Biotechnological interventions in sericulture: A Review of current trends and future scope

Introduction

One of the oldest agro-based industries in Asia especially in China, India, Japan and Thailand is sericulture which involves raising domesticated silkworms (Bombyx mori L.) for the production of silk (Shahzadi, Tahir, Ali, Bhatti, Azizullah, Khan, & Khaliq, 2022) [33] . Because sericulture is labor-intensive but low-investment, it creates jobs for women and smallholder farmers, which contributes significantly to rural development beyond textiles (Dasaraj, 2021) [6] . Selective breeding of mulberry cultivars and silkworm strains has historically been used to increase the output and quality of silk. However, advanced technological interventions are required due to the growing demand for premium silk, climate unpredictability and silkworm infections (Pan, Deng, Chen, Cai, Duan, & Xia, 2020) [25] . In sericulture, biotechnology which includes tissue culture, molecular markers, transgenics, genome editing and microbial engineering has become a game-changer. Biotechnological techniques including genetic transformation and micropropagation have sped up the production of high-yielding, stress-tolerant mulberry cultivars (Shahzadi et al., 2022) [33] . New approaches to improving silk fiber properties, strengthening resistance to viral pathogens and deciphering intricate physiological processes have been made possible by developments in functional genomics, transcriptomics and genome editing technologies like CRISPR/Cas9 in silkworms (Zhang, Xia, Day, Harris, Oliveira, Knittel, Yarger & Lewis, 2019; Sun, Chen, Wang, Liu, Zhang, & Li, 2024) [47,36] . Additionally, it has been shown that CRISPR/Cas12a-based editing can give resistance to the Bombyx mori nucleopolyhedrovirus (BmNPV) underscoring the usefulness of genome engineering for disease control (Pan et al., 2020) [25] . While lowering reliance on chemical interventions, omics-driven methodologies and RNA interference (RNAi) tactics also provide innovative options for controlling pests and pathogens (Gao, Li, Tong, Liu, Xu, Zhang, & Lin, 2020) [10] . Meanwhile, biotechnological procedures allow for the sustained value-adding of sericulture by-products including sericin and pupal protein, which are becoming more and more recognized for their biomedical, cosmetic and nutraceutical uses (Aramwit, 2017) [1] . Green solvents and enzymatic degumming are two new environmentally friendly methods that support sustainable silk manufacturing and the circular bioeconomy. Furthermore, biotechnology is being integrated with sericulture practices through digital innovations like artificial intelligence, Internet of Things (IoT)-based monitoring and biosensors made from silk fibroin (Wang, Zhu, Yu, Li, Han, & Tao, 2021) [41] . This ensures resilience against market and environmental challenges.

Biotechnology in Mulberry Improvement

Mulberry (Morus spp.) the main food source for silkworms (Bombyx mori L.) is essential to sericulture since cocoon yield and silk quality are directly impacted by the productivity and quality of the leaves. Although conventional breeding has yielded a number of better varieties, biotechnological treatments are required for rapid development because to the lengthy breeding cycle, limited genetic base and susceptibility to climate stress (Sarkar, Mogili, & Sivaprasad, 2017; Taha, Ghazy, & Gabr, 2020) [38] .

Tissue Culture and Micropropagation

Tissue culture methods, including as somatic embryogenesis, nodal culture and shoot tip culture are frequently used to produce homogeneous, disease-free and true-to-type planting material. Elite cultivars can multiply quickly through micropropagation while retaining genetic stability. Furthermore, in vitro culture-induced somaclonal variation has been used to produce novel variability especially for traits like stress tolerance (Bhattacharya & Vijayan, 2018) [4] .

Molecular Markers and Marker-Assisted Selection (MAS)

Molecular markers such as RAPD, AFLP, SSRs and SNPs have been applied to characterize mulberry germplasm, assess genetic diversity and identify quantitative trait loci (QTLs) associated with leaf yield, protein content and stress tolerance. Marker-assisted selection (MAS) has enhanced the precision of breeding programs allowing rapid introgression of desirable alleles and the development of varieties with improved adaptability and disease resistance (Das, Singh, & Singh, 2016) [5] .

Genetic Transformation and Genome Editing

Genes that confer abiotic stress tolerance (salinity, drought) and enhanced photosynthetic efficiency have been introduced by transgenic techniques, such as Agrobacterium-mediated transformation (Wani, Bhat, Kamilli, Malik, Mir, Iqbal, Mir, Hassan, & Hamid, 2013) [43] . Although field-level applications are still mostly in the experimental stage recent developments in CRISPR/Cas9 genome editing offer the possibility of targeted modification of important genes regulating lignin biosynthesis, leaf nutritional quality and stress resilience (Zhang, Xia, Day, Harris, Oliveira, Knittel, Yarger, & Lewis, 2019) [47] .

Somaclonal variation

In vitro culture-induced variability Stress tolerance, leaf yield Generation of novel genotypes with enhanced adaptability Taha, Ghazy, & Gabr, 2020 [38] Molecular markers & MAS RAPD, SSRs, SNPs, QTL mapping Drought tolerance, leaf quality, disease resistance Identification of marker-trait associations; precision breeding Das, Singh, & Singh, 2016 [5] Genetic transformation Agrobacteriummediated gene transfer Stress tolerance, photosynthetic efficiency Transgenic mulberry with improved abiotic stress tolerance Wani, Bhat, Kamilli, Malik, Mir, Iqbal, Mir, Hassan, & Hamid, 2013 [43] Genome editing CRISPR/Cas9, TALENs Lignin biosynthesis, leaf nutrient quality

Targeted modification of genes; potential for designer varieties Zhang, Xia, Day, Harris, Oliveira, Knittel, Yarger, & Lewis, 2019 [47] Silkworm Genetics and Functional Genomics Because of its commercial significance and well-sequenced genome, the domesticated silkworm (Bombyx mori L.) has become a model lepidopteran insect for genetics and biotechnology. Functional genomics research in sericulture is now possible thanks to developments in next-generation sequencing (NGS) which has produced high-resolution genome assemblies and transcriptome datasets. A strong basis for trait discovery, especially in silk protein synthesis, disease resistance and growth regulation has been established by high throughput sequencing of the B. mori genome, which has identified over 14,600 protein-coding genes in addition to non-coding RNAs, repetitive elements and structural variants (Kawamoto, Jouraku, Toyoda, Yokoi, Minakuchi, Katsuma, Shimada, & Fujiwara, 2019) [14] .

Research on silkworms has been transformed by CRISPR/Cas gene editing, which enables targeted editing of immunity related genes to confer pathogen resistance, precise modification of silk fibroin genes to improve mechanical properties and manipulation of pigmentation genes for designer silk (Zhang, Zhang, Liu, Li, & Xia, 2021) [19] . RNA interference (RNAi) has been used in parallel studies to functionally shut down pathogenic targets showing promise for managing viral infections such the BmNPV (Bombyx mori nucleopolyhedrovirus) (Xu, Li, Zhang, Chen, & Wu, 2020) [25] . [19] P25 (Fibrohexamerin) CRISPR/Cas9 Improved silk stability and secretion Li, Chen, Zhang, Wang, & Xia, 2022 [18] Yellow-e, Yellow-d CRISPR/Cas9 Altered silk pigmentation (golden, reddish silk) Dong, Huang, Zhang, & Xia, 2019 [8] BmNPV Immediate Early Genes RNAi Resistance to viral infection Xu, Li, Zhang, Chen, & Wu, 2020 [42] Antimicrobial Peptide Genes (Cecropin, Attacin) CRISPR activation Enhanced immunity against bacterial pathogens Liu, Zhang, Li, & Xia, 2021 [12] Disease and Pest Management through Biotechnology Silkworm illnesses (pebrine, grasserie, flacherie and muscardine) and pests pose serious problems for sericulture resulting in significant production and financial losses. Efficiency, sustainability and environmental safety are all limited by traditional management techniques including chemical treatments and hygienic measures. Recent developments in biotechnology have made it possible to implement accurate and environmentally responsible treatments for disease resistance, pest management and diagnostics.

Molecular Diagnostics

For the quick and accurate identification of silkworm infections, such as Nosema bombycis (pebrine), Bombyx mori nucleopolyhedrovirus (BmNPV; grasserie) and bacterial agents that cause flacherie, PCR, qPCR and loopmediated isothermal amplification (LAMP) assays are frequently utilized. By enabling early diagnosis prior to the onset of obvious symptoms, these methods help stop disease epidemics (Selvakumar, Vijayalakshmi, & Arumugam, 2019) [32] .

Probiotics and Microbiome Engineering

Manipulating the silkworm gut microbiome through probiotics (e.g., Lactobacillus, Bacillus spp.) enhances immunity, digestion and disease tolerance. Microbiome based approaches are emerging as sustainable solutions for silkworm health management (Guo, Chen, Wu, Wang, & Li, 2021) [12] .

RNA Interference (RNAi)

A promising method for managing silkworm viral infections, especially BmNPV is the use of RNA interference (RNAi) and dsRNA sprays that target viral genes. RNA interference (RNAi) lowers viral mortality and reproduction by squelching essential viral transcripts (Nandi, Kar, Ghosh, & Mandal, 2020) [24] .

Biopesticides and Pheromone-Based Integrated Pest Management (IPM)

The treatment of silkworm pests is increasingly incorporating natural biocontrol agents such as pheromone traps and entomopathogenic fungi like Beauveria bassiana and Metarhizium anisopliae. These methods reduce the need of chemical pesticides and offer mulberry and silkworm crops environmentally friendly protection (Suresh, Rao, & Reddy, 2018) [37] . [32] LAMP assays Pebrine, flacherie bacteria Field-level diagnostics Wang, Li, Chen, & Zhang, 2020 [25] Probiotics (Lactobacillus, Bacillus)

Gut pathogens, bacterial infections Enhanced immunity and digestion Guo, Chen, Wu, Wang, & Li, 2021 [12] RNAi/dsRNA sprays BmNPV Gene silencing, reduced viral load Nandi, Kar, Ghosh, & Mandal, 2020 [24] Biopesticides (Beauveria bassiana, Metarhizium anisopliae)

Silkworm pests

Eco-friendly pest suppression Suresh, Rao, & Reddy, 2018 [37] Pheromone traps Uzi fly (Exorista bombycis) Monitoring and population control Ramesh, Kumar, & Singh, 2019 [28] Bioprocessing and Green Technologies The chemical-intensive processes used in traditional silk reeling and processing procedures produce a lot of effluent and break down important silk proteins like sericin. The goals of recent developments in green technology and bioprocessing techniques are to enhance silk quality, reduce environmental effects, and recover bioactive chemicals for a variety of uses.

Enzymatic Degumming

Enzymatic degumming with proteases, lipases, and pectinases offers a gentle and targeted removal of sericin from silk fibroin fibers in contrast to alkaline or soap-based chemical degumming. This lowers the chemical load in effluents while maintaining the mechanical strength and molecular integrity of fibroin (Aramwit, Srichana, Srichana, & Sangcakul, 2013) [3] .

Ionic Liquids (ILs) and Deep Eutectic Solvents (DESs)

These novel solvents enable eco-friendly fibroin dissolution and reconstitution avoiding harsh chemicals. ILs such as 1butyl-3-methylimidazolium chloride ([Bmim]Cl) and DESs composed of choline chloride with urea or glycerol offer recyclable, tunable systems for silk processing (Zhang, Li, Wang, & Xia, 2021) [12] .

Sericin Recovery and Valorization

Sericin, which was formerly regarded as a waste product is now valued as a high-value biomaterial that finds use in nutraceuticals, biomedicine and cosmetics. Membrane filtration, spray drying and freeze-drying are recovery methods that guarantee the functional preservation of moisturizing and antioxidant qualities (Dash, Sahoo, & Nayak, 2020) [7] . Together, these developments lessen the environmental impact and maximize the use of silk byproducts in sericulture, thereby promoting a circular bioeconomy. [16, 30] . Additionally, functionalized silk nanoparticles are being investigated for cancer treatments and gene delivery.

Cosmetic Applications

Because of its moisturizing, UV-protective and anti-aging properties sericin is being utilized more and more in skincare products including creams, lotions and masks. It is a valued element in international cosmetic markets because to its capacity to create a thin protective layer on the skin (Aramwit, Kanokpanont, & Kanokpanont, 2017) [1] .

Industrial Applications

Packaging, coatings and sensor technologies use biodegradable silk fibroin films, polymer blends and composites. Hydrogels and nanoparticles made from sericin have applications in nanotechnology and environmentally friendly sectors (Vepari & Kaplan, 2007) [40] . In addition to improving sericulture's financial sustainability, the conversion of silk proteins into high-value goods advances the creation of a circular bioeconomy. [29] .

Artificial intelligence (AI) applications

To identify early signs of silkworm diseases (such muscardine and grasserie) using digital photos, machine learning models are being trained. AI also makes it possible to estimate cocoon yield and predict the stages of larval growth, which lowers human error and improves decisionmaking (Patil et al., 2021) [26] .

Biosensors for pathogen detection

Rearing rooms equipped with biosensors are able to quickly identify pathogenic agents (bacterial infections, BmNPV and Nosema bombycis). They offer on-site diagnostics in conjunction with smartphone-based readouts enabling prompt actions (Kumari et al., 2019) [15] . All of these developments point to a move toward precision sericulture, which lowers risks, boosts productivity and promotes sustainable silk production.

Sustainability and Circular Bioeconomy

Sericulture is not only a source of silk but also generates a wide range of by-products such as pupal protein, frass (silkworm excreta) and reeling waste, which were once considered low-value residues. Recent advances in sustainable practices have transformed these by-products into valuable resources, contributing to a circular bioeconomy.

Sericulture By-Products  Silkworm pupae: Rich in protein, essential amino acids and polyunsaturated fatty acids, pupae are utilized in animal feed, aquaculture and nutraceutical industries  Silkworm frass: A nitrogen-and phosphorus-rich organic material, frass serves as an effective biofertilizer, improving soil fertility and plant growth  Reeling waste: Pierced cocoons, floss and waste silk fibers are repurposed into spun silk yarn, nonwoven textiles and sericin extraction (Aramwit, Srichana, Srichana, & Sangcakul, 2013) [3] .

Circular Economy Practices

By converting waste into animal feed, fertilizers, cosmetics and biomedical materials sericulture enhances resource efficiency and minimizes waste. For instance, sericin recovered from wastewater is used in cosmetics and wound healing formulations, creating additional income streams and reducing pollution load.

Life Cycle Assessment (LCA)

LCA studies of the silk industry highlight the environmental impacts of cocoon production, degumming and dyeing. Adoption of green technologies such as enzymatic degumming, renewable energy in reeling and eco-friendly dyes can significantly lower the carbon footprint and enhance sustainability. Thus, sericulture represents a model bio-based industry where every stage of production can be integrated into a zero-waste circular system

Adoption, Policy and Ethics

Although promising, there are a number of obstacles to the widespread use of biotechnology in sericulture. Smallholder farmers, who control the majority of the sericulture industry are unable to use biotechnological techniques due to their high initial costs (e.g., molecular markers, CRISPR-based breeding, IoT monitoring). Integration of these innovations into conventional operations is further hampered by a lack of technical training and awareness (Lakshmi & Kumar, 2021) [17] . Additionally, intellectual property (IP) issues around patented biotech tools restrict open access and create inequity in technology dissemination.

When it comes to silkworm biotechnology, biosafety and ethical are crucial. Genetic modifications such as transgenic silkworms for enhanced silk quality, disease resistance or recombinant protein production raise concerns about ecological risks (e.g., accidental release, biodiversity impact) and ethical debates over animal manipulation. To strike a balance between innovation and ecological responsibility, strict biosafety procedures and open regulatory frameworks are required (Nagaraju & Goldsmith, 2002) [23] . Public-private partnerships (PPPs) are becoming more and more crucial in bridging the divide between industry, government and research organizations. PPPs can ensure adherence to ethical standards while facilitating the commercialization of innovations, farmer training and the provision of reasonably priced biotech toolkits. Scaling acceptance is greatly aided by government involvement through initiatives like capacity-building programs, subsidies, and biotech-friendly laws. Lakshmi & Kumar, 2021 [17] Farmer training & awareness programs Knowledge dissemination Enhances understanding of biotech benefits and safe practices [23] Public-private partnerships (PPPs)

Collaborative innovation & commercialization

Accelerates lab-to-field translation and market readiness Sharma & Raina, 2020

Future Directions and Research Gaps

The application of biotechnology in sericulture has opened new avenues for crop and silkworm improvement, disease management, and value addition. However, several research gaps and future directions remain to be addressed for realizing the full potential of these technologies.

a) Multi-omics integration (genomics, proteomics, metabolomics

Although the Bombyx mori genome has been sequenced and mulberry genomics is advancing, integrating multi-omics platforms remains limited. A systems biology approach combining genomics, transcriptomics, proteomics, and metabolomics can provide deeper insights into complex traits such as silk protein synthesis, stress tolerance, and host-pathogen interactions (Mita et al., 2004; Goldsmith et al., 2005) [4,11] . Future research should focus on building comprehensive omics databases for mulberry and silkworm to accelerate molecular breeding and functional validation. [39] . Similarly, developing silkworm strains with resilience against temperature fluctuations and disease outbreaks through MAS, genome editing, and hybridization will ensure stable silk production under changing environmental conditions.

c) Field validation of CRISPR and RNAi technologies.

While laboratory studies have demonstrated the potential of CRISPR/Cas and RNAi in enhancing silk yield, modifying fibroin/sericin genes, and improving disease resistance, field-scale validation is still lacking (Meng et al., 2017) [21] . Translating these gene-editing technologies into practical sericulture requires optimization of delivery systems, biosafety assessments, and long-term monitoring of ecological impacts.

D) Green processing scale-up and regulatory frameworks.

Eco-friendly silk processing techniques such as enzymatic degumming, ionic liquids, and sericin valorization have been demonstrated at the laboratory scale (Zhang et al., 2020) [42] . However, their industrial scalability, costeffectiveness, and environmental trade-offs require further research. Regulatory frameworks must also evolve to evaluate and promote green bioprocessing technologies within the silk sector, ensuring alignment with global sustainability goals. [21] Green silk processing Pilot studies with enzymes & ionic liquids

Industrial scale-up challenges

Develop cost-effective ecofriendly protocols Zhang et al.,

Conclusion

Biotechnology is increasingly emerging as a transformative force in sericulture offering innovations across the value chain from mulberry improvement and silkworm genomics to disease management, green bioprocessing, and silk biomaterial applications. Tools such as CRISPR/Cas, RNAi, molecular markers and omics platforms have demonstrated significant potential in enhancing productivity, resilience and value addition. However, the successful future of sericulture biotechnology depends on its balanced integration with digital technologies (e.g., IoT, AI, biosensors) and supportive policy frameworks that address barriers like cost, awareness, biosafety and intellectual property rights. Public private partnerships and farmer training programs will be critical for ensuring that innovations translate effectively from laboratories to rearing houses and markets. Future research must prioritize scalability, sustainability and farmer-centric adoption. Developing climate-resilient mulberry varieties and silkworm breeds, validating geneediting and RNAi tools at field scale and promoting ecofriendly processing technologies will be essential to strengthen the resilience of the silk sector in the face of climate change and global sustainability demands. In essence, the convergence of biotechnology, digital sericulture and circular bioeconomy practices can position sericulture as not only a traditional livelihood source but also a future-ready bio-based industry contributing to rural development, environmental sustainability and global silk markets.

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