The climate impact and land use of cultivated meat: Evaluating agricultural feedstock production

forests, wetlands, or savannas. When these ecosystems are altered or converted, for instance into crop fields or pastures, the carbon that could have been sequestered by the native vegeta- tion is a forgone benefit and therefore must be considered as an opportunity cost when quanti- fying the climate impact of a food product [104]. Searchinger et al. (2018) [68] suggest using the “carbon cost of consumption” (CCC) to evaluate the climate impact of food products. CCC is the sum of production emissions and car- bon opportunity cost of a consumed product. Most previous studies only focus on production emissions. By factoring in carbon opportunity cost, we can quantify the climate impact of not only the production emissions from agriculture, but also the forgone benefit of native vegeta- tion. This is especially relevant when comparing conventional animal agriculture with alterna- tive food production methods like plant-based meat alternatives or CM, which potentially require less land and thereby incur lower carbon opportunity cost. Incorporating carbon opportunity cost into climate impact assessments offers a more nuanced and comprehensive understanding of the climate impact of food production systems, particularly highlighting the value of native vegetation for climate change mitigation [68, 105]. The foregone carbon sequestrations potential of arable land in Germany is assumed with 2.9 t CO2 ha-1 yr-1. Multiplying this factor with 44/12, which is the conversion factor from C to CO2 [106], and the land requirements results in the carbon opportunity cost per output. The sum

of total production emissions and carbon opportunity cost results in the carbon cost of con- sumption. Data for environmental impact of soy was taken from DonauSoja (2022) [107] and data of foregone carbon sequestration of arable land was taken from Fehrenbach & Bu¨rck (2022) [108]. 4. Results The analysis of CM production, factoring in energy consumption and land use, yields compel- ling insights. We have found that solar energy integration and land utilization, coupled with crop rotation restrictions, play a critical role in the sustainability and feasibility of CM. In the ensuing results and discussion section, we elaborate on these findings, positioning them in the context of the existing body of research. The crops’ share in the 100-hectare reference land area following linear optimization is shown in Fig 1. In terms of crop distribution, all four scenarios utilized soybeans with a maxi- mum cultivation area of 50 hectares. The cultivation of rapeseed was maximized within crop rotation limitations at 20 hectares for all scenarios. For wheat, scenario 1 had a cultivation area of 13.33 hectares, while scenario 2 had a cultivation area of 7.59 hectares. In contrast, scenarios 3 and 4 had a wheat cultivation area of 13.33 hectares and 8.75 hectares, respectively. For maize, scenario 1 had a cultivation area of 8.10 hectares and scenario 2 had a cultivation area of 9.63 hectares. In contrast, scenarios 3 and 4 have a cultivation area of 0 hectares for maize. Sunflower was cultivated at 13.33 hectares in scenarios 3 and 4 but was not cultivated in sce-

narios 1 and 2. Potato cultivation was 8.57 hectares for scenario 1, 7.04 hectares for scenario 2, and 3.33 hectares for scenarios 3 and 4. In terms of energy production, scenario 2 and 4 uti- lized solar panels with an area devoted to solar panels amounting to 5.74 hectares and 4.58 hectares, respectively. A comparison of different scenarios for CM production and traditional pork farming was conducted, presented in Table 7. We made use of the review by Gislason et al. (2023) [21] to compare scenario outcomes to pork production worldwide. The review analyzed 31 LCAs cal- culating land use efficiency for pork production. Boxplots for land use efficiency reported in CM LCAs, Gislason et al. (2023) [21] and the scenarios of this study are visualized in Fig 2. PLOS ONE Evaluating agricultural feedstock for cultivated meat PLOS ONE | https://doi.org/10.1371/journal.pone.0316480 January 22, 2025 15 / 27

The results showed differences in land use efficiency across the four scenarios. Scenario 1, which involved maximum CM production without solar panel energy and includes the extrac- tion of all nutrients from the crops, had the highest efficiency of 1,785.16 kg/ha. Scenario 2, which added solar panels to the mix, showed a slightly lower efficiency of 1,727.14 kg/ha. Sce- narios 3 and 4, which sourced nutrients only from respective protein and glucose plants, had efficiencies of 1,378.28 kg/ha and 1,378.28 kg/ha respectively. With scenarios 3 and 4, the pro- duction was inhibited by amino acid availability, more precisely by the lack of availability of lysine and alanine. Table 7 lists results of various LCAs to calculate a corrected area efficiency to fit various assumption of the observed LCAs. We used a carcass yield of 79% following Reckmann et al. (2013) [98]. On average, pork production also needs more land per kilogram than CM production look- ing at existing LCAs. Therefore, we can generally report slightly higher land use performance for CM which was also reported in aforementioned LCAs. To calculate the carbon cost of consumption, which are the sum of production emissions and carbon opportunity cost [68], we used production emissions per area for the crops that are used to create the pig feed concentrates and the cell medium for each of the four scenarios. The results are shown in Table 8. Given the land use requirements for feed and cell mediums, we calculate the total production emissions per area. This result is combined with the output

per area to derive the feed concentrate and cell medium production emissions. Regarding energy use, for pigs we use data based on Wirsenius et al. (2020) [105]. Manure management and enteric fermentation numbers for the pig farm are from Wirsenius et al. (2020) [105]. For CM, we assume no emissions in these two categories. All four mentioned categories of produc- tion emissions make up the total production emissions. Besides production emissions, we con- sider the carbon opportunity cost to quantify the carbon cost of consumption [68] for meat from pigs and the various CM scenarios. Fig 1. Crop rotation for maximum cell media production in the four scenarios. https://doi.org/10.1371/journal.pone.0316480.g001 PLOS ONE Evaluating agricultural feedstock for cultivated meat PLOS ONE | https://doi.org/10.1371/journal.pone.0316480 January 22, 2025 16 / 27

The results indicate that when considering production emissions and carbon opportunity cost, scenarios 1 and 2 show slightly lower carbon cost of consumption (9.0 and 9.1 t CO2e t-1) as well as scenarios 3 and 4 (10.8 and 10.7 t CO2e t-1) compared to the emissions of German pork production (11.24 t CO2e t-1) reported by Wirsenius et al. (2020) [105]. 5. Discussion 5.1 Comparison with existing literature Our investigation into CM production in southern Germany reveals limited land use advan- tages when compared with traditional pork production when feedstock is solely taken from crops. Our findings contrast with the assertions of several previously published studies which have not taken into account crop rotation restrictions or pure plant-based cell media [40, 41, 50]. The examined scenarios highlight that crop rotation restrictions along with protein and sugar content of specific crops are crucial factors determining the land use efficiency of CM. If CM were fed by cell media produced from plant-based crops to a large extent, land use effi- ciency would not be substantially better than it would be for conventional pork production. It is important to keep in mind that the cell culture medium accounts for the highest environ- mental impact in CM production [42]. The higher energy demand of CM production translates into additional land requirements when renewable energy sources, namely ground-mounted photovoltaic systems, are employed. Table 7. Comparison of land use efficiency of different European pork production systems and cultivated meat production.

Scenario/ LCA Protein source Energy source Solar panel use Area efficiency (m2/kg) Scenario 1 (max. extraction/ no solar energy) Maximum yield from all crops Maximum yield from crops No 5.6 Scenario 2 (max. extraction/ solar energy) Maximum yield from all crops Maximum yield from crops Yes 5.79 Scenario 3 (specific extraction/no solar energy) Proteins only from protein crops Sugar from energy crops No 7.26 Scenario 4 (specific extraction/solar energy) Proteins only from protein crops Sugar from energy crops Yes 7.26 Mean 6.48 Examples from LCAs for Cultivated Meat Production Smetana et al., 2015 Cyanobacteria Cyanobacteria - 0.4 Vergeer et al., 2021 Soy Maize - 1.8 Sinke et al., 2023 75% Soy hydrolysate/ 25% chemical & microbial production Maize 2.48 Mattick, et al., 2015 Soy Maize - 5.4 Tuomisto et al., 2022 Soy Maize - 6.89 Mean (without ranges) 3.39 Average from LCAs for pork production Gislason et al., 2023 31 studies from various countries 8.02 (adjusted from live weight) Mean (without ranges) 7.93 Note: m2/kg = square meters per kilogram; LCA = Life Cycle Assessment. https://doi.org/10.1371/journal.pone.0316480.t007 PLOS ONE Evaluating agricultural feedstock for cultivated meat PLOS ONE | https://doi.org/10.1371/journal.pone.0316480 January 22, 2025 17 / 27

This is due to the additional land that solar panels would require. Since land use is a critical determinant of the climatic impact of pork (up to 75%) [105], substantial environmental bene- fits from CM in this aspect cannot be readily anticipated when agricultural feedstock is used. Also, for the carbon cost of consumption (production emissions and carbon opportunity cost) there is only a limited advantage of the considered CM scenarios in comparison to pig farming in Germany. In previous LCAs Sinke et al., 2023 brings up freed up land, which can only be realized if precision fermentation is employed as they use 25% for media production through biotechnology and 75% amounts of amino acids through soy hydrolysate which again on a global scope would not reflect a sustainable crop rotation [41]. As a consequence, public and private research needs to focus on CM production systems where proteins for cell media feed- stock are produced through synthetic processes independent from agricultural products, like precision fermentation [61, 70]. Precision fermentation is essential for supplying nutrients to CM bioreactors, but scaling this technology presents major obstacles. Global fermentation capacity is currently inadequate to meet future demands, and engineering highly productive, robust microbial strains remains complex and costly [109]. Production costs are still too high, and significant regulatory hurdles persist, although they are somewhat less severe than those for CM [110]. Ensuring the purity of amino acids for bioreactor use is another challenge, particularly

when crop-based substrates require extensive refinement, potentially leading to higher cost in precision fermentation [111, 112]. Additionally, fermentation and chemical processes are water-intensive, complicating sustainability efforts [41]. Fig 2. Boxplots of land use efficiency for previous cultivated meat LCAs, pork production LCAs, and cultivated meat scenarios from this study. https://doi.org/10.1371/journal.pone.0316480.g002 PLOS ONE Evaluating agricultural feedstock for cultivated meat PLOS ONE | https://doi.org/10.1371/journal.pone.0316480 January 22, 2025 18 / 27

5.2 Limitations While our study provides valuable insights into the land use implications of CM, it is impor- tant to note a few limitations. Our case study analysis is based on southern Germany, so the results may not be representative of other geographical locations with different agricultural systems, climates, and land accessibility. More agricultural crops could be grown and used as media bases, even in a German context. However, every agricultural site offers different basic requirements. We limited our model to a set of regular crops used in Germany. Furthermore, our data was based on German yield averages. Future integrated and regener- ative agricultural practices could lead to different results, although lower crop yields and there- fore lower sugar and amino acid yields are to be expected with these cultivation methods [113]. Moreover, conservation agriculture does not generally lead to N2O emission mitigation [114]. Table 8. Land use, carbon opportunity cost, and carbon cost of consumption of 100 ha arable land for cultivated meat. Scenario 1 Scenario 2 Scenario 3 Scenario 4 Unit Crop yield Crop production emissions (max. extraction/ (max. extraction/ (specific extraction/ (specific extraction/ no solar energy) solar energy) no solar energy) solar energy) t/ha t CO2e t-1 fw Sources Land use Soybeans 50.0 50.0 50.0 50.0 ha 3.2 0.4 LFL, 2023; DonauSoja, 2022 Winter oilseed rapeseed 20.0 20.0 20.0 20.0 ha 3.6 0.5 LFL, 2023 Sunflower - - 13.3 13.3 ha 2.7 0.8 LFL, 2023 Maize for grain 8.1 9.6 - - ha 10.5 0.3 LFL, 2023

Winter wheat 13.3 7.6 13.3 8.8 ha 7.5 0.3 LFL, 2023 Potatoes 8.6 7.0 3.3 3.3 ha 42.3 0.2 LFL, 2023 Photovoltaic cells - 5.7 - 4.6 ha Bo¨hm et al., 2022 Total land use 100.0 100.0 100.0 100.0 ha Production emissions per area 2.2 2.0 1.8 1.7 t CO2e ha-1 Own calculation Land requirement per output 5.6 5.8 7.3 7.3 m2/kg Own calculation Output per area 1.8 1.7 1.4 1.4 t/ha Own calculation Production emissions per output Feed concentrates or medium production emissions 1.2 1.2 1.3 1.2 t CO2e t-1 Own calculation Energy use, scaffold, equipment & other 1.8 1.8 1.8 1.8 t CO2e t-1 Sinke et al., 2023 Total production emissions 3.0 3.0 3.1 3.0 t CO2e t-1 Own calculation Carbon Opp. Cost Foregone carbon sequestration of arable land 2.9 2.9 2.9 2.9 t C/ ha*a Fehrenbach & Bu¨rck, 2022 Foregone carbon sequestration of arable land 10.6 10.6 10.6 10.6 t CO2/ ha*a Own calculation Carbon opportunity cost 6.0 6.2 7.7 7.7 t CO2e t-1 Own calculation Carbon cost of consumption 9.0 9.1 10.8 10.7 t CO2e t-1 Own calculation https://doi.org/10.1371/journal.pone.0316480.t008 PLOS ONE Evaluating agricultural feedstock for cultivated meat PLOS ONE | https://doi.org/10.1371/journal.pone.0316480 January 22, 2025 19 / 27

The economic feasibility of CM production is heavily influenced by the cost of growth media, which remains a major financial bottleneck. Current estimates indicate that media expenses account for up to 99% of material costs, driven by expensive amino acids and growth factors produced through precision fermentation or chemical synthesis [37, 41, 115]. Even as bioreactor scale increases, reducing media costs is critical to achieving competitive prices with conventional meat [116]. While precision fermentation has lower land use impacts, its com- plexity and high production costs makes it still a challenge. Chemical production of growth factors adds to this economic burden. Emerging research suggests that plant-based media could offer a more cost-effective solution, though this remains speculative as hydrolysates have not fully proven feasibility for cell media formulation [37, 115]. The study assumes direct extraction of amino acids and glucose from crops, which oversim- plifies real-world processes and might overestimate yields, thus affecting production efficiency and land use metric. Additionally, incorporating biotechnological methods for amino acid production could realistically enhance efficiency. Other studies using precision fermentation technologies showed more optimistic scenarios that could be implemented in the future. Those hybrid media formulations could decrease land use and efficiency [41]. On the other hand, involving synthetic amino acid supply for pig feed would also boost efficiency of land use for pork production [117, 118].

In general, we did not consider the upcycling of by-products from food processing, such as soybean meal, rapeseed meal, or sugar beet pulp. There may still be potential to increase land use efficiency in the production of plant-based cell media. Many co-products are created by industrial food processing leading to sugar and protein rich feed that could be used for culti- vated as they are used for animal feed now. But many sustainability criteria must be reconsid- ered for those side streams anew [119]. While land use and emissions could be decreased, economic feasibility often remains a constraint when it comes to adopting side stream prod- ucts and their usage. Currently available solutions include alternatives to fermentable sugar, nitrogen, and phosphates, which could also be converted into proteins through precision fer- mentation [120]. During the production of hydrolysates and glucose from crops, byproducts are generated including solid residues (stillage), fiber, proteins, lipids, ash, and in the case of maize, maize gluten meal, maize gluten feed, and maize oil. One limitation of our study lies in the fact that we did not account for the metabolites [50] produced by cells during the CM pro- duction process. These metabolites, primarily consisting of compounds such as urea, ammo- nia, lactic acid, and other organic acids [121], could potentially be utilized as a form of manure or soil conditioner. This factor was not incorporated into our current model and could repre- sent an area of interest for further investigation.

In terms of energy, CM comes with higher energy needs than animal meats, including pork [39]. We only considered land use scenarios if solar energy was used, leading to direct influ- ence on land use. While it might use less land per output, the use of fossil energy for produc- tion of CM would have a far greater impact on the environment [122]. Furthermore, the model considers only land use and does not account for other critical environmental impacts, such as water usage, influence on biodiversity, or ecosystem structures. 5.3 Future research directions There are several opportunities for future research on agricultural involvement in CM produc- tion. Agriculture will be involved in CM production in any matter. It is also critical to under- stand the social acceptance and economic viability of CM. Thus, future studies could focus on consumer perceptions, market dynamics, and the broader socio-economic implications of a shift from conventional meat production to CM. PLOS ONE Evaluating agricultural feedstock for cultivated meat PLOS ONE | https://doi.org/10.1371/journal.pone.0316480 January 22, 2025 20 / 27

Future research could build upon the findings of this study by examining the environmen- tal impacts of CM in a more comprehensive and holistic manner, such as by incorporating a range of sustainability metrics beyond land use and energy consumption. To examine CMs full sustainability impacts this will be crucial. This could include additional factors, such as water use, biodiversity impacts, and further potential for carbon sequestration. Given the constraints discussed above regarding information on extraction efficiencies, research is needed to provide more precise estimates of nutrient extraction efficiencies. This would enhance the accuracy and relevance of the model and therefore the results. Taken together, these research directions would contribute to a more comprehensive understanding of the potential of CM as a sustainable alternative to conventional meat production and pro- vide valuable insights for policy makers, industry stakeholders, and consumers alike. 6. Conclusions This study evaluates the potential role of agriculture to produce the feedstock for CM produc- tion. Using a linear programming model, we investigated different scenarios of CM produc- tion based on agricultural feedstock in southern Germany, considering crop rotation restrictions and energy requirements, particularly the integration of solar energy. Our findings indicate that CM production using crops as raw materials for cell media does not offer significant land use efficiency improvements over conventional pork production.

Scenarios with maximum nutrient extraction from crops showed only minor land use benefits, while scenarios relying on specific crops for glucose and amino acids were less efficient. Addi- tionally, the higher energy demands of CM production, especially when integrating solar energy, translate into further land requirements, thus not significantly advancing climate change mitigation. This study suggests that agriculture’s role in directly providing protein feedstock for CM production should be small to ensure sustainability of CM. Policymakers and industry stake- holders should focus on exploring advanced biotechnological methods i.e. precision fermenta- tion for cell media production to maximize the efficiency of CM. Further research should investigate systems where protein sources are based on precision fermentation rather than directly from crops. Author Contributions Conceptualization: Peter Breunig. Data curation: Hanno Kossmann, Thorsten Moess. Formal analysis: Hanno Kossmann, Thorsten Moess. Methodology: Hanno Kossmann. Visualization: Hanno Kossmann. Writing – original draft: Hanno Kossmann. Writing – review & editing: Hanno Kossmann, Peter Breunig. References 1. Larsen CS. Animal source foods and human health during evolution. J Nutr. 2003; 133(11):3893S– 3897S. https://doi.org/10.1093/jn/133.11.3893S PMID: 14672287 2. Murimi M. What is the place of beef in a healthful sustainable Homo sapiens diet? Am J Health Promot. 2022; 36(5):898–900. https://doi.org/10.1177/08901171221088661c PMID: 35531995 PLOS ONE Evaluating agricultural feedstock for cultivated meat

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