces the need for further studies to evaluate the applicability of these responses in extreme environments, such as the lunar and Martian regolith.

There is significant interest in exploring how fungi can be employed to extract metals and remediate soils on celestial bodies such as the Moon and Mars, mainly due to their bioremedial potential in degraded areas (Soares et al., 2011).

6 Extremophile fungi and their applications in bioremediation

Fungal extremophiles such as Cryomyces antarcticus could be used in future space missions for bioremediation, since their resilience to extreme environmental stressors, including radiation and their ability to survive exposure to high doses of ferric cations, has been previously documented (Aureli et al., 2020; Simões et al., 2023). This resilience may contribute to its role in bioremediation and other applications in space exploration, although the science surrounding space biomining and bioremediation is still in its early stages (Simões et al., 2023; Shevtsov, 2021).

Cryomyces antarcticus produces melanin, which is crucial for its survival in extreme conditions such as high UV radiation and desiccation (Pacelli et al., 2020; Catanzaro et al., 2024). Fungi, particularly those with a melanized state, show promise in breaking down many different compounds, such as persistent organic pollutants, due to their ability to produce enzymes that break down these contaminants outside of their cells (Soares et al., 2011). A recent study investigated the mechanisms of action of radiotrophic fungi, present in extreme environments such as Chernobyl, Antarctic as well as the International Space Station. The main objective was to understand the absorption of ionizing radiation and the deposition of radioisotopes in the cell walls of these organisms. Additionally, their effectiveness as bioremediation agents was evaluated. The research suggests that these fungi have potential for both bioremediation and protective applications and can reduce human and equipment exposure to ionizing radiation, in addition to acting as biosensors for its detection (Tibolla and Fischer, 2025).

7 A survey of fungi reported from the international space station belonging to plant growth–promoting genera

Microorganisms found on the ISS have been able to survive the stressful conditions of the space environment. This suggest they could be used to evaluated in controlled regolith-based experiments to determine their potential effects on substrate properties and nutrient availability (Simões et al., 2023). It is believed that once established in the plant units of the ISS, fungi will be able to persist in the plant microbiome outside of their natural environment (Hummerick et al., 2021; Koehl et al., 2023). We consider that species PGPB of the same genus found in the ISS may share similar adaptations to the spaceflight environment.

Microbial resilience is addressed as a prerequisite for future experimental investigation; however, it is important to emphasize the current lack of experimental evidence integrating fungi, regolith-based substrates, and plants under space-relevant conditions. By outlining this knowledge gap, this review seeks to highlight priorities for targeted experimental studies, rather than to suggest any inherent advantage of fungi reported from the International Space Station (ISS) for regolith-based agricultural applications.

The benefits of plant growth-promoting microorganisms (PGPMs) are not limited to improving growth and development but also enhancing tolerance/resistance to abiotic and biotic stress to the plants with which they are associated (Raja Gopalan et al., 2023). The chemicals secreted by the microorganisms modulate the expression of key stress-responsive genes and their corresponding proteins; important metabolic pathways that regulate phytohormone signaling in plants.

Table 4 presents the previously reported beneficial potential under terrestrial conditions of genera or species of fungi isolated on the ISS or identified at the station by metagenomics. Since phylogenetically related species frequently share functional traits, fungal species from genera detected on the ISS may likewise have the capacity to survive and support plant growth under space conditions.

According to Contreras-Cornejo et al. (2024), fungi of the genus Trichoderma can cause significant biochemical changes in the plant content of carbohydrates, amino acids, organic acids, and lipids, as detected in Arabidopsis thaliana, corn (Zea mays), tomato (Solanum lycoperscicum), and barley (Hordeum vulgare), which can improve plant health throughout its life cycle.

The species Penicillium chrysogenum indicated in Table 4 is of special interest for space agriculture. Isolated from Antarctic plants and, therefore, characterized as a psychrophilic fungus, with diverse enzymatic activities at 4 °C, including nitrogen mineralization, suggesting that root endophytes play a key ecological role in accelerating the nitrogen cycle, improving nutrient acquisition, and thus promoting plant growth in Antarctic terrestrial ecosystems (Oses-Pedraza et al., 2020). It is also important to highlight that this species has optimal growth at 27 °C and can tolerate an increased temperature of 31 °C, and can be useful in other environments (Ramšak and Kück, 2022).

The most studied fungal genera for promoting plant growth are Trichoderma, Penicililum, and Aspergillus, all found on the ISS (Argumedo-Delira et al., 2022; Simões et al., 2023). However, caution is needed for the rigorous selection of appropriate fungal strains to promote plant growth because some strains of these genera are pathogenic to plants, animals and humans (Pitt, 1994). Aspergillus fumigatus for example, can cause allergic reactions to severe invasive infections, particularly in immunocompromised patients (Morrissey et al., 2024). Pfordt et al. (2025) analyzed some species of Trichoderma and found that T. asperellum, T. atroviride, and T. guizhouense exhibit pathogenicity in corn cobs with disease severity reaching 92%. During apple inoculation experiments, Penicillium expansum strains produced patulina, a mycotoxin, which is a toxic secondary metabolite produced by certain fungi, with concentrations ranging from 2.31 to 88.67 mg/kg in the tissues of the rotten fruit, levels considerably lower than those observed in PDA medium according to Shen et al. (2024). Therefore, it is worth emphasizing the importance of considering biosafety aspects related to its application in controlled environments (Fackrell et al., 2024).

Moreover, because fungal endophytes and other seed-associated fungi are frequently transmitted with plant propagules and are difficult to fully eliminate even under sterile conditions (Salamon et al., 2025), their unintended introduction into regolith-based extraterrestrial production systems should be expected, making fungal–fungal and fungal–plant interactions an important consideration for system stability and performance.

We believe that the beneficial fungi and their different mechanisms of plant growth promotion already elucidated on Earth have the potential to act in a similar way in lunar and Martian regolith and may contribute to space agricultural practices in the future. Such research represents a vast field of investigation to be explored by the scientific community and that can be applied in terrestrial conditions or analogous missions with the use of regolith simulators.

8 Arbuscular mycorrhizal fungi (AMF) as candidates to metabolize regolith and promote plant development

The fungi found on the ISS belong to the phyla Ascomycota and Basidiomycota, with no genera from the phylum Glomeromycota, that comprise the obligate biotrophic arbuscular mycorrhizal fungi (AMF) which associate with most terrestrial plants.

AMF species isolated in terrestrial soils rich in heavy metals, especially iron and aluminum, as well as those that occur in poor edaphic environments, actually thrive in conditions comparable to regolith. We propose that the strategic inclusion of AMFs in an integrated approach that considers the plant-associated microbiota can generate significant advances for space missions and the development of extraterrestrial agricultural ecosystems. In addition, although focused on space exploration, RBA can also provide valuable insights for agriculture in impoverished soils on Earth, contributing to sustainable and resilient practices in the terrestrial biosphere.

Arbuscular mycorrhizal fungi are well known to enhance plant resistance to a wide range of abiotic stresses, including drought, salinity, nutrient limitation, and heavy metal toxicity, as well as to biotic stresses such as soil-borne pathogens (Joner et al., 2000). AMF are therefore interesting candidates for metabolizing regolith. Despite their well-recognized positive influence on plant development, few studies have involved the use of AMF in regolith or other analogous substrates.

Kozyrovska et al. (2004) evaluated the influence of a microbial consortium (bacteria and the mycorrhizal fungus Glomus) in the development and flowering of Tagetes patula L. under limiting factors in a substrate similar to lunar rocks, showing that the intercropping favored the germination of the plants, which flowered at 54 days of co-cultivation, while the control plants, not inoculated, died after 3–4 weeks. Recently, Atkin and Oliveira Pedro dos Santos (2024) created a fertile matrix in lunar regolith simulant using AMF and vermicompost (in different mixtures with regolith) in chickpeas (Cicer arietinum). The species of AMF used by these authors were: Rhizophagus intraradices, Funneliformis mosseae, Claroideoglomus claroideum and Claroideoglomus etunicatum. Chickpea plants inoculated with AMF flowered, produced seeds, and showed elevated chlorophyll concentrations, which suggests that the fungi helped mitigate the stress caused by the simulant.

Moreover, the co-occurrence of diazotrophs (nitrogen-fixing bacteria) and AMF can enhance biological nitrogen fixation and plant nutrient uptake, indicating a beneficial consortial relationship. For example, Mirsha et al. (2016) showed synergistic effects of AMF and plant growth promoting rhizobacteria in bioremediation of iron contaminated soils. Such synergistic microbial relationships, well documented in terrestrial soils where phosphorus and nitrogen are limiting, also suggest a promising strategy for nutrient mobilization in nutrient-poor regolith environments, where biological consortia may jointly improve access to nitrogen and other essential elements.

The ability to incorporate AMF into RBA food production systems helps bolster the case for this crop system’s inclusion in mission planning. However, ensuring food safety/security depends on redundant crop production efforts, supporting the inclusion of both hydroponic and RBA systems in any space agriculture plans to support an off-world settlement. With that in mind, the ongoing development and optimizations for the hydroponic cultivation with AMF, especially in small-sized systems such as Das et al. (2020), are potentially beneficial to these systems as well.

8.1 AMF and iron uptake by plants

Iron plays fundamental biological roles in processes such as photosynthesis and respiration and is frequently one of the most limiting essential micronutrients for plant growth, even in natural terrestrial habitats, because its low solubility and strong association with insoluble Fe³⁺ compounds that restrict its bioavailability (Vélez-Bermúdez and Schmidt, 2023).

Plants have evolved two major strategies for the absorption of iron. Model plants like A. thaliana, as well as many agricultural crops such as tomatoes, rely on strategy I, in which Fe³⁺ solubility is increased by acidification of the rhizosphere by H⁺ translocation or organic compounds, followed by reduction to Fe²⁺ by ferric chelate reductase at the root surface. Alternatively, grasses such as barley, rice and corn developed strategy II, generating and secreting phytosiderophores, organic molecules with affinity for Fe. Fe³⁺ is chelated by these compounds then transported across the membrane (Taiz et al., 2017). Both in the rhizosphere and in the simulated regolith, iron is present mostly as iron oxides and, therefore, has low solubility.

In long-distance transportation within the plant, translocation, the driving force is mainly transpiration. In organs that transpire little or at night (closed stomata) or when there is high humidity, the driving force is the positive pressure of the root due to the concentration of ions in the rhizosphere (guttation). In this case of Fe, it is transported as Fe³⁺ citrate in the xylem (upward movement, from root to shoot) and as complexed Fe²⁺ to nicotianamine in the phloem. In plant metabolism, Fe is important in the biosynthesis of chlorophyll, it is part of classes of proteins (cytochromes) and enzymatic constituents (cofactors) that transport electrons, therefore important for photosynthesis and respiration. Redistribution (or re-translocation), which involves transporting nutrients from old leaves (source) to growing organs or storage, via phloem, is inefficient because iron is poorly mobile, precipitated as insoluble oxides, and therefore its deficiency appears in young leaves (chlorosis between the veins) (Taiz et al., 2017).

A recent review by Liu et al. (2023) presents a number of studies showing that AMF-associations with plants can increase iron uptake. For example, AMF promoted an increase in iron concentrations in the roots and stems of sunflowers (Helianthus annuus), a model Strategy 1 plant, under Fe deficiency relative to controls (Kabir et al., 2020). Similar AMF treatments stimulated improvements in the absorption of iron in T. patula nana (Haselwandter et al., 2020) and Medicago sativa (Rahman et al., 2020), both Strategy 1 plants, relative to controls in iron-limiting conditions. According to Liu et al. (2023) there is a possibility that iron is also mobilized by the organic acids released by AMF under a low Pi supply and are easier to absorb by host plants.

In terms of Strategy 2 plants, transcriptomic analysis of AMF-colonized Triticum aestivum (wheat) roots showed upregulation in the gene expression of two phytosiderophore transporters, Traes_6AL_E36FCEF64 (homologous to a barley iron-phytosiderophore transporter) and Traes_6BL_D65EC1432 (Yellow Stripe-Like 15, YSL15 homolog) (Li et al., 2018). AMF-treated Maize upregulated the expression of genes associated with both the maintenance of iron content as well as the expression of Fe-related transporter genes (OPT8a and OPT8b) (Chorianopoulou et al., 2015; Kobae et al., 2014). Similarly, AMF-treatments stimulated Sorghum bicolor to increase the release of phytosiderophores as well as the expression of genes related to iron absorption (Prity et al., 2020). Oryza sativa (rice) is particularly interesting as it displays elements of both Strategy I and II in managing iron uptake and here also AMF-associations have shown improvement through a ferric-nicotianamine transporter (Kobae et al., 2014).

In addition to stimulating the production of root exudates that can chelate and solubilize iron (Sardans et al., 2023), under iron deficiency conditions, plants in symbiosis with AMF exhibit improved stress tolerance by activating stress-associated signaling pathways (Rahman et al., 2020). AMF associations can also regulate micronutrient concentrations in plant tissues, offering a mechanism to mitigate the risks of deficiency or toxicity (Mahmud and Chong et al., 2022). This could be of particular interest in regolith-based agriculture systems. Indeed, studies by Gill and Tuteja (2010) and Emamverdian et al. (2015), suggest that mycorrhizae-induced antioxidants (both enzymatic and secondary metabolites) in host plants can help eliminate excess reactive oxygen species (ROS) produced by heavy metal exposures.

In addition to enhancing plant mechanisms for iron uptake and maintenance, many mycorrhizal fungi produce their own siderophores or modifications to endogenous iron uptake pathways to perform these tasks which can then be conferred to plants. For example, the ferric reductase (RiFRE1) and the high affinity Fe permeases (RiFTR1-2) were found in the extraradical mycelia (ERM) of Rhizophagus irregularis associated to maize plants grown without Fe (Tamayo et al., 2018). Later, it was shown that the coupled to the expression of an extracellular multicopper oxidase (MCO) can facilitate the reductive iron update mechanism by these fungi. Tamayo et al. (2025) suggested that RiMCO1 and RiMCO3 might be the partners of the Fe permease RiFTR1. Meanwhile, mycorrhizal fungi such as ericoids, orchidaceans and ectomycorrhizals use hydroxamate-type siderophores. AMF also have siderophores typical of the phylum Zygomycota, such as rhizoferrin which can be translocated into plant tissues where their presence is indicative of active AMF smybioses (Brundrett and Tedersoo et al., 2018; Rajapitamahuni et al., 2023). However, there is still speculation about the involvement of bacteria in the synthesis of siderophores by AMF. For example, bacterial strains capable of producing siderophores have been observed in the spores of R. intraradices (Battini et al., 2016). Similarly, the siderophore producing obligate endobacteria Candidates Glomeribacter gigasporarum (Burkholderiaceae) has been observed as a symbiont in the hyphae and spores of the AMF Gigaspora margarita (Desiro et al., 2014). This underscores the potential need to incorporate both bacteria and fungi into any engineered microbiome for space agriculture.

8.2 AMF in metal-contaminated soils

Arbuscular mycorrhizal fungi propagate by producing mycelium (hyphae) or spores. These indigenous young propagules always persist in the soil and are resistant to heavy metals, increasing the heavy metal tolerance of host plants after colonization. The associations of these fungi with plant roots in metal-contaminated soils have been widely recognized (Karimi et al., 2011). Mycorrhizae have a glycoprotein, called glomalin, that promotes soil aggregation and sequesters heavy metals, reducing their bioavailability to plants (Rillig, 2004). In Brazil, areas affected by dam failures such as Fundão, in Mariana-MG, showed an increase in the diversity of AMF species in revegetated areas, with a predominance of Glomus and Rhizophagus species (Oliveira Prado et al., 2019). Species such as Acaulospora mellea and Paraglomus occultumwere frequent in other areas of the Iron Quadrangle or ecosystems impacted by iron mining activity (Teixeira et al., 2017; Vieira et al., 2018; Melloni et al., 2003). In addition to these, Acaulospora morrowiae has been considered efficient in promoting plant growth in soils with a high level of metals (Schneider et al., 2013). In the review by Khalid et al. (2021), studies and AMF species in the metabolization of metals and trace elements including Cu²⁺, Cd²⁺, Ni²⁺, Zn²⁺, Pb²⁺, Fe²⁺/Fe³⁺, Mn²⁺ and Co²⁺ were related. For example, the direct influences of AMF on chromium (Cr) translocation and transformation in the soil-plant continuum have been studied, showing that AMF can immobilize quantities of Cr via reduction of Cr⁶⁺ to Cr³⁺, forming Cr³⁺-phosphate analogues (Wu et al., 2015). We highlight the species: Glomus aureum, Rhizophagus irregulares, and Rhizophagus clarus, which increased the accumulation of Cd in Iris pseudacorus plants (Weżowicz, 2015). R. irregulares also regulated the composition of the rhizosphere bacterial community and promoted the growth of potentially beneficial microorganisms, thereby enhancing the resistance of perennial ryegrass (Lolium perenne L.) to Cd stress (Yang et al., 2024). Plants vetiver grass Chrysopogon zizanioides (L.), colonized by Glomus mosseae exhibit better growth (increase in plant biomass), but also significantly increase Pb uptake in root and higher translocation to the shoot (Punamiya et al., 2010). Analysis of the R. irregularis genome revealed the presence of the Fe/Zn influx mediator (RiFTR1, RiZRT) (Tisserant et al., 2013; Tamayo et al., 2014), but there is currently no evidence as to its exact location and specificity. We speculate that these transporters are involved in the homeostasis of metals.

9 Conclusion

Including plant growth-promoting fungi into lunar or Martian regolith-based agriculture systems would present a strategic enhancement to space crop production and the establishment of human settlements beyond Earth. Fungi such as Trichoderma and the various AMF (Glomeromycota) stand out for their ability to relieve abiotic stresses, mobilize essential nutrients, and potentially improve the physicochemical structure of regolith substrates. These microorganisms offer a promising biotechnological tool to transform the regolith environment (inorganic composition) and positively impact the engineered microbiome introduced to inhospitable substrates. This impact may positively affect the germination rate of seeds, the growth and development of seedlings, flowers, fruits, and positively impact the rhizosphere and the modification of the regolith. The different modes of action in iron metabolization by Trichoderma and AMF fungi that occur in the rhizosphere of plants on Earth are summarized in Figure 1, illustrating the hypothesis that, in lunar and Martian regolith, they may act in a similar way and promote improvement in plant growth, development and health, as well as stress tolerance. However, to harness this potential there are several knowledge gaps which must be resolved. The main gaps involve: (i) the need for growth studies under controlled conditions involving, microgravity, cosmic radiation, extreme temperatures, CO₂ levels, since how these particular fungi tolerate or respond to exposure to space conditions is poorly characterized; (ii) the development of standardization protocols for evaluating the symbiotic viability of fungi throughout the life cycle of plants in high-fiedlity regolith simulants; (iii) evaluation of the biosafety of fungal strains, especially those with a history of pathogenicity; and (iv) the improvement of inoculation technologies compatible with agricultural systems in closed and extreme environments.

FIGURE 1

Furthermore, additional tests are required for fungi of genera such as Trichoderma, Beauveria and Penicillium as well as AMF genera such as Rhizophagus, Glomus and Claroideoglomus, as these have previously demonstrated the potential for improving metal tolerance and plant growth promotion (Atkin and Oliveira Pedro dos Santos, 2024; Abdelrhim, 2024; Sui et al., 2023; Bello et al., 2022; Tamayo et al., 2018; Weżowicz, 2015). It is essential to validate these interactions under realistic cultivation conditions, which combine the physical, chemical and biological challenges of regolith based agriculture, most notably by following growth from germination to harvest through multiple generations.

We propose the implementation of an integrated research programs that combines the selection of fungal species capable of promoting plant growth as well as the bioweathering/mitigation of regolith as a substrate. Such a research pipeline begins with the development of standardized methods for the introduction and phenotypic analysis of these inoculants in high fidelity regolith simulants. Likely target species can then be incorporated into bacterial consortia currently being developed to support plant growth both by regolith-based agriculture as well as hydroponic systems. Once these approaches have been developed, the introduction of additional abiotic factors that simulate extraterrestrial conditions (pressure, temperature, atmospheric composition, etc.) will determine when and how fungi and their partners (plant or bacterial) could be incorporated into mission planning. This integrated interdisciplinary approach, will contribute not only to food security and other aspects of bioregenerative life support in future space missions but provide sustainable solutions for the recovery of degraded soils on Earth bridging the divide between terrestrial and space agriculture.

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