Black Soldier Fly Larvae: A Sustainable Solution to Antimicrobial Resistance in Livestock Farming

An article about antimicrobial resistance in livestock farming by Ishaya Gadzama & Penuel Panuel

Introduction 

Black soldier fly larvae (BSFL) have gained attention for their potential as a valuable source of protein and fat for poultry, pigs, and aquaculture (Singh & Kumari, 2019; Veldkamp et al., 2022; Gadzama, 2024). The larvae are not only a source of nutrients but also possess antimicrobial peptides (AMPs) and bioactive molecules that could enhance animal health (Veldkamp et al., 2022; Belhadj Slimen et al., 2023).

This process begins with the BSFL naturally growing on organic food and animal waste, exposing them to various microorganisms. Despite this, BSFL thrives due to their innate immune system (specialized defense mechanisms), which produces substances like peptides (antibacterial compounds on their exoskeleton or in their digestive tracts) that protect them against pathogenic microbes such as bacteria, fungi, and viruses (Lalander et al., 2015; Elhag et al., 2017; Feng et al., 2020; Zabulionė et al., 2023). Most recently, Elhag et al. (2022) demonstrated that BSFL can reduce the populations of many zoonotic pathogens in dairy and cattle manure, showcasing their potential in waste management and disease control. There are several successful case studies where BSFL has been integrated into animal feed systems, demonstrating their practicality and benefits when used on a larger scale.

Figure 1: Adult Black Soldier Fly (Hermetia illucens)

The Role of BSFL in Sustainable Agriculture

The Black Soldier Fly has emerged as a significant player in sustainable agriculture and waste management due to its larvae's ability to thrive on decomposing organic matter. Recognizing the potential of BSFL, researchers and agricultural specialists have been developing methods for optimizing the large-scale production of BSFL as an alternative to traditional feed sources (Fahmy et al., 2024). The discovery of their role in producing AMPs and other bioactive substances (Veldkamp et al., 2022) has opened new avenues in the fight against microbial resistance, further solidifying their importance in a sustainable future. 

The bioactive substances derived from insects can be categorized into three main groups: 

• Antimicrobial peptides
• Fatty acids
• Polysaccharides

The studies by Elhag et al. (2022) confirmed the potent antibacterial properties found within the BSFL's immune system, particularly in the hemolymph. This component of their immune system has shown effectiveness against various Salmonella species, indicating the potential for BSFL to contribute to healthier livestock and, by extension, a more resilient food system.

Figure 2. Antimicrobial Resistance

Source: https://www.nps.org.au/consumers/antibiotic-resistance-the-facts

What are Antimicrobial Peptides (AMPs)

Antimicrobial peptides (AMPs) are integral components of the innate immune system in all living organisms. These small, potent molecules, typically fewer than 50 amino acids in length and with molecular weights of around 10 kDa, serve as a natural defense mechanism (Huan et al., 2020). Insects, for instance, rely on a diverse group of AMPs to fend off bacterial, fungal, and viral invaders. It's like having an internal army ready to combat microscopic threats at a moment's notice. However, the overuse of antibiotics in medical and agricultural applications has led to the development of a new challenge: antimicrobial resistance (AMR). Pathogens began to outsmart the drugs designed to kill them, thereby posing a significant challenge to public health (Llor & Bjerrum, 2014). 

Unlike traditional antibiotics, which pathogens can become resistant to, emphasizing the need for alternative strategies to combat pathogens effectively without contributing to the rise of AMR. 

Antimicrobial peptides offer a multifaceted approach to attacking and neutralizing harmful microorganisms, making them a crucial subject of study for future medical breakthroughs and the ongoing battle against infectious diseases. So, while humans were busy creating antibiotics, nature had already equipped insects with an ancient and powerful form of protection: the remarkable antimicrobial peptides.

Categories of AMPs

Insects are a goldmine for antimicrobial peptides (AMPs) (Yi et al., 2014), small proteins that help them fight off infections. These AMPs are crucial for insects and have potential for use in medicine, especially as alternatives to antibiotics, given the growing problem of antibiotic resistance (Koehbach & Craik, 2019).

When insects face a microbial threat, their immune system kicks into gear, using pathways like Toll, Imd, and JAK-STAT to trigger AMP production. Structurally, insect AMPs fall into four main categories (Figure 3):

1. Linear α-helical peptides
2. β-sheet peptides
3. Linear extended peptides without α- or β-structures
4. Peptides with a combination of α and β-elements

Common AMPs Found in Insects

One insect of particular interest is the Black Soldier Fly (BSF). Its larvae produce a variety of AMPs, some of which can be enhanced by feeding them bacteria or organic materials. Researchers have identified 57 potential AMPs in BSF, and studies have confirmed their broad-spectrum antimicrobial activity against bacteria (Li et al., 2017; Xu et al., 2020; Fahmy et al., 2023), fungi, and parasites.

Among the AMPs found in insects, defensins are the most studied. These small peptides can neutralize a wide range of bacteria by disrupting their cell membranes (Koehbach et al., 2019). Other notable insect AMPs include cecropins, which also disrupt bacterial membranes, and proline-rich peptides and attacins, which interfere with bacterial functions. In BSF larvae, defensins make up the largest share of AMPs, followed by cecropins and lysozymes. These peptides offer diverse protection against pathogens  (Figure 4).

Commonly found AMP families in insects include:

• Defensins
• Cecropins
• Proline-rich peptides
• Attacins

Gloverins and moricins have been identified only in Lepidoptera (Yi et al., 2014). Each peptide plays a unique role in the insect immune system, contributing to a collective barrier against various pathogens. The mechanism of action is part of a larger pattern of AMPs' ability to target and neutralize threats to the insect host. 

Figure 4. Pie chart showing defensin as the largest peptide identified from BSFL 

Adapted from Moretta et al. (2020)

The study by Moretta et al. (2020) presents a pie chart analysis of AMPs in BSFL. It reveals that defensins are the most prevalent, constituting 44% of the AMPs identified. Cecropins and lysozymes each comprise 18%, followed by attacins at 7%. The chart also indicates that knottin-like peptides represent 5%, while alo1-like and diptericins each account for 3%. Stomoxyn-like peptides are the least represented at 2%. AMPs are diverse in nature and are categorized into families based on their structure, amino acid composition, the organism they are sourced from, and their specific activity. This classification is detailed in Figure 5. 

Figure 5. Classification of AMPs

Antimicrobial Peptides (AMPs) Derived from BSFL

Despite insects making up about 90% of all animal species, only a small fraction of known AMPs come from them, leaving much potential for discovery (Yi et al., 2014; Li et al., 2017). In recent years, the focus has shifted to BSF larvae due to their promising antimicrobial properties, particularly in treating parasitic diseases and as a replacement for antibiotics in animal feed. Studies have revealed that peptides with an α-helical structure lacking cysteine (Cys) residues are effective against malaria (Lacerda et al., 2016). In animal nutrition, AMPs from BSFL are emerging as a viable substitute for traditional antibiotics, which are losing their effectiveness due to the proliferation of antibiotic-resistant bacteria (Andersson et al., 2016).

Exploring insect-derived peptides for therapeutic applications, particularly for parasitic diseases, is still in the early stages but shows immense promise. One of the most recent discoveries in this field is a defensin-like peptide 4 (DLP4), identified and extracted from various immune tissues of BSF larvae (Li et al., 2017). Like other insect AMPs, this peptide is synthesized by ribosomes and composed of natural amino acids, ensuring that its antimicrobial action within the animal digestive system does not have systemic effects on the host animals (Andersson et al., 2016). 

The Antimicrobial Properties of BSFL Fat

Black soldier fly larvae are a rich source of lipids, with fat content ranging from 15% to 49% (Marusich et al., 2020; Zabulionė et al., 2023). BSF larvae fats also exhibit antimicrobial properties, suggesting their use as natural preservatives in food. These fats can remain stable for long periods and protect against common food pathogens.

Lauric acid (a natural antimicrobial compound) is the predominant fatty acid in BSFL (Figure 6; Shu et al., 2024). BSFL contains 40.93% lauric acid, followed by palmitic (19.11%), oleic (17.34%), linolenic (8.79%), myristic (6.49%), and palmitoleic acids (3.89%) (Zabulionė et al., 2023). 

Black soldier fly larvae fats exhibit antimicrobial properties, particularly against common food pathogens and spoilage microorganisms (Zabulionė et al., 2023). This characteristic suggests that BSFL fats could serve as a natural preservative in high-moisture, refrigerated food products, which typically have a short shelf-life or require preservatives. In addition, BSFL fats could remain stable for up to a year without signs of chemical or microbiological spoilage. 

Modes of Action of BSFL AMPs

Antimicrobial peptides (AMPs), like defensins, primarily target Gram-positive bacteria, though some also work against Gram-negative bacteria and fungi (Wu et al., 2018). Most insect-derived AMPs carry a positive charge, which helps them latch onto the negatively charged surfaces of bacterial cells, such as lipopolysaccharides in Gram-negative bacteria and teichoic acids in Gram-positive bacteria. This interaction disrupts the bacterial membrane, effectively killing the bacteria without easily allowing them to develop resistance (Luo & Song, 2021; Haidari et al., 2023).

Research by Patrulea et al. (2020) further supports that bacteria find it difficult to develop resistance to AMPs because it's challenging for them to alter their membranes to avoid disruption. Specifically, Moretta et al. (2021) found that AMPs from Black Soldier Fly larvae are cationic, meaning they carry a positive charge. These AMPs disrupt bacterial membranes through electrostatic interactions, leading to cell death. This mechanism is particularly effective because it reduces the likelihood of bacteria developing resistance, offering a significant advantage over traditional antibiotics.

Conclusion:

Black Soldier Fly Larvae (BSFL) offers immense potential to address some of the most pressing challenges in modern agriculture and healthcare. As a sustainable protein source, they help reduce the environmental impact of traditional animal feed. Moreover, their ability to produce antimicrobial peptides (AMPs) and bioactive molecules positions them as a formidable tool in the fight against antimicrobial resistance (AMR), providing a natural alternative to conventional antibiotics. The dual role of BSFL in enhancing animal health and contributing to waste management makes them a valuable asset in creating a more sustainable and resilient agricultural system. As research progresses, BSFL could play a pivotal role in shaping the future of both agriculture and medicine.

References

• Abro, Z., Macharia, I., Mulungu, K., Subramanian, S., Tanga, C. M., & Kassie, M. (2022). The potential economic benefits of insect-based feed in Uganda. Frontiers in Insect Science, 2, 968042. https://doi.org/10.3389/finsc.2022.968042
• Andersson, D. I., Hughes, D., & Kubicek-Sutherland, J. Z. (2016). Mechanisms and consequences of bacterial resistance to antimicrobial peptides. Drug Resistance Updates, 26, 43-57.
• Attia, Y. A., Bovera, F., Asiry, K. A., Alqurashi, S., & Alrefaei, M. S. (2023). Fish and Black Soldier Fly Meals as Partial Replacements for Soybean Meal Can Affect Sustainability of Productive Performance, Blood Constituents, Gut Microbiota, and Nutrient Excretion of Broiler Chickens. Animals, 13(17), 2759. https://doi.org/10.3390/ani13172759
• Batool, M., Tabinda, A. B., & Tahir, A. (2024). Black soldier fly larvae feed suitability for chicken and prospects for economic valuation and viability aspects. Journal of Insects as Food and Feed, 1(aop), 1-21.
• Belhadj Slimen, I., Yerou, H., Ben Larbi, M., M'Hamdi, N., & Najar, T. (2023). Insects as an alternative protein source for poultry nutrition: a review. Frontiers in Veterinary Science, 10, 1200031. https://doi.org/10.3389/fvets.2023.1200031
• Bosch, G., & Swanson, K. S. (2021). Effect of using insects as feed on animals: pet dogs and cats. Journal of Insects as Food and Feed, 7(5), 795-805.
• Dabbou, S., Ferrocino, I., Gasco, L., Schiavone, A., Trocino, A., Xiccato, G., Barroeta, A. C., Maione, S., Soglia, D., Biasato, I., Cocolin, L., Gai, F., & Nucera, D. M. (2020). Antimicrobial Effects of Black Soldier Fly and Yellow Mealworm Fats and Their Impact on Gut Microbiota of Growing Rabbits. Animals, 10(8), 1292. https://doi.org/10.3390/ani10081292
• Elhag, O., Zhang, Y., Xiao, X., Cai, M., Zheng, L., Jordan, H. R., ... & Zhang, J. (2022). Inhibition of zoonotic pathogens naturally found in pig manure by black soldier fly larvae and their intestine bacteria. Insects, 13(1), 66.
• Elhag, O., Zhou, D., Song, Q., Soomro, A. A., Cai, M., Zheng, L., ... & Zhang, J. (2017). Screening, expression, purification and functional characterization of novel antimicrobial peptide genes from Hermetia illucens (L.). PloS One, 12(1), e0169582.
• Fahmy, L., Ali, Y. M., Seilly, D., McCoy, R., Owens, R. M., Pipan, M., ... & Grant, A. J. (2023). An attacin antimicrobial peptide, Hill_BB_C10074, from Hermetia illucens with anti-Pseudomonas aeruginosa activity. BMC Microbiology, 23(1), 378.
• Fahmy, L., Generalovic, T., Ali, Y. M., Seilly, D., Sivanesan, K., Kalmar, L., ... & Grant, A. J. (2024). A novel family of defensin-like peptides from Hermetia illucens with antibacterial properties. BMC Microbiology, 24(1), 167. https://doi.org/10.1186/s12866-024-03325-1
• Fatima, S., Afzal, A., Rashid, H., Iqbal, S., Zafar, R., Khalid, K., Rauf, A., Majeed, M., Malik, A., & Carter, C. G. (2023). Dietary replacement of soybean meal with black soldier fly larvae meal in juvenile Labeo rohita and Catla catla: Effects on growth, nutritional quality, oxidative stress biomarkers and disease resistance. PloS One, 18(11), e0294452. https://doi.org/10.1371/journal.pone.0294452
• Feng, M., Fei, S., Xia, J., Labropoulou, V., Swevers, L., & Sun, J. (2020). Antimicrobial peptides as potential antiviral factors in insect antiviral immune response. Frontiers in Immunology, 11, 2030.
• Freel, T. A., McComb, A., & Koutsos, E. A. (2021). Digestibility and safety of dry black soldier fly larvae meal and black soldier fly larvae oil in dogs. Journal of Animal Science, 99(3), skab047. https://doi.org/10.1093/jas/skab047
• Gadzama, I. U. (2024). Influence of rearing substrate on black soldier fly larvae nutritional value for animal feed. Retrieved from https://wikifarmer.com/influence-of-rearing-substrate-on-
• Gadzama, I. U., Malcolm, J., & Malcolm, C. (2024). Mineral composition in black soldier fly larvae: A promising alternative to fishmeal and soybean meal in livestock feed. In Proceedings of the 35th Biennial Conference of the Australian Association of Animal Sciences and the 20th Asian-Australasian Association of Animal Production Societies (Vol. 35, pp. 334-335). Melbourne: Australian Association of Animal Sciences. Available from: https://www.researchgate.net/publication/383122248
• Haidari, H., Melguizo-Rodríguez, L., Cowin, A. J., & Kopecki, Z. (2023). Therapeutic potential of antimicrobial peptides for treatment of wound infection. American Journal of Physiology-Cell Physiology, 324(1), C29-C38.
• Huan, Y., Kong, Q., Mou, H., & Yi, H. (2020). Antimicrobial peptides: classification, design, application and research progress in multiple fields. Frontiers in Microbiology, 11, 582779.
• Kar, S. K., Schokker, D., Harms, A. C., Kruijt, L., Smits, M. A., & Jansman, A. J. M. (2021). Local intestinal microbiota response and systemic effects of feeding black soldier fly larvae to replace soybean meal in growing pigs. Scientific Reports, 11(1), 15088. https://doi.org/10.1038/s41598-021-94604-8
• Kierończyk, B., Rawski, M., Mikołajczak, Z., Szymkowiak, P., Stuper-Szablewska, K., & Józefiak, D. (2023). Black soldier fly larva fat in broiler chicken diets affects breast meat quality. Animals, 13(7), 1137. https://doi.org/10.3390/ani13071137
• Koehbach, J., & Craik, D. J. (2019). The vast structural diversity of antimicrobial peptides. Trends in pharmacological sciences, 40(7), 517-528.
• Lacerda, A. F., Pelegrini, P. B., de Oliveira, D. M., Vasconcelos, É. A., & Grossi-de-Sá, M. F. (2016). Anti-parasitic peptides from arthropods and their application in drug therapy. Frontiers in Microbiology, 7, 91.
• Lalander, C. H., Fidjeland, J., Diener, S., Eriksson, S., & Vinnerås, B. (2015). High waste-to-biomass conversion and efficient Salmonella spp. reduction using black soldier fly for waste recycling. Agronomy for Sustainable Development, 35, 261-271.
• Lei, X. J., Kim, T. H., Park, J. H., & Kim, I. H. (2019). Evaluation of Supplementation of Defatted Black Soldier Fly Larvae Meal in Beagle Dogs. Annals of Animal Science, 19(3), 767-777. https://doi.org/10.2478/aoas-2019-0021
• Leni, G., Tedeschi, T., Faccini, A., Pratesi, F., Folli, C., Puxeddu, I., ... & Sforza, S. (2020). Shotgun proteomics, in-silico evaluation and immunoblotting assays for allergenicity assessment of lesser mealworm, black soldier fly and their protein hydrolysates. Scientific Reports, 10(1), 1228.
• Li, Z., Mao, R., Teng, D., Hao, Y., Chen, H., Wang, X., ... & Wang, J. (2017). Antibacterial and immunomodulatory activities of insect defensins-DLP2 and DLP4 against multidrug-resistant Staphylococcus aureus. Scientific Reports, 7(1), 12124.
• Liu, C., Yao, H., & Wang, C. (2021). Black soldier fly larvae can effectively degrade oxytetracycline bacterial residue by means of the gut bacterial community. Frontiers in Microbiology, 12, 663972.
• Llor, C., & Bjerrum, L. (2014). Antimicrobial resistance: risk associated with antibiotic overuse and initiatives to reduce the problem. Therapeutic Advances in Drug Safety, 5(6), 229-241.
• Luo, Y., & Song, Y. (2021). Mechanism of antimicrobial peptides: antimicrobial, anti-inflammatory and antibiofilm activities. International Journal of Molecular Sciences, 22(21), 11401.
• Marusich, E., Mohamed, H., Afanasev, Y., & Leonov, S. (2020). Fatty acids from Hermetia illucens larvae fat inhibit the proliferation and growth of actual phytopathogens. Microorganisms, 8(9), 1423.
• Miao, F., Li, Y., Tai, Z., Zhang, Y., Gao, Y., Hu, M., & Zhu, Q. (2021). Antimicrobial peptides: the promising therapeutics for cutaneous wound healing. Macromolecular Bioscience, 21(10), 2100103.
• Moretta, A., Salvia, R., Scieuzo, C., Di Somma, A., Vogel, H., Pucci, P., ... & Falabella, P. (2020). A bioinformatic study of antimicrobial peptides identified in the Black Soldier Fly (BSF) Hermetia illucens (Diptera: Stratiomyidae). Scientific Reports, 10(1), 16875.
• Moretta, A., Scieuzo, C., Petrone, A. M., Salvia, R., Manniello, M. D., Franco, A., ... & Falabella, P. (2021). Antimicrobial peptides: A new hope in biomedical and pharmaceutical fields. Frontiers in Cellular and Infection Microbiology, 11, 668632.
• Munthali, M. G., Chilora, L., Goliath, M., Burke, W. J., Benbow, E., Kang'ombe, J., & Safalaoh, A. (2023). The Economic Cost-benefit Analysis of Black Soldier Fly as an Alternative Animal and Fish Feed Ingredient in Malawi. https://ageconsearch.umn.edu/record/338590/files/WP-23-01%20BSF%20%28fv%29.pdf
• Muslykhah, U., Phupaboon, S., Suriyapha, C., Matra, M., & Wanapat, M. (2024). Encapsulation of protein-based bioactive from black soldier fly for ruminant feeding. Journal of Agriculture and Food Research, 101325.
• Patrulea, V., Borchard, G., & Jordan, O. (2020). An update on antimicrobial peptides (AMPs) and their delivery strategies for wound infections. Pharmaceutics, 12(9), 840.
• Pfalzgraff, A., Brandenburg, K., & Weindl, G. (2018). Antimicrobial peptides and their therapeutic potential for bacterial skin infections and wounds. Frontiers in Pharmacology, 9, 281.
• Raman, S. S., Stringer, L. C., Bruce, N. C., & Chong, C. S. (2022). Opportunities, challenges and solutions for black soldier fly larvae-based animal feed production. Journal of Cleaner Production, 373, 133802.
• Sepriadi, S., Hanafi, N. D., & Sadeli, A. (2022). The effect of milk replacer containing black soldier fly on pre-weaned Ettawa crossbreed goats on performance and physiological response. In IOP Conference Series: Earth and Environmental Science, 977 (1), 5. IOP Publishing. https://iopscience.iop.org/article/10.1088/1755-1315/977/1/012132
• Shu, M. K., Li, C. M., Furtado, W. E., Huang, Q., St-Hilaire, S., & Kenéz, Á. (2024). Antibacterial properties of oil extracts of black soldier fly larvae reared on bread waste. Animal Production Science, 64(8).
• Simjee, S., White, D. G., Meng, J., Wagner, D. D., Qaiyumi, S., Zhao, S., ... & McDermott, P. F. (2002). Prevalence of streptogramin resistance genes among Enterococcus isolates recovered from retail meats in the Greater Washington DC area. Journal of Antimicrobial Chemotherapy, 50(6), 877-882.
• Singh, A., & Kumari, K. (2019). An inclusive approach for organic waste treatment and valorisation using Black Soldier Fly larvae: A review. Journal of Environmental Management, 251, 109569. 
• Smets, R., Goos, P., Claes, J., & Van Der Borght, M. (2021). Optimisation of the lipid extraction of fresh black soldier fly larvae (Hermetia illucens) with 2-methyl tetrahydrofuran by response surface methodology. Separation and Purification Technology, 258, 118040.
• Spranghers, T., Michiels, J., Vrancx, J., Ovyn, A., Eeckhout, M., De Clercq, P., & De Smet, S. (2018). Gut antimicrobial effects and nutritional value of black soldier fly (Hermetia illucens L.) prepupae for weaned piglets. Animal Feed Science and Technology, 235, 33-42. https://doi.org/10.1016/j.anifeedsci.2017.08.012
• Starr, C. G., & Wimley, W. C. (2017). Antimicrobial peptides are degraded by the cytosolic proteases of human erythrocytes. Biochimica et biophysica acta. Biomembranes, 1859(12), 2319–2326. https://doi.org/10.1016/j.bbamem.2017.09.008
• Sypniewski, J., Kierończyk, B., Benzertiha, A., Mikołajczak, Z., Pruszyńska-Oszmałek, E., Kołodziejski, P., Sassek, M., Rawski, M., Czekała, W., & Józefiak, D. (2020). Replacement of soybean oil by Hermetia illucens fat in turkey nutrition: effect on performance, digestibility, microbial community, immune and physiological status and final product quality. British Poultry Science, 61(3), 294–302. https://doi.org/10.1080/00071668.2020.1716302
• Van Huis, A., & Oonincx, D. G. A. B. (2017). The environmental sustainability of insects as food and feed. A review. Agronomy for Sustainable Development, 37(5), Article 43. https://doi.org/10.1007/s13593-017-0452-8
• Veldkamp, T., Dong, L., Paul, A., & Govers, C. C. F. M. (2022). Bioactive properties of insect products for monogastric animals–a review. Journal of Insects as Food and Feed, 8(9), 1027-1040. https://doi.org/10.3920/JIFF2021.0031
• Vogel, H., Müller, A., Heckel, D. G., Gutzeit, H., & Vilcinskas, A. (2018). Nutritional immunology: diversification and diet-dependent expression of antimicrobial peptides in the black soldier fly Hermetia illucens. Developmental & Comparative Immunology, 78, 141-148.
• Wu, N., Wang, X., Xu, X., Cai, R., & Xie, S. (2020). Effects of heavy metals on the bioaccumulation, excretion and gut microbiome of black soldier fly larvae (Hermetia illucens). Ecotoxicology and Environmental Safety, 192, 110323.
• Wu, Q., Patočka, J., & Kuča, K. (2018). Insect antimicrobial peptides, a mini review. Toxins, 10(11), 461.
• Xia, J., Ge, C., & Yao, H. (2021). Antimicrobial Peptides from Black Soldier Fly (Hermetia illucens) as Potential Antimicrobial Factors Representing an Alternative to Antibiotics in Livestock Farming. Animals, 11(7), 1937. https://doi.org/10.3390/ani11071937
• Xu, J., Luo, X., Fang, G., Zhan, S., Wu, J., Wang, D., & Huang, Y. (2020). Transgenic expression of antimicrobial peptides from black soldier fly enhance resistance against entomopathogenic bacteria in the silkworm, Bombyx mori. Insect Biochemistry and Molecular Biology, 127, 103487.
• Yan, Y., Zhang, J., Chen, X., & Wang, Z. (2023). Effects of Black Soldier Fly Larvae (Hermetia illucens Larvae) Meal on the Production Performance and Cecal Microbiota of Hens. Veterinary sciences, 10(5), 364. https://doi.org/10.3390/vetsci10050364
• Yi, H. Y., Chowdhury, M., Huang, Y. D., & Yu, X. Q. (2014). Insect antimicrobial peptides and their applications. Applied Microbiology and Biotechnology, 98, 5807-5822.
• Zabulionė, A., Šalaševičienė, A., Makštutienė, N., & Šarkinas, A. (2023). Exploring the antimicrobial potential and stability of black soldier fly (Hermetia illucens) larvae fat for enhanced food shelf-life. Gels, 9(10), 793. https://doi.org/10.3390/gels9100793
• Zhan, S., Fang, G., Cai, M., Kou, Z., Xu, J., Cao, Y., ... & Huang, Y. (2020). Genomic landscape and genetic manipulation of the black soldier fly Hermetia illucens, a natural waste recycler. Cell Research, 30(1), 50-60.