Can lunar and martian soils support food plant production? Effects of horse/swine monogastric manure fertilisation on regolith simulants enzymatic activity, nutrient bioavailability, and lettuce growth
Open accessAbstract
To make feasible the crewed missions to the Moon or Mars, space research is focusing on the development of bioregenerative life support systems (BLSS) designed to produce food crops based on in situ resource utilisation (ISRU), allowing to reduce terrestrial input and to recycle organic wastes. In this regard, a major question concerns the suitability of native regoliths for plant growth and how their agronomic performance is affected by additions of organic matter from crew waste. We tested plant growth substrates consisting of MMS-1 (Mars) or LHS-1 (Lunar) simulants mixed with a commercial horse/swine monogastric manure (i.e., an analogue of crew excreta and crop residues) at varying rates (100:0, 90:10, 70:30, 50:50, w/w). Specifically, we measured: (i) lettuce (Lactuca sativa L. cultivar ‘Grand Rapids’) growth (at 30 days in open gas exchange climate chamber with no fertilisation), plant physiology, and nutrient uptake; as well as (ii) microbial biomass C and N, enzymatic activity, and nutrient bioavailability in the simulant/manure mixtures after plant growth. We discussed mechanisms of different plant yield, architecture, and physiology as a function of chemical, physico-hydraulic, and biological properties of different substrates. A better agronomic performance, in terms of plant growth and optically measured chlorophyll content, nutrient availability, and enzymatic activity, was provided by substrates containing MMS-1, in comparison to LHS-1-based ones, despite a lower volume of readily available water (likely due to the high-frequency low-volume irrigation strategy applied in our experiment and foreseen in space settings). Other physical and chemical properties, along with a different bioavailability of essential nutrients for plants and rhizosphere biota, alkalinity, and release of promptly bioavailable Na from substrates, were identified as the factors leading to the better ranking of MMS-1 in plant above and below-ground mass and physiology. Pure Mars (MMS-1) and Lunar (LHS-1) simulants were able to sustain plant growth even in absence of fertilisation, but the amendment with the monogastric manure significantly improved above- and below-ground plant biomass; moreover, the maximum lettuce leaf production, across combinations of simulants and amendment rates, was obtained in treatments resulting in a finer root system. Increasing rates of monogastric manure stimulated the growth of microbial biomass and enzymatic activities, such as dehydrogenase and alkaline phosphomonoesterase, which, in turn, fostered nutrient bioavailability. Consequently, nutrient uptake and translocation into lettuce leaves were enhanced with manure supply, with positive outcomes in the nutritional value of edible biomass for space crews. The best crop growth response was achieved with the 70:30 simulant/manure mixture due to good availability of nutrients and water compared to low amendment rates, and better-saturated hydraulic conductivity compared to high organic matter application. A 70:30 simulant/manure mixture is also a more sustainable option than a 50:50 mixture for a BLSS developed on ISRU strategy. Matching crop growth performance and (bio)chemical, mineralogical, and physico-hydraulic characteristics of possible plant growth media for space farming allows a better understanding of the processes and dynamics occurring in the experimental substrate/plant system, potentially suitable for an extra-terrestrial BLSS.
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Previous studies by the authors on Martian and Lunar regolith simulants showed that both pure regoliths are unsuitable to sustain plant growth even with fertigation.
“Results revealed that both pure regoliths are unsuitable to sustain the plant growth, even in the presence of fertigation”
From Fava bean plant performance and fertility dynamics in Mars regolith simulant-based substrates for space farming (2025) · cites this paper as Caporale et al., 2022
The Conclusions (page not provided) state the opposite: pure Mars and Lunar simulants sustained lettuce growth even without fertilisation. The paper notes that their properties can hinder growth and that manure improves it, but does not say the pure simulants cannot sustain growth with fertigation. (LLM, conf=0.90)
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Caporale, A. G., Paradiso, R., Liuzzi, G., Palladino, M., Amitrano, C., Arena, C., Arouna, N., Verrillo, M., Cozzolino, V., De Pascale, S., et al. (2023) · Plant and Soil · doi:10.1007/s11104-022-05860-0 · Cited by 27 · see citing works
Self-citationBoudjabi, S., & Chenchouni, H. (2023) · Heliyon · doi:10.1016/j.heliyon.2023.e14615 · Cited by 25 · see citing works
Duri, L. G., Pannico, A., Petropoulos, S. A., Caporale, A. G., Adamo, P., Graziani, G., Ritieni, A., Pascale, S. D., & Rouphael, Y. (2022) · Frontiers in Nutrition · doi:10.3389/fnut.2022.890786 · Cited by 21 · see citing works
Self-citationDuri, L. G., Romano, I., Adamo, P., Rouphael, Y., Pannico, A., Ventorino, V., Pepe, O., Pascale, S. D., & Caporale, A. G. (2025) · Biology and Fertility of Soils · doi:10.1007/s00374-025-01923-3 · Cited by 12 · see citing works
Self-citationRaimondo, M., Simeone, G. D. R., Coppola, G. P., Zaccardelli, M., Caracciolo, F., & Rao, M. A. (2023) · Journal of Agriculture and Food Research · doi:10.1016/j.jafr.2023.100845 · Cited by 11 · see citing works
Self-citationCaporale, A. G., Paradiso, R., Liuzzi, G., Arouna, N., Pascale, S. D., & Adamo, P. (2022) · Plants · doi:10.3390/plants12010064 · Cited by 8 · see citing works
Self-citationCaporale, A. G., Paradiso, R., Palladino, M., Arouna, N., Izzo, L., Ritieni, A., De Pascale, S., & Adamo, P. (2024) · Plants · doi:10.3390/plants13050747 · Cited by 7 · see citing works
Self-citationVezzola, M., Tosi, S., Doria, E., Bonazzi, M., Alvaro, M., & Sanfilippo, A. (2023) · Journal of Fungi · doi:10.3390/jof9100976 · Cited by 7 · see citing works
Space Agriculture and Mechatronic Technologies: Micro-Review and Multi-Collaborative Study
Barreto, R., Cornejo, J., Palomares, R., Cornejo, J., Suárez-Quispe, J. C., Vargas, M., Valenzuela, C., Chavez, J. C., & Valdivia, J. (2023) · International Conference for Innovation in Technology (INOCON) · doi:10.1109/inocon57975.2023.10101312 · Cited by 7 · see citing works
Moss-derived microbiome improves crop growth in lunar and Martian soil simulants
Park, J., Kang, D., Jang, J. W., Choi, S., Jeong, S., Kim, S., & Kim, D. (2025) · Ecological Indicators · doi:10.1016/j.ecolind.2025.114023 · Cited by 6 · see citing works
Coker, H. R., Denvir, A. C., Robertson, I. J., Shackelford, C. E. B., Li, W., Lin, C.-W., Watters, R. M., Sparks, D. L., Smith, A. P., & Howe, J. A. (2025) · ACS Earth and Space Chemistry · doi:10.1021/acsearthspacechem.4c00312 · Cited by 5 · see citing works
Emerging Chemical and Biological Materials Technologies in the Extraplanetary Environment
Jiang, Q., Wang, B., Cheng, Y., Wang, Y., Zhao, H. F., & Lu, Y. (2026) · Nano-Micro Letters · doi:10.1007/s40820-025-01979-8 · Cited by 5 · see citing works
Mei, C., Xie, G., Ya, H., & Peng, M. (2025) · Acta Astronautica · doi:10.1016/j.actaastro.2025.09.033 · Cited by 5 · see citing works
Zhao, Y., Luo, R., Zhang, H., Li, Y., Fang, X., Tong, X., Qian, Y., Zhou, Z., Yang, Y., Zeng, X., et al. (2024) · Plant and Soil · doi:10.1007/s11104-024-06970-7 · Cited by 4 · see citing works
Liao, B., Yao, Z., Feng, J., & Liu, H. (2025) · Acta Astronautica · doi:10.1016/j.actaastro.2025.06.018 · Cited by 3 · see citing works
Cheatham, R. W., Sultana, A. I., & Reza, M. T. (2025) · Journal of Analytical and Applied Pyrolysis · doi:10.1016/j.jaap.2025.107064 · Cited by 3 · see citing works
Biochar improves lettuce seedling growth by influencing the nutrient content of simulated lunar soil
Mei, C., Xie, G., Ya, H., & Peng, M. (2025) · iScience · doi:10.1016/j.isci.2025.113327 · Cited by 3 · see citing works
A systematic review of plant cultivation and growth in extraterrestrial regolith simulants
Elkamouh, A., Chami, D. E., Abdelmouneim, A. A., Khadra, R., & Pascale, S. D. (2026) · Life Sciences in Space Research · doi:10.1016/j.lssr.2026.04.005 · Cited by 2 · see citing works
Servetto, G. P., Botta, P., Rossi, G., Bernasconi, A., Caldo, F., Vigliaturo, R., & Maffei, M. E. (2026) · Scientific Reports · doi:10.1038/s41598-026-50840-4 · Cited by 2 · see citing works
Caporale, A. G., Paradiso, R., Arouna, N., Amitrano, C., Tagliamonte, S., Chiacchio, M. F., Vitaglione, P., De Pascale, S., & Adamo, P. (2025) · Frontiers in Plant Science · doi:10.3389/fpls.2025.1676285 · Cited by 1 · see citing works
Self-citationNedbalová, L., Mihál, M., Procházková, L., Kvíderová, J., & Lukavský, J. (2025) · Czech Polar Reports · doi:10.5817/cpr2024-2-19 · Cited by 1 · see citing works
Hou, S., Wang, Z., Zhu, Y., Liu, H., & Feng, J. (2025) · Acta Astronautica · doi:10.1016/j.actaastro.2025.09.080 · Cited by 1 · see citing works
Mansouri, F., Barros, A. N., Bordoni, L., Fedeli, D., Pontetti, G., Roncolato, E., Mannozzi, C., Pompei, F., Vittori, S., Pettinelli, E., et al. (2025) · Life Sciences in Space Research · doi:10.1016/j.lssr.2025.11.008 · Cited by 1 · see citing works
Exploring differential gene correlations in Arabidopsis thaliana for sustainable space farming
D’Annunzio, F., Fiscon, G., Conte, F., Paci, P., Bianco, M. D., & Ioio, R. D. (2026) · Biotechnology Reports · doi:10.1016/j.btre.2026.e00949 · Cited by 1 · see citing works
Comparison of two laser wavelengths for LIBS bioimaging of plants grown in lunar regolith
Vozár, T., Čechová, L., Buday, J., Fuleky, M., Molnárová, K., Lenža, L., Mužík, J., Koshiba, Y., Smrž, M., Pořízka, P., et al. (2026) · Spectrochimica Acta Part B Atomic Spectroscopy · doi:10.1016/j.sab.2026.107513 · Cited by 1 · see citing works
Simeone, G. D. R., Scala, G., Scarpato, M., & Rao, M. A. (2023) · Heliyon · doi:10.1016/j.heliyon.2023.e22894
Self-citationRetractedFlórez-Velasco, N., Sánchez-Reinoso, A. D., Bernal-Romero, S., Marín-Peña, Y. A., Peña-Luna, C. R., & Fischer, G. (2023) · International Journal of Vegetable Science · doi:10.1080/19315260.2023.2289977
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Self-citationSingh, J., Durga, A. V. U., Bodh, S., Vikanksha, V., Bakshi, M., & Verma, P. K. (2024) · Agricultural Science Digest - A Research Journal · doi:10.18805/ag.d-5903
Infrastructure and Master Planning
Inocente, D. (2025) · Wiley eBooks · doi:10.1002/9781394318131.ch17
Mendoza, J. E., Lemke, N. B., & Tomberlin, J. K. (2026) · New Space · doi:10.1177/21680256261504179
Sustainable plant cultivation in Lunar regolith: Feasibility study
Dias, J. E. L., May, Z., Mohai, M. P., & Barkó, G. (2026) · Progress in Agricultural Engineering Sciences · doi:10.1556/446.2026.00321
Agricultural potential of lunar regolith – a review on cultivation methodologies
Dias, J. E. L., Oszlányi, R., & Barkó, G. (2026) · The Journal of Agricultural Science · doi:10.1017/s0021859626100811
Oroian, I., & Tripathi, A. (2026) · Extreme Life Biospeology & Astrobiology · doi:10.67784/0043
Akuku, V., Loredo, A., Howe, J. A., Sparks, D., Knappett, P. S. K., Northrup, P., & Szerlag, K. D. (2026) · ACS Earth and Space Chemistry · doi:10.1021/acsearthspacechem.6c00270
Addesso, R., Caporale, A. G., Adamo, P., Labella, R., Bochicchio, R., Rossi, R., & Amato, M. (2026) · Frontiers in Soil Science · doi:10.3389/fsoil.2026.1999715
Self-citation
- Authors
- Caporale, A. G., Amato, M., Duri, L. G., Bochicchio, R., De Pascale, S., Simeone, G. D. R., Palladino, M., Pannico, A., Rao, M. A., Rouphael, Y., & Adamo, P.
- Affiliations
- University of Naples Federico II (IT)
- University of Basilicata (IT)
- Department of Agricultural Sciences, University of Naples Federico II, 80055 Portici, Italy
- Corresponding author
- Caporale, Antonio G.
- Rouphael, Youssef
- ORCID
- Venue
- Plants
- Publisher
- MDPI AG
- Published
- 2022-12-02
- Volume
- 11
- Issue
- 23
- Pages
- 3345
- ISSN
- 2223-7747 (ISSN-L)
- PMC
- PMC9740528
- OpenAlex
- W4311107385
- Cited by
- 36
- References
- 91
- Citations by year
- 2026: 11 · 2025: 12 · 2024: 4 · 2023: 7 · 2022: 2
- FWCI
- 3.37
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- Plants (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- CINECA IRIS Institutional Research Information System (University of Basilicata) (submittedVersion, other-oa)
- Funders
- Italian Space Agency (F74I16000000005)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Soil Science › Plant Growth and Agriculture Techniques
- Physical Sciences › Engineering › Aerospace Engineering › Space Exploration and Technology
- Life Sciences › Agricultural and Biological Sciences › Soil Science › Agricultural Science and Fertilization
- Keywords
- lunar regolith simulant, bioregenerative life support, in-situ resource utilization, lettuce growth, nutrient bioavailability, microbial biomass, plant nutrient uptake, water availability, sodium release, space agriculture, chlorophyll
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