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<journal-id journal-id-type="publisher-id">Front. Bacteriol.</journal-id>
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<journal-title>Frontiers in Bacteriology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Bacteriol.</abbrev-journal-title>
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<issn pub-type="epub">2813-6144</issn>
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<article-id pub-id-type="doi">10.3389/fbrio.2026.1746114</article-id>
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<subj-group subj-group-type="heading">
<subject>Review</subject>
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<title-group>
<article-title>Bacterial pigments as sustainable and functional colorants in cosmetic applications: advances and prospects</article-title>
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<name><surname>Chakraborty</surname><given-names>Pritha</given-names></name>
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<name><surname>Kaur</surname><given-names>Mandheer</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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<name><surname>Thazeem</surname><given-names>Basheer</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<name><surname>Thomas</surname><given-names>Jithin</given-names></name>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
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<name><surname>S</surname><given-names>Hemavathy</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
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<name><surname>Sharma</surname><given-names>Deepak</given-names></name>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>*</sup></xref>
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<aff id="aff1"><label>1</label><institution>School of Allied Healthcare and Sciences, JAIN (Deemed to be University)</institution>, <city>Bangalore</city>, <state>Karnataka</state>,&#xa0;<country country="in">India</country></aff>
<aff id="aff2"><label>2</label><institution>Department of Biotechnology, Chandigarh College of Technology, Chandigarh Group of Colleges Landran</institution>, <city>Mohali</city>, <state>Punjab</state>,&#xa0;<country country="in">India</country></aff>
<aff id="aff3"><label>3</label><institution>Postgraduate and Research Department of Zoology, Maharaja&#x2019;s College</institution>, <city>Ernakulam</city>, <state>Kerala</state>,&#xa0;<country country="in">India</country></aff>
<aff id="aff4"><label>4</label><institution>Department of Biotechnology, Mar Athanasius College</institution>, <city>Kothamangalam</city>, <state>Kerala</state>,&#xa0;<country country="in">India</country></aff>
<aff id="aff5"><label>5</label><institution>Biomaterials and Tissue Engineering Laboratory (BITE LAB), Department of Community Medicine, Saveetha Medical College and Hospital, Saveetha Institute of Medical and Technical Sciences (SIMATS)</institution>, <city>Chennai</city>, <state>Tamil Nadu</state>,&#xa0;<country country="in">India</country></aff>
<aff id="aff6"><label>6</label><institution>Advanced Therapeutics and Sensing Laboratory, Department of General Surgery, Saveetha Medical College and Hospital, Saveetha Institute of Medical and Technical Sciences (SIMATS)</institution>, <city>Chennai</city>, <state>Tamil Nadu</state>,&#xa0;<country country="in">India</country></aff>
<author-notes>
<corresp id="c001"><label>*</label>Correspondence: Deepak Sharma, <email xlink:href="mailto:deepaksharma2899@gmail.com">deepaksharma2899@gmail.com</email>; <email xlink:href="mailto:deepaksharma.smc@saveetha.com">deepaksharma.smc@saveetha.com</email></corresp>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2026-02-11">
<day>11</day>
<month>02</month>
<year>2026</year>
</pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2026</year>
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<volume>5</volume>
<elocation-id>1746114</elocation-id>
<history>
<date date-type="received">
<day>14</day>
<month>11</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>13</day>
<month>01</month>
<year>2026</year>
</date>
<date date-type="rev-recd">
<day>04</day>
<month>01</month>
<year>2026</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2026 Chakraborty, Kaur, Thazeem, Thomas, S and Sharma.</copyright-statement>
<copyright-year>2026</copyright-year>
<copyright-holder>Chakraborty, Kaur, Thazeem, Thomas, S and Sharma</copyright-holder>
<license>
<ali:license_ref start_date="2026-02-11">https://creativecommons.org/licenses/by/4.0/</ali:license_ref>
<license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License (CC BY)</ext-link>. The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</license-p>
</license>
</permissions>
<abstract>
<p>The bacterial world is a promising source for the production of colored metabolites known as bacterial pigments. Synthetic dyes are responsible for health problems such as allergies, cancer, toxicity, and hyperactivity and for environmental issues such as pollution of waterways, disruption in aquatic ecosystems, inhibition of photosynthesis, and bioaccumulation in food chains. These concerns have provided the microbial world a chance of being used as a source of natural pigment for industrial applications, including cosmetics. Cosmetic products such as lipsticks, nail polishes, hair dyes, soaps, body washes, face washes, creams, and lotions utilize various colored compounds of chemical origin and may pose adverse effects on their unregulated or overuse. Bacterial pigments can be an alternate and sustainable option to replace these chemical moieties in these cosmetic products. Bacteria from diverse habitats with a broad scale of colors such as carotenoids, prodigiosin, melanin, violaceins, quinones, and indigoidines have been reported for their beneficial properties such as color, antioxidant, emulsifying, antiaging, and UV protection. These pigments have multiple shades and also possess nutritional and therapeutic properties. Although most of the information in this field is based on primary research at a laboratory scale and very limited attempts have been made to improve these bacterial strains and processes for enhanced pigment production, it highlights a significant scope for research and innovations in this field. The integration of advanced genetic and metabolic engineering technology such as CRISPR/Cas, along with the utilization of artificial intelligence and machine learning-based methods, can transform this field and boost pigment production. Therefore, the current review article aims to provide a state-of-the-art overview of bacterial pigments with the potential for application in cosmetic products. Moreover, it also highlights the existing challenges and outlines future research directions.</p>
</abstract>
<kwd-group>
<kwd>bacterial pigments</kwd>
<kwd>chemical toxicity</kwd>
<kwd>cosmetics</kwd>
<kwd>natural colors</kwd>
<kwd>sustainable source</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declared that financial support was not received for this work and/or its publication.</funding-statement>
</funding-group>
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<meta-name>section-at-acceptance</meta-name>
<meta-value>One Health in Bacteriology</meta-value>
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</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>The history of dyeing dates back to the Indus Valley civilization, approximately 2600&#x2013;1900 BC, showing the interest of humans in color. Before the 19th century, most of the dyes were natural and curated from fruits, plants, and berries using various techniques (<xref ref-type="bibr" rid="B500">Alegbe and Uthman, 2024</xref>). In 1850, the first synthetic dye was produced, and almost 80 different synthetic dyes were produced and used in different sectors by the 1900s (<xref ref-type="bibr" rid="B501">Rather et&#xa0;al., 2023</xref>). The great industrial revolution had quickly turned the attention toward synthetic dyes due to their cheaper cost and availability with limited understanding of their long-term effect. Although synthetic dyes are known to be carcinogenic and harmful, at present, they are widely used in food, textile, agricultural, and cosmetic products (<xref ref-type="bibr" rid="B15">Cheng et&#xa0;al., 2022</xref>). The presence of synthetic dyes in cosmetic products is a well-known fact. Human skin is a very sensitive organ that loses moisture and elasticity over time. Constant exposure to ultraviolet (UV) light and chemicals can result in faster aging of the skin. Thus, skin care is important to maintain its barrier function (<xref ref-type="bibr" rid="B502">Resende et&#xa0;al., 2021</xref>). The synthetic dyes present in cosmetic products have many harmful effects such as skin damage and tumor formation. Heavy metals such as chromium and cadmium are found in cosmetic products like lipsticks, eye shadow, eyeliner, and blush, which can lead to skin allergies, blindness, and even death (<xref ref-type="bibr" rid="B15">Cheng et&#xa0;al., 2022</xref>). These carcinogenic compounds are absorbed by the skin and act as neurotoxins, endocrine disruptors, and mutagens, affecting reproductive and overall human health. According to <xref ref-type="bibr" rid="B504">Wijesekara and Xu (2024)</xref>, several synthetic colorants approved by the Food and Drug Administration (FDA) were found to be carcinogenic. Furthermore, in addition to skin contact, humans are also exposed to synthetic dyes through contaminated water and the food chain. These dyes trigger allergies, asthma, dermatitis, central nervous system disorders, organ dysfunction, and cancer risks. The bioaccumulation of these dyes in fish fatty tissues poses zoonotic threats (<xref ref-type="bibr" rid="B89">Sudarshan et&#xa0;al., 2023</xref>). Environmental contamination from these dyes is caused by the discharge of wastewater from various industries, including the textile and cosmetic industries. In the aquatic system, these dyes exhibit carcinogenic and mutagenic properties along with high biological oxygen demand and chemical oxygen demand (<xref ref-type="bibr" rid="B88">Srivastava et&#xa0;al., 2022</xref>). These dyes also reduce light penetration in waterways, thus inhibiting microalgae photosynthesis, causing growth inhibition, ecosystem collapse, and toxicity to fish (<xref ref-type="bibr" rid="B42">Krishna Moorthy et&#xa0;al., 2021</xref>).</p>
<p>Natural pigments are usually not harmful but are more expensive than the synthetic ones. Lately, there is a growing concern among consumers of skin care products regarding the safety and increasing demand for natural pigments, which are not harmful to the skin (<xref ref-type="bibr" rid="B57">Mary et&#xa0;al., 2024</xref>). The advantage of natural pigments is their antioxidant and antimicrobial properties, which boost their demand in the market. The market demand for natural pigments has grown to nearly 2.5 billion USD in 2025 and is expected to increase to approximately 55 billion USD by 2027 (<xref ref-type="bibr" rid="B504">Anshi et&#xa0;al., 2024</xref>). The need for bio-based, less harmful, and high-performance colorants has driven the demand for natural pigments. The ever-expanding cosmetic industry requires numerous sources of natural pigments (<xref ref-type="bibr" rid="B5">Agarwal et&#xa0;al., 2023</xref>). The growing demand for natural pigments in Asia is mainly due to industrialization and increasing population, and the demand for natural pigments is also increasing equally in European and American markets. Carotenoids are one of the pigments most widely used across food, cosmetics, medicine, textiles, beverages, and dietary supplements. The global demand remains high at 23.5% in 2023, and 80%&#x2013;90% of the carotenoids present in the market are synthetic in nature (<xref ref-type="bibr" rid="B1">Abdelaziz et&#xa0;al., 2023</xref>). There are a few microbial-sourced pigments already on the market, such as riboflavin, alpha-carotene, and astaxanthin from <italic>Eremothecium gossypii</italic>, <italic>Bacillus trispora</italic>, and <italic>Xanthophyllomyces dendrorhous</italic>, respectively. The extraction of natural pigments from novel sources and their application have gained momentum among researchers at present (<xref ref-type="bibr" rid="B505">Kharkhota et&#xa0;al., 2022</xref>).</p>
<p>Pigments derived from microbes are identified as microbial pigments that bear health benefits for humans through essential nutrient supply, and their antimicrobial, anti-inflammatory, and anticancer activities are an added advantage (<xref ref-type="bibr" rid="B506">Di Salvo et&#xa0;al., 2023</xref>). Their low cost, durability, sustainability, availability, environment-friendly nature, and numerous sources and applications across various industries are some of the reasons for their high demand (<xref ref-type="bibr" rid="B507">Dufoss&#xe9;, 2019</xref>). Microbes produce a range of pigments to adapt to the ecosystem, exhibiting unique characteristics like protection against solar or UV radiation. Even though fungi, microalgae, and yeast produce pigments, bacteria-derived pigments attract the majority of the market due to their abundance and ready availability (<xref ref-type="bibr" rid="B508">Singh et&#xa0;al., 2021</xref>). Quinones, melanin, flavin, monascine, carotenoids, phenazines, and violacein are some of the examples of bacterial pigments with provitamin A and antitumor activity and temperature, pH, and photostability (<xref ref-type="bibr" rid="B501">Rather et&#xa0;al., 2023</xref>). <italic>Serretia mercescens</italic> produces the red-colored pigment prodigiosin with anticancer and antimicrobial activity in the presence of glucose and glycine. <italic>Dietzia natronolimnaea</italic> HS-1 produces the dark red-colored canthaxanthin used in the pharmaceutical, food, and cosmetic industries using cheese whey as a substrate, while <italic>Duganella</italic> sp., <italic>Janthinobacterium lividum</italic>, and <italic>Chromobacterium</italic> sp. produce the purple-colored violacein pigment that exhibits antitumoral, antioxidant, antiparasitic, and immunomodulatory activities. While red and yellow pigments are abundantly produced by several bacteria, only a few bacteria can produce blue pigments. <italic>Corynebacterium insidiosum</italic> produces indigoidine, which is blue in color and exhibits protection against oxidative stress. The blue-green color pigment pyocyanin, which helps in reducing inflammation, is produced by <italic>Pseudomonas aeruginosa</italic> (<xref ref-type="bibr" rid="B95">Venil et&#xa0;al., 2020</xref>).</p>
<p>The Green Revolution has hit the cosmetic industry, advocating for health and safety and introducing a term called cosmeceuticals. It offers a new perspective besides beauty and attracts vibrant business ideas. In addition to their antioxidant and antiaging effects, bacterial pigments offer vibrant colors in cosmetic products (<xref ref-type="bibr" rid="B509">Pagels et&#xa0;al., 2022</xref>). Sun damage to the skin leads to wrinkles, aging, and even skin cancer. Regular sunblock products available in the market absorb UV rays rather than reflecting them, causing allergies and skin irritation (<xref ref-type="bibr" rid="B510">Mendes-Silva et&#xa0;al., 2020</xref>). Carotenoids curated from bacteria have gained attraction due to their anti-UV properties and antioxidant activities, causing less skin damage by reducing free radical production. It can be used as a natural sunscreen replacing chemical ingredients (<xref ref-type="bibr" rid="B511">Terao, 2023</xref>). Beta-carotene, lycopene, and astaxanthin are some of the naturally sourced carotenoid pigments. Antiaging creams are one of the most popular skin care products, and consumers are turning toward natural components (<xref ref-type="bibr" rid="B512">Chavda et&#xa0;al., 2023</xref>). The study by <xref ref-type="bibr" rid="B37">Kiki (2023)</xref> showed that beta-cryptoxanthin from <italic>Dunaliella salina</italic> and lycopene from <italic>Anabaena vaginicola</italic> can stimulate hyaluronic acid production, increasing skin hydration. <italic>Haematococcus pluvialis</italic> produces astaxanthin, which is known to improve skin moisture content, texture, and wrinkles (<xref ref-type="bibr" rid="B513">Liu, 2022</xref>). Japanese and Swedish skin care companies have marketed products containing astaxanthin for antiaging solution. Skin whitening creams are also very popular among Asian countries (<xref ref-type="bibr" rid="B514">Pollock et&#xa0;al., 2021</xref>). Tyrosinase inhibitors lead to decreased melanin production, catalyzing the pigmentation process. These compounds are being used to treat albinism. Fucoxanthin from <italic>Laminaria japonica</italic> and astaxanthin from <italic>Haematococcus pluvialis</italic> are superior antioxidants due to their catalase and superoxide dismutase capabilities. Topical and oral astaxanthin can improve hyperpigmentation problems by stopping melanin production (<xref ref-type="bibr" rid="B515">Ara&#xfa;jo et&#xa0;al., 2021</xref>). Chemicals like benzene and toluene are used for the vibrant colors of lipsticks and nail polish, which can be replaced by bacterial pigments offering safety and stability (<xref ref-type="bibr" rid="B516">Kalra et&#xa0;al., 2020</xref>). Marine microbes are being extensively studied for their cosmetic applications. Phycocyanin from marine algae is being used in eye shadow and eye makeup. Pigments extracted from red microalgae are used in red, purple, and pink lipstick and eye makeup (<xref ref-type="bibr" rid="B517">Yarkent et&#xa0;al., 2020</xref>). Microbial pigments from several environmental niches have attracted attention and are being studied for their use in the cosmetic industry. This review intends to explore the classification and biosynthetic pathways of bacterial pigments, highlighting the scope of genetic engineering for the enhanced production of pigments. It explores the use of bacterial pigments in cosmetic products and their advantages. This review also discusses the recent advances and technological innovations in pigment production from different bacterial sources. The challenges, limitations, and future perspectives of the revolutionary role of bacterial pigments in the cosmetic industry are also explored in this study.</p>
</sec>
<sec id="s2">
<label>2</label>
<title>Bacterial pigments</title>
<p>Bacterial pigments are characterized as diverse metabolites, which are generated by various microbial species. These pigments impart characteristic colors, but they also play physiological, ecological, and industrial roles. The broad classification of bacterial pigments is mainly based on color, chemical nature, biological function, and solubility (<xref ref-type="bibr" rid="B5">Agarwal et&#xa0;al., 2023</xref>).</p>
<sec id="s2_1">
<label>2.1</label>
<title>Classification of bacterial pigments</title>
<sec id="s2_1_1">
<label>2.1.1</label>
<title>Carotenoids</title>
<p>Carotenoids are yellow, red, or orange pigments that mainly protect bacterial species from oxidative stress. <italic>Micrococcus luteus</italic> and <italic>Serratia marcescens</italic> produce carotenoids (yellow color) and prodigiosin (red color), respectively. Carotenoids are isoprenoid compounds that are normally lipid-soluble (<xref ref-type="bibr" rid="B53">L&#xf3;pez et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s2_1_2">
<label>2.1.2</label>
<title>Melanin</title>
<p>These are black or darkish brown, which are generated through phenolic compound oxidation. Melanin normally protects bacteria from free radicals, enzymatic lysis, and UV radiation. Some of the bacteria that produce melanin are <italic>Vibrio cholerae</italic> and <italic>Shewanella colwelliana</italic> (<xref ref-type="bibr" rid="B14">Celed&#xf3;n and D&#xed;az, 2021</xref>; <xref ref-type="bibr" rid="B92">Thakur et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s2_1_3">
<label>2.1.3</label>
<title>Quinones</title>
<p>These pigments usually work as electron carriers during bacterial respiration and photosynthesis. Some of the common quinones in bacterial species are menaquinones, ubiquinones, chlorobiumquionones, and rhodoquinones. Some of the common bacteria that produce these pigments are <italic>Escherichia coli</italic> (ubiquinones), <italic>P. aeruginosa</italic> (ubiquinones), and <italic>Staphylococcus aureus</italic> (menaquinones) (<xref ref-type="bibr" rid="B14">Celed&#xf3;n and D&#xed;az, 2021</xref>).</p>
</sec>
<sec id="s2_1_4">
<label>2.1.4</label>
<title>Phenazine pigments</title>
<p>These are heterocyclic pigments possessing antimicrobial properties. The most common examples are pyocyanin (blue green) and pyorubin (red pigment), which are produced by <italic>P. aeruginosa</italic>. Also, these pigments act as electron carriers in redox reactions (<xref ref-type="bibr" rid="B1">Abdelaziz et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s2_1_5">
<label>2.1.5</label>
<title>Other pigments</title>
<p>Certain bacteria generate unique pigments that do not fit into any of the above classifications, and examples include <italic>Janthinobacterium lividum</italic>, which produces violacein (purple color), and <italic>Streptomyces coelicolor</italic>, which produces actinorhodin (blue color) (<xref ref-type="bibr" rid="B55">Lyakhovchenko et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s2_1_6">
<label>2.1.6</label>
<title>The Indian Himalayan region as a source of pigment-producing bacteria</title>
<p>Although the high-altitude Himalayan region visibly looks barren and lifeless, beneath its surface, rich and diverse microbial communities are found. These trans-Himalayan niches experience environmental stresses such as fluctuating temperatures, extreme cold, frequent freeze&#x2013;thaw cycles, oxidative stress, high UV radiation, low oxygen levels, and low nutrient availability. The pigment production in the microbes of this region is considered an important adaptive strategy against these extreme conditions. There are a number of pigment-producing bacteria that have been identified from this region, and these are illustrated in <xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>. The list of pigment-producing bacteria/phyla is summarized in <xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Visual presentation of pigment-producing bacteria from the high-altitude trans-Himalayan region: <bold>(a)</bold> <italic>Iodobacter</italic> sp. PCH194, <bold>(b)</bold> <italic>Streptomyces</italic> sp. PCH436, <bold>(c)</bold> <italic>Streptomyces</italic> sp. PCH436, <bold>(d)</bold> <italic>Janthinobacterium</italic> sp. PCH410, <bold>(e)</bold> <italic>Kocuria</italic> sp. PCH206, <bold>(f)</bold> <italic>Pedobacter</italic> sp. PCH18, <bold>(g)</bold> <italic>Pseudomonas</italic> sp. PCH 413, <bold>(h)</bold> <italic>Arthrobacter</italic> sp., <bold>(i)</bold> <italic>Bacillus</italic> sp. PCH164, <bold>(j)</bold> <italic>Flavobacterium</italic> sp. PCH19, <bold>(k)</bold> <italic>Arthrobacter</italic> sp. PCH30, and <bold>(l)</bold> <italic>Leifsonia</italic> sp. PCH178. Reprinted from <italic>Microbial pigments: learning from the Himalayan perspective to industrial applications</italic> by <xref ref-type="bibr" rid="B49">Kumar et al. (2022)</xref>, licensed under Creative Commons CC BY license, <uri xlink:href="https://doi.org/10.1093/jimb/kuac017">https://doi.org/10.1093/jimb/kuac017</uri>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fbrio-05-1746114-g001.tif">
<alt-text content-type="machine-generated">Twelve images show different bacterial colonies on agar plates. Each has varying colors and patterns: (a) Iodobacter sp. PCH194, (b) and (c) Streptomyces sp. PCH436, (d) Janthinobacterium sp. PCH410, (e) Kocuria sp. PCH206, (f) Pedobacter sp. PCH18, (g) Pseudomonas sp. PCH413, (h) Arthrobacter sp., (i) Bacillus sp. PCH164, (j) Flavobacterium sp. PCH19, (k) Arthrobacter sp. PCH30, and (l) Leifsonia sp. PCH178. Each species displays distinct colony morphology and pigmentation.</alt-text>
</graphic></fig>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>List of pigment-producing bacteria/phyla along with pigment name.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Pigment(s)</th>
<th valign="middle" align="left">Bacteria/phylum</th>
<th valign="middle" align="left">Application/activity</th>
<th valign="middle" align="left">References</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Prodigiosin</td>
<td valign="middle" align="left"><italic>Serratia plymuthica</italic> and <italic>Serratia marcescens</italic></td>
<td valign="middle" align="left">Textile dyeing</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B84">Simsek Geyik et&#xa0;al. (2025)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Prodigiosin</td>
<td valign="middle" align="left"><italic>Serratia marcescens</italic> ATCC 8100</td>
<td valign="middle" align="left">Textile dyeing</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B107">Zhang et&#xa0;al. (2024)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Staphyloxanthin</td>
<td valign="middle" align="left"><italic>Staphylococcus aureus</italic> ATCC 6538</td>
<td valign="middle" align="left">Textile dyeing</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B107">Zhang et&#xa0;al. (2024)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Carotenoid</td>
<td valign="middle" align="left"><italic>Paracoccus marcusii</italic> RSPO1</td>
<td valign="middle" align="left">Textile dyeing, antimicrobial and antiallergic activities</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B60">Naik and Gupte (2024)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Pyocyanin</td>
<td valign="middle" align="left"><italic>Pseudomonas aeruginosa</italic></td>
<td valign="middle" align="left">Antibacterial, antioxidant, and anticancer activities</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B1">Abdelaziz et&#xa0;al. (2023)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">&#x3b2;-Carotene</td>
<td valign="middle" align="left"><italic>Citricoccus parietis</italic> AUCs</td>
<td valign="middle" align="left">Antibacterial, antioxidant, and antidiabetic activities</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B30">Hagaggi and Abdul-Raouf (2023)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Blue-green pigment</td>
<td valign="middle" align="left"><italic>Pseudomonas aeruginosa</italic></td>
<td valign="middle" align="left">Fabric dyeing, antimicrobial and antiallergic activities</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B78">Sengupta and Bhowal (2023)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Prodigiosin</td>
<td valign="middle" align="left"><italic>Serratia plymuthica</italic></td>
<td valign="middle" align="left">Fabric dyeing, antibacterial and antioxidant activities</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B7">Amorim et&#xa0;al. (2022)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Flexirubin-type pigment</td>
<td valign="middle" align="left"><italic>Chryseobacterium shigense</italic></td>
<td valign="middle" align="left">Fabric dyeing, antibacterial and antioxidant activities</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B7">Amorim et&#xa0;al. (2022)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Prodigiosin</td>
<td valign="middle" align="left"><italic>Enterobacter</italic> sp. PWN1</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B71">Poddar et&#xa0;al. (2021)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Total carotenoids</td>
<td valign="middle" align="left"><italic>R. toruloides</italic> ATCC204091, <italic>R. glutinis</italic> Y1</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B86">Sinha et&#xa0;al. (2021)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Violacein, deoxyviolacein</td>
<td valign="middle" align="left"><italic>Iodobacter</italic> sp. PCH194</td>
<td valign="middle" align="left">Anticancer and antimicrobial activities</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B44">Kumar et&#xa0;al. (2021)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Total carotenoids</td>
<td valign="middle" align="left"><italic>R. toruloides</italic>, <italic>L. starkeyi</italic></td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B52">Liu et&#xa0;al. (2020)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Torularhodin, &#x3b2;-carotene, torulene</td>
<td valign="middle" align="left"><italic>R. mucilaginosa</italic></td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B26">Ghilardi et&#xa0;al. (2020)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Lycopene</td>
<td valign="middle" align="left"><italic>R. faecalis</italic> PA2</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B69">Patthawaro et&#xa0;al. (2020)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Torularhodin, &#x3b2;-carotene, torulene</td>
<td valign="middle" align="left"><italic>R. mucilaginosa</italic> MTCC-1403</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B79">Sharma and Ghoshal (2020)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">&#x3b2;-Carotene</td>
<td valign="middle" align="left"><italic>R. gracilis</italic> ATCC 10788</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B40">Kot et&#xa0;al. (2020)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Red pigment</td>
<td valign="middle" align="left"><italic>Rhodonellum psychrophillum</italic></td>
<td valign="middle" align="left">Fabric dyeing and antimicrobial activity</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B13">Bisht et&#xa0;al. (2020)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">&#x3b2;-Carotene</td>
<td valign="middle" align="left"><italic>B. trispora</italic> MTCC 884</td>
<td valign="middle" align="left">Antioxidant activity</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B36">Kaur et&#xa0;al. (2019)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Yellow color</td>
<td valign="middle" align="left"><italic>Flavobacterium bomense</italic> sp.</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B51">Liu et&#xa0;al. (2019)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Yellow, orange, brown, violet, and pinkish-red</td>
<td valign="middle" align="left">Proteobacteria, Firmicutes, Actinobacteria, Bacteroidetes</td>
<td valign="middle" align="left">Antioxidant activity</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B67">Panwar et&#xa0;al. (2019)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Carotenoids (various colors)</td>
<td valign="middle" align="left">Proteobacteria, Actinobacteria, Bacteroidetes, and Firmicutes</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B80">Shen et&#xa0;al. (2018)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Violacein</td>
<td valign="middle" align="left"><italic>C. violaceum</italic> UTM5</td>
<td valign="middle" align="left">Anticancer and antimicrobial activities</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B8">Aruldass et&#xa0;al. (2018)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Prodigiosin (red pigment)</td>
<td valign="middle" align="left"><italic>Serratia nematodiphila</italic> RL2</td>
<td valign="middle" align="left">Antimicrobial activity</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B27">Gondil et&#xa0;al. (2017)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Total carotenoids</td>
<td valign="middle" align="left"><italic>R. glutinis</italic></td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B35">Kanzy et&#xa0;al. (2015)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Carotenoids (yellow-orange)</td>
<td valign="middle" align="left"><italic>Sanguibacter suarezii</italic> KK6, <italic>Kocuria turfanensis</italic> KK7, <italic>Kocuria rosea</italic> KK12, <italic>Planococcus maritimus</italic> KK21</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B45">Kushwaha et&#xa0;al. (2014)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Various pigments</td>
<td valign="middle" align="left">Firmicutes, alpha- and gamma-Proteobacteria, Actinobacteria</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B81">Shen et&#xa0;al. (2012)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Total carotenoids</td>
<td valign="middle" align="left"><italic>R.</italic> sp<italic>haeroides</italic> O.U.001</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B23">Eroglu et&#xa0;al. (2010)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Violacein (violet)</td>
<td valign="middle" align="left"><italic>Janthinobacterium lividum</italic> XT1</td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B54">Lu et&#xa0;al. (2009)</xref></td>
</tr>
<tr>
<td valign="middle" align="left">Yellow pigment</td>
<td valign="middle" align="left"><italic>Leifsonia pindariensis</italic></td>
<td valign="middle" align="left">Pigment production</td>
<td valign="middle" align="left"><xref ref-type="bibr" rid="B75">Reddy et&#xa0;al. (2008)</xref></td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Biosynthetic pathways of pigment production in bacteria</title>
<p>Bacteria generate various pigments through special biosynthetic pathways, which originate from primary metabolic intermediates. The biosynthetic pathways are often regulated by environmental and genetic factors. Understanding bacterial biosynthetic pathways is crucial in aiding metabolic engineering for enhanced pigment synthesis. Several biosynthetic routes differ on the basis of the chemical nature of the pigment (<xref ref-type="bibr" rid="B5">Agarwal et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B10">Barreto et&#xa0;al., 2023</xref>).</p>
<sec id="s2_2_1">
<label>2.2.1</label>
<title>Biosynthesis of carotenoids</title>
<p>The synthesis of carotenoids begins with isopentenyl pyrophosphate (IPP) and dimethylallyl pyrophosphate (DMAPP) condensation, to form geranylgeranyl pyrophosphate (GGPP). Two molecules of geranylgeranyl pyrophosphate combine to form phytoene, which later undergoes cyclization and desaturation reactions, producing carotenoids such as lycopene, zeaxanthin, and &#x3b2;-carotene. Certain bacteria such as <italic>Rhodobacter</italic> spp. and <italic>M. luteus</italic> follow this biosynthetic pathway (<xref ref-type="bibr" rid="B100">Wang et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s2_2_2">
<label>2.2.2</label>
<title>Biosynthesis of phenazine</title>
<p>Pigments such as pyocyanin are generated in <italic>P. aeruginosa</italic> through the shikimate pathway. Chorismic acid functions as a key intermediate and undergoes various enzymatic reactions, which is catalyzed by the <italic>phz</italic> (phenazine biosynthetic) gene cluster. This leads to the production of phenazine-1-carboxylic acid (PCA), which can be later modified into pyorubin and pyocyanin (<xref ref-type="bibr" rid="B1">Abdelaziz et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s2_2_3">
<label>2.2.3</label>
<title>Biosynthesis of melanin</title>
<p>Melanin is formed through oxidation of tyrosine, with the help of the enzyme tyrosinase, leading to the generation of dopaquinone, which gradually polymerizes into dark melanin. Certain other bacteria produce melanin through the 1,8-dihydroxynaphthalene (DHN) or homogentisate pathways (<xref ref-type="bibr" rid="B70">Pavan et&#xa0;al., 2020</xref>).</p>
</sec>
<sec id="s2_2_4">
<label>2.2.4</label>
<title>Biosynthesis of prodigiosin</title>
<p><italic>Serratia marcescens</italic> synthesizes the red pigment prodigiosin through the tripyrrole pathway. 2-Methyl-3-n-amyl-pyrrole and 4-methoxy-2,2&#x2032;-bipyrrole-5-carboxyaldehyde are the two intermediates formed in this pathway. These two intermediates are condensed with the help of certain enzymes to produce prodigiosin (<xref ref-type="bibr" rid="B101">Williams, 1973</xref>).</p>
</sec>
<sec id="s2_2_5">
<label>2.2.5</label>
<title>Biosynthesis of violacein</title>
<p><italic>Chromobacterium violaceum</italic> synthesizes violacein from the amino acid tryptophan. The amino acid tryptophan is converted into indole derivatives using an enzyme coded by the vioABCDE gene cluster. Indole derivatives gradually undergo oxidative coupling in order to form violacein (<xref ref-type="bibr" rid="B34">Hui et&#xa0;al., 2022</xref>).</p>
</sec>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Genetic and metabolic engineering approaches for enhanced pigment production</title>
<p>Most of the bacterial pigments such as carotenoids, melanin, and violacein have significant applications in the cosmetic, pharmaceutical, textile, and food industries (<xref ref-type="bibr" rid="B90">Sundar and Sivaperumal, 2022</xref>). However, natural pigment production is mostly limited due to low yields, suboptimal metabolic fluxes, and regulatory constraints. In order to overcome these, genetic and metabolic engineering approaches are employed nowadays to enhance the biosynthesis of bacterial pigments (<xref ref-type="bibr" rid="B47">Lee and Schmidt-Dannert, 2002</xref>).</p>
<sec id="s2_3_1">
<label>2.3.1</label>
<title>Genetic engineering</title>
<p>This includes direct manipulation of bacterial pigment-producing genes and metabolic pathways. Some of the techniques such as heterologous expression, gene deletion, and overexpression have transformed pigment biosynthesis. In case of gene overexpression, genes controlling rate-regulating enzymes are amplified to increase bacterial pigment yield. For instance, in <italic>Rhodobacter</italic> sp<italic>haeroides</italic>, overexpression of the <italic>crtY</italic> gene increases &#x3b2;-carotene synthesis. An overview of the promoter optimization strategy is shown in <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref> (<xref ref-type="bibr" rid="B72">Qiang et&#xa0;al., 2019</xref>). In another work, from <italic>C. violaceum</italic>, overexpression of the genes vioA&#x2013;E increases the violacein production in <italic>E. coli</italic> (<xref ref-type="bibr" rid="B9">Balibar and Walsh, 2006</xref>). In this mechanism, VioA (flavoenzyme) and VioB (heme protein) cooperatively oxidize and dimerize L-tryptophan to result in an intermediate that can yield chromopyrrolic acid off-pathway. VioE presence precedes the reaction via [1,2]-indole rearrangement to prodeoxyviolacein. The final flavin-dependent oxygenases, VioD and VioC, sequentially hydroxylate and oxidize the indole rings at positions 5 and 2, respectively, completing violacein biosynthesis.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Graphical presentation of the promoter optimization strategy in the engineered <italic>Rhodobacter</italic> sp<italic>haeroides</italic> for &#x3b2;-carotene synthesis. Reprinted with permission from <xref ref-type="bibr" rid="B72">Qiang et&#xa0;al. (2019)</xref>. Copyright (2019) by the American Chemical Society. Opt, codon optimization; CrtY, lycopene cyclase (from <italic>P. agglomerans)</italic>; Zwf, glucose-6-phosphate dehydrogenase; Dxs, 1-deoxy-D-xylulose-5-phosphate synthase.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fbrio-05-1746114-g002.tif">
<alt-text content-type="machine-generated">Metabolic pathway diagram showing the conversion of glucose to β-carotene. Key intermediates include G6P, G3P, DXP, MEP, FPP, phytoene, lycopene, and β-carotene. Enzymes zwf and dxs are shown in red. Promoter optimization for opt crtY is illustrated with different promoters (PJ95025, PJ95026, PJ95027, Ptac, PrrnB) in a separate section.</alt-text>
</graphic></fig>
<p>In another study, transcription factor and promoter engineering was performed to enhance prodigiosin production in <italic>Serratia marcescens</italic> JNB5-1 (<xref ref-type="bibr" rid="B66">Pan et&#xa0;al., 2022</xref>). In this study, first, a Tn5G transposon insertion library identified the response regulator BVG89_19895 (OmpR) as a positive regulator of prodigiosin synthesis. Thereafter, RNA-Seq, GFP reporter, and RT-qPCR analyses revealed the promoter P17 (P<sub>RplJ</sub>) as a strong constitutive promoter. Overexpression of OmpR and PsrA in the strain JNB5&#x2013;1 generated a recombinant strain PG-6. The generated recombinant strain achieved a prodigiosin titer of 10.25 g/L, which was 1.62-fold higher than the parent strain (6.33 g/L).</p>
</sec>
<sec id="s2_3_2">
<label>2.3.2</label>
<title>Metabolic engineering approaches</title>
<p>The rise in the demand for carotenoid pigments has laid the foundation for the increased production of these pigments through metabolic engineering of their biosynthesis pathways. The first attempt for lycopene production was carried out in a fungus (<italic>S. cerevisiae</italic>) with the carotenogenic genes of the Eubacterium <italic>Pantoea ananatis</italic> (<xref ref-type="bibr" rid="B103">Yamano et&#xa0;al., 1994</xref>). However, considerable efforts have been made in the metabolic engineering of <italic>E. coli</italic> to produce carotenoids (lycopene, &#x3b2;-carotene, and zeaxanthin) at a higher titer (<xref ref-type="bibr" rid="B20">Das et&#xa0;al., 2007</xref>). <italic>Escherichia coli</italic> has been observed as a choice of host for the heterologous carotenoid production due to the fact that it can accommodate and express the carotenogenic genes from bacteria, fungi, and plants. In this context, the improved production of carotenoid in <italic>E. coli</italic> has been achieved through metabolic engineering of the endogenous MEP (2-C-methyl-D-erythritol4-phosphate) pathway and through introducing a heterologous mevalonate (MVA) pathway. The engineering of these pathways is intended to increase the intracellular supply of isopentenyl diphosphate (IPP), which is a key precursor for carotenoid biosynthesis. Furthermore, improvements in carotenoid production have been made by engineering the prenyl diphosphate and central metabolic pathways (<xref ref-type="bibr" rid="B47">Lee and Schmidt-Dannert, 2002</xref>). The MVA and MEP pathways begin with substrates, viz., acetyl-CoA, pyruvate, and glyceraldehyde-3-phosphate (G3P), produced in the glycolytic pathway (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3</bold></xref>). Therefore, regulating the production of these metabolites can improve carotenoid production. Pyruvate is considered a more crucial precursor, as it finds itself in multiple metabolic routes and is more abundant than G3P for isoprenoid biosynthesis. The MEP pathway utilizes pyruvate and G3P in equal amounts for IPP synthesis. Hence, modulation of these intermediate enzymes can alter the metabolic flux between these intermediates to enhance lycopene production in <italic>E. coli</italic> (<xref ref-type="bibr" rid="B24">Farmer and Liao, 2001</xref>). In these studies, it was observed that G3P is a limiting factor in lycopene biosynthesis. Therefore, inactivation of the competing pathways branching at the pyruvate and acetyl-CoA nodes increases 45% of lycopene production in the engineered <italic>E. coli</italic> strain as compared to the parent <italic>E. coli</italic> strain (<xref ref-type="bibr" rid="B93">Vadali et&#xa0;al., 2005</xref>). Production was further improved by incorporating the MVA pathway from the <italic>Streptomyces</italic> sp. strain CL190 into the <italic>E. coli</italic> strain (inactivated). This bioengineering fostered the acetyl-CoA pool for IPP synthesis, which eventually resulted in a 2-fold increase in lycopene production as compared to the native MEP pathway. In another study, the co-expression of the exogenous MEP pathway genes (<italic>dxs</italic> and <italic>idi</italic>) in <italic>E. coli</italic> promoted the transcription of isoprenoid pathway genes (<xref ref-type="bibr" rid="B99">Wang et&#xa0;al., 2015</xref>). In this case, there was a 16.5 times increase in the yield (20.57 mg/L) of lycopene.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Schematic overview of &#x3b2;-carotene biosynthesis via the MEP and MVA pathways. Reprinted from <xref ref-type="bibr" rid="B104">Yang and Guo (2014)</xref>; article is under Commons Attribution License (<ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0">http://creativecommons.org/licenses/by/4.0</ext-link>).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fbrio-05-1746114-g003.tif">
<alt-text content-type="machine-generated">Diagram showing the MEP and MVA pathways for isoprenoid synthesis. The MEP pathway converts G-3-P to IPP and DMAPP via DXP, with key enzymes like dxs and dxr. The MVA pathway converts acetyl-CoA to IPP using enzymes such as mvaE and mvaS. Both pathways converge at IPP, leading to β-carotene synthesis through intermediates like GGPP, requiring enzymes such as crtE, crtB, and crtI.</alt-text>
</graphic></fig>
<p>Introduction of a heterologous MVA pathway into <italic>E. coli</italic> represents a promising strategy for enhancing carotenoid biosynthesis. The MVA pathway can be functionally divided into two segments: the upper (top) and lower (bottom) pathways. The final step of the upper pathway, catalyzed by HMG-CoA reductase, serves as a rate-limiting step in the process. In this regard, promoter-driven overexpression of the <italic>cHMG1</italic> gene from <italic>S. cerevisiae</italic> in <italic>Neurospora crassa</italic> has been shown to increase the production of lycopene and neurosporaxanthin (<xref ref-type="bibr" rid="B98">Wang and Keasling, 2002</xref>). Similarly, <italic>E. coli</italic> has also been metabolically engineered for the co-overexpression of <italic>crtY</italic>, <italic>crtZ</italic>, and <italic>ZEP</italic> encoding the enzymes lycopene &#x3b2;-cyclase, &#x3b2;-carotene 3-hydroxylase, and zeaxanthin epoxidase, respectively (<xref ref-type="bibr" rid="B102">Xinrui et&#xa0;al., 2023</xref>). The engineered strain was able to produce a violaxanthin yield of 25.28 &#xb1; 3.94 mg/g DCW along with a lowering in the accumulation of by-products. <xref ref-type="bibr" rid="B15">Cheng et&#xa0;al. (2022)</xref> introduced the heterologous lycopene biosynthetic pathway into <italic>E. coli</italic>, and this was organized into three modules. In the modules, MVA and dimethylallyl pyrophosphate (DMAPP) acted as connecting intermediates. The intermodule balance was optimized by controlling the expression level of the genes, viz., <italic>MVK</italic>, <italic>PMK</italic>, <italic>MVD</italic>, and <italic>IDI</italic> (involved in downstream), through a ribosome binding site library with defined expression strengths. The bioengineering resulted in 4.6 times more yield of lycopene (219.7&#xa0;mg/g DCW) as compared to the reference strain.</p>
</sec>
<sec id="s2_3_3">
<label>2.3.3</label>
<title>Synthetic biology and advanced engineering tools</title>
<p>CRISPR/Cas9 is an exciting technology for bioengineering the metabolic pathway of bacteria for desired applications. However, it has been less explored for pigment production in bacteria. CRISPR has been used to enhance &#x3b2;-carotene production from the IPP and dimethylallyl pyrophosphate (DMAPP) through the MEP pathway. The technology has been used to engineer the central metabolic routes and regulate the metabolic fluxes by targeted gene deletions and strategic gene overexpression (<xref ref-type="bibr" rid="B16">Cho et&#xa0;al., 2018</xref>). In another work, <italic>E. coli</italic> strains possessing the biosynthetic MVA pathway and terpenoid synthases (plant-derived) have been engineered using the CRISPR interference (CRISPRi) system for suppressing the expression of acetoacetyl-CoA thiolase enzyme. This enzyme is responsible for catalyzing the first step of the MVA pathway (<xref ref-type="bibr" rid="B39">Kim et&#xa0;al., 2016</xref>). This bioengineering has been observed to enhance the production of (&#x2212;)-&#x3b1;-bisabolol (C15) and lycopene (C40). Furthermore, it also mitigated cell growth inhibition and accumulation of toxic intermediates within the MVA pathway. Similarly, CRISPRi-based library targeting of 74 genes was done in <italic>Corynebacterium glutamicum</italic> (<xref ref-type="bibr" rid="B29">G&#xf6;ttl et&#xa0;al., 2021</xref>). The results indicated that CRISPRi-mediated repression of the carotenogenesis repressor gene <italic>crtR</italic> led to enhanced pigmentation and increased accumulation of the native carotenoid pigment decaprenoxanthin. CRISPRi screening identified 14 genes whose repression influenced decaprenoxanthin biosynthesis. Out of the 14 genes identified, repression of 11 genes reduced carotenoid production, and repression of 3 genes promoted decaprenoxanthin synthesis. Furthermore, deletion of <italic>pgi</italic> and <italic>gapA</italic> genes enhanced decaprenoxanthin production by 43- and 9-fold, respectively. Another study utilized the CRISPR/Cas9-based genome editing system for actinomycetes for two chromogenic reporter systems (GusA and IdgS) (<xref ref-type="bibr" rid="B29">G&#xf6;ttl et&#xa0;al., 2021</xref>). The researchers deleted a single gene, i.e., <italic>actIORFI</italic> (SCO5087 of the actinorhodin gene cluster), in <italic>S. coelicolor</italic> M145. They also deleted the small gene cluster of 5.5 kb responsible for orange-pigmented carotenoid production and a relatively large gene cluster (61 kb) corresponding to the abyssomicin biosynthetic pathway in <italic>Verrucosispora</italic> sp. MS100137.</p>
</sec>
</sec>
</sec>
<sec id="s3">
<label>3</label>
<title>Applications in cosmetic formulations</title>
<sec id="s3_1">
<label>3.1</label>
<title>Cosmetic products incorporating bacterial pigments</title>
<p>Biodegradability, stability, and bioactivity have placed bacterial pigments in the spotlight as alternatives to synthetic dyes in the cosmetics sector and as being environmentally friendly, sustainable, and natural (<xref ref-type="bibr" rid="B10">Barreto et&#xa0;al., 2023</xref>). These pigments are incorporated into a broad category of cosmetic products such as lipsticks, skin creams and lotions, nail polishes, hair dyes, soaps, and body washes.</p>
<sec id="s3_1_1">
<label>3.1.1</label>
<title>Lipsticks, nail polishes, and hair dyes</title>
<p>Microbial carotenoids and prodigiosin-like pigments have color intensities ranging between red and orange. These pigments can be natural alternatives to azo dyes (<xref ref-type="bibr" rid="B21">Devi et&#xa0;al., 2024</xref>). Their high coloring intensity along with their antioxidant activity can be the reason for their acceptance in lip care applications. Furthermore, these natural pigments reduce the risk of direct skin contact and ingestion of synthetic chemicals, hence are biocompatible (<xref ref-type="bibr" rid="B33">Huang et&#xa0;al., 2024b</xref>). Similarly, these microbial pigments can also be used for nail polishes, as they cover a broad range of colors. These natural pigments derived from bacteria are biocompatible, and their use in place of chemical-based nail polishes can help minimize adverse effects associated with synthetic colorants. Furthermore, these colors could be the best option for the manufacturing of natural hair colors.</p>
</sec>
<sec id="s3_1_2">
<label>3.1.2</label>
<title>Skin creams and lotions</title>
<p>It is well known that UV radiation from the sun can lead to adverse health effects such as pigmentation, erythema, immunosuppression, photoaging, and even skin cancer (<xref ref-type="bibr" rid="B17">Choksi et&#xa0;al., 2020</xref>). Therefore, sunscreens are commonly used to protect the skin from UV rays. Most sunscreens in the market utilize chemicals for the screening of these UV rays, but due to the growing awareness and demand for natural alternatives in recent years, there is a need for natural and effective pigments for skin products. Researchers are also focusing on identifying new, cost-effective natural sunscreen agents from bacterial and other natural sources (<xref ref-type="bibr" rid="B28">Goswami et&#xa0;al., 2013</xref>). There are a number of microorganisms that produce pigments and provide protection against UV-induced damage (<xref ref-type="bibr" rid="B91">Suryawanshi et&#xa0;al., 2015</xref>). Bacterial pigments such as prodigiosin, violacein, and melanin have been reported by various research groups for their strong UV-protective effect (<xref ref-type="bibr" rid="B83">Siezen, 2011</xref>; <xref ref-type="bibr" rid="B19">Darshan and Manonmani, 2015</xref>; <xref ref-type="bibr" rid="B41">Kothari et&#xa0;al., 2017</xref>). These pigments also exhibit strong antioxidant activity, making them suitable for incorporation into creams and lotions to enhance skin protection. In a study, the bacterium <italic>Virgibacillus salarius</italic> strain 19.PP.Sc1.6 has been identified for carotenoid production (<xref ref-type="bibr" rid="B46">Kusmita et&#xa0;al., 2021</xref>). The carotenoid pigments from the bacterium were utilized in the development of antiaging creams and demonstrated significant antiaging activity.</p>
</sec>
<sec id="s3_1_3">
<label>3.1.3</label>
<title>Soaps and body washes</title>
<p>Soaps and body washes have not been documented with bacterial pigments, but there are a few reports where fungal pigments and other plant pigments were utilized in these products. Asnani and Diastuti developed a glycerine soap utilizing the red pigment from <italic>Streptomyces</italic> K-4 B and Dayak onion (<italic>Eleutherine palmifolia</italic> (L.) Merr) (<xref ref-type="bibr" rid="B31">Herlina and Diastuti, 2017</xref>). The results of the developed product were assessed based on color, aroma, texture, foam, and rough impression upon usage and after usage and were observed to be in the &#x201c;like to very like soap&#x201d; category (score of 4.67 in the scale of 1 to 5) and &#x201c;like to very like&#x201d; category (score of 4.30 in the scale of 1 to 5). Additionally, bacterial glycolipids and lipopeptides have been reported for their emulsifying, detergency, foaming, and skin moisturizing capabilities. The pigments from bacteria can be an alternative to chemical colors along with their emulsifying, moisturizing, and antioxidant properties. However, this area of pigment-producing bacteria is not well established, and still a lot of biotechnological interventions that can be made.</p>
</sec>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Functional benefits beyond coloring</title>
<p>In addition to their ability to color, bacterial pigments have functional activities such as antioxidant, antimicrobial, and antiaging activities, which have contributed to increasing the therapeutic properties of cosmetics.</p>
<sec id="s3_2_1">
<label>3.2.1</label>
<title>Antioxidant activity</title>
<p>Carotenoid, violacein, and prodigiosin pigments possess an effective free-radical scavenging and lipid peroxidation inhibitory effect, playing a role in photoprotection of the skin and related repair (<xref ref-type="bibr" rid="B37">Kiki, 2023</xref>; <xref ref-type="bibr" rid="B21">Devi et&#xa0;al., 2024</xref>). Incorporated into topical preparations, these pigments prevent oxidative stress caused by UV radiation and pollutants, hence preserving the dermal homeostasis (<xref ref-type="bibr" rid="B95">Venil et&#xa0;al., 2020</xref>).</p>
</sec>
<sec id="s3_2_2">
<label>3.2.2</label>
<title>Antimicrobial activities</title>
<p>Numerous bacterial pigments, such as prodigiosin and pyocyanin, have neutral antimicrobial activity against <italic>S. aureus</italic>, <italic>Candida albicans</italic>, and <italic>E. coli</italic> (<xref ref-type="bibr" rid="B106">Yusuf et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B76">Salman et&#xa0;al., 2025</xref>). In a study, <italic>Micrococcus</italic> species of bacteria isolated from soil samples were reported to produce a yellow pigment (<xref ref-type="bibr" rid="B85">Sinha et&#xa0;al., 2017</xref>). The pigments were observed for their antibacterial activity against <italic>S. aureus</italic> (23&#xa0;mm), <italic>Candida</italic> (14&#xa0;mm), and <italic>S. typhi</italic> (10&#xa0;mm), which was comparable to penicillin (19&#xa0;mm) and gentamicin (17&#xa0;mm) against <italic>S. aureus</italic>. In another study, the bacterial pigment carotenoid from <italic>Rhodotorula glutinis</italic> (PTCC 5256) was able to exhibit significant antimicrobial activity against <italic>Salmonella enteritidis</italic> and <italic>E. coli</italic> (<xref ref-type="bibr" rid="B105">Yolmeh, 2016</xref>). Likewise, <italic>S. marcescens</italic>, <italic>C. violaceum</italic>, and <italic>P. aeruginosa</italic> were observed to produce pigments such as prodigiosin, violacein, and pyocyanin, respectively (<xref ref-type="bibr" rid="B76">Salman et&#xa0;al., 2025</xref>). These pigments have been able to inhibit the growth of 30 clinical isolates of <italic>Enterococcus faecalis</italic>, <italic>Klebsiella</italic> spp., and <italic>Candida</italic> spp.</p>
</sec>
<sec id="s3_2_3">
<label>3.2.3</label>
<title>Antiaging activities</title>
<p>Microbial pigments as antiaging bioactives reduce oxidative stress and preserve collagen integrity. Carotenoid creams and violacein emulsions increase the elasticity of the skin and inhibit photoaging by suppressing reactive oxygen species (<xref ref-type="bibr" rid="B63">Orlandi et&#xa0;al., 2022</xref>). Furthermore, they have the potential to suppress the activity of tyrosinase, creating skin-brightening effects, and thus extending their uses in antiaging and cosmeceutical collections. The bacteriochlorophyll <italic>a</italic> pigment from <italic>R.</italic> sp<italic>haeroides</italic> has been observed for its antioxidant activity of 63.8% in aqueous extract form (<xref ref-type="bibr" rid="B38">Kim et&#xa0;al., 2015</xref>). Similarly, s pigment&#x2013;protein complex from <italic>Chlorella pyrenoidosa</italic> has been observed to inhibit the inflammatory cytokines TNF-&#x3b1; and IL-6 and inflammatory mediator nitric oxide formation in simulated conditions (<xref ref-type="bibr" rid="B108">Zhang et&#xa0;al., 2019</xref>). Likewise, carotenoid pigments from <italic>Flavobacterium</italic> sp. and <italic>Brevibacterium</italic> sp. were able to provide significant protection against UVA-B to UVC radiation (<xref ref-type="bibr" rid="B68">Patki et&#xa0;al., 2021</xref>). These pigments further exhibited good antimicrobial activity against <italic>E. coli</italic>, <italic>Corynebacterium diphtheriae</italic>, and <italic>S. aureus</italic> along with remarkable antioxidant activity. These multifunctional properties from pigment-producing bacteria show their potential use in cosmetic products. However, limited studies have been done in this field, and more research is required for its global acceptance and recognition.</p>
</sec>
</sec>
</sec>
<sec id="s4">
<label>4</label>
<title>Recent advances and technological innovations</title>
<p>Refinement of innate pigment-producing bacterial strains could be achieved through biotechnological (genetic engineering) strategies for short-term fermentations, giving maximal yields. This is possible through appropriate screening of the best (hyper) pigment-producing bacterial strains, targeting the genetic matter that acts as a regulator/determinant of the respective bacterial pigments (<xref ref-type="bibr" rid="B63">Orlandi et&#xa0;al., 2022</xref>). Genetic engineering of pigment-producing bacterial strains may involve random mutagenesis or specific site-directed mutagenesis to increase the desired pigment production. This strain improvement was attained using microwave (bacterial prodigiosin from <italic>S. marcescens</italic>) and ultraviolet radiations and chemicals like 1-methyl-3-nitro-1-nitrosoguanidine and ethyl methane sulfonate, to mention a few (<xref ref-type="bibr" rid="B82">Shrestha et&#xa0;al., 2025</xref>). Violacein, from the strain <italic>Pseudogulbenkiania ferrooxidans</italic>, has contributed to the cosmetic industry, with its whole genome data relied upon to better understand its biosynthesis pathway (<xref ref-type="bibr" rid="B58">Mehta et&#xa0;al., 2025</xref>). Progressive advances in the field of genetic engineering have paved the way for positive shifts in innate bacterial strains, leading to the production of pigments of interest to researchers/industrialists. To introduce a specific case, the blue pigment-producing <italic>S. coelicolor</italic> might be manipulated genetically to produce red, brilliant yellow, yellow, and orange pigments, thus being capable of generating inherent and acquired colors (pigments) (<xref ref-type="bibr" rid="B58">Mehta et&#xa0;al., 2025</xref>).</p>
<p>Metabolic engineering and microbial bioconversions aid fermentation microbiologists in scaling up the laboratory bioprocesses (<xref ref-type="bibr" rid="B56">Mandal and Majumdar, 2023</xref>). Alternative biotechnological preference focused on the augmentation of the pigment-producing strains&#x2019; culture medium with applicable stimulators, thereby physically and statistically modeling and optimizing the fermentation pathways for shorter periods and maximum bacterial pigment gains. Pardhan et&#xa0;al. reported the psychrotolerant <italic>Paenibacillus</italic> sp. BPW19 for the first time, which produced intracellular pink pigment and was isolated from wastewater, belonging to the <italic>Paenibacillus</italic> genus (<xref ref-type="bibr" rid="B65">Padhan et&#xa0;al., 2021</xref>). In a cost-effective (zero-energy) manner, the production of the pigment and its solvent extraction resulted in high pigment gains. In the last half decade, the pigment-producing bacterial strains&#x2019; responses to their vicinity and surroundings (microbial communications) are well-studied in terms of quorum sensing and evocation, as a model for the extra production of bacterial pigments at an industrial scale, particularly prodigiosin and violacein pigments, as well as deep blue pigments (<italic>Pantoea agglomerans</italic>) (<xref ref-type="bibr" rid="B25">Fujikawa and Akimoto, 2011</xref>). Gene cloning is another prominent strategy to produce positive clones of the desired pigment producers with a suitable expression vector. The effective application of bacterial pigments as colorants in cosmetics may be restrained due to weak water solubility, poor chemical stability, delicate skin absorption, and minimal bioavailability factors. Over and above that, few biopigments (biocolorants) have high molecular weight, which can hinder their penetration (permeation) into the <italic>stratum corneum</italic> (the foremost skin layer) (<xref ref-type="bibr" rid="B96">Venil et&#xa0;al., 2013</xref>). In light of the current circumstances, it becomes mandatory for an industrialist to appropriately design product delivery systems in an efficient and scientific way that allows easy and safe penetration into the skin. Such highly unstable, less bioavailable, poorly soluble biopigments need advanced tools for systematic delivery and best skin outcomes. In the recent past, bacterial pigments like astaxanthin and carotenoids were capable of defending against the aging processes, preventing skin protein degradation. To further enhance the pigment delivery process, tetraethyl orthosilicate inside lecithin vesicles formed silicified liposomes, also known as silicated phospholipids, which served as the transporters of astaxanthin in cosmetic products (<xref ref-type="bibr" rid="B50">Liu and Nizet, 2009</xref>; <xref ref-type="bibr" rid="B59">Moura-Alves et&#xa0;al., 2014</xref>). This combination, along with boron nitride&#x2014;one of the prevailing cosmetic ingredients, gives&#xa0;the best blend for superior adherence to smooth skin and eventually fine-tuning the skin texture (<xref ref-type="bibr" rid="B49">Li et&#xa0;al., 2024</xref>). Such astaxanthin liposomes could be incorporated in lipsticks, concealers, and even eye skin care products (<xref ref-type="bibr" rid="B87">Souto et&#xa0;al., 2020</xref>). Following microencapsulation, nanoencapsulation, liposomes by spray drying,&#xa0;emulsions by spray drying, lyophilization (freeze-drying) encapsulation, hydrogel encapsulation, and co-crystallization encapsulation are popular delivery systems of bacterial pigments, universally in the case of carotenoids (<xref ref-type="bibr" rid="B12">Bhandari et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B22">D&#xed;az-Montes, 2025</xref>). Considering economic viability, scalability, and safeness, researchers choose prevalent encapsulation methodologies with utmost care, understanding the characteristics and nature of the&#xa0;bacteria-derived pigments being encapsulated. The unification and implementation of the newest tools and technologies in the fields of protein engineering, systems biology, bioprocess technology for optimized yields, process engineering, high-efficiency methodologies for pigment producers&#x2019; cultivation, survival-of-the-fittest screening and genome analyses, and metabolic engineering of pigment-producing bacteria for high yields is anticipated to become all-powerful and scientifically influential (<xref ref-type="bibr" rid="B97">Vieira et&#xa0;al., 2020</xref>). Along with the advancements and innovations in biotechnology, synthetic biology, chemistry, and forthcoming metabolic engineering, the production of microbes, particularly bacteria, will enormously amplify the number of biopigments that could be produced within stipulated fermentation periods economically (<xref ref-type="bibr" rid="B4">Acharya et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s5">
<label>5</label>
<title>Challenges and future perspectives</title>
<p>Although there are several pigment-producing microbes existing in diverse environmental conditions, the production and yield of pigments by bacteria and other microorganisms are highly dependent on factors such as pH, temperature, media constituents/substrate, seasonal climatic variations, location, habitats, and genetic constitution (<xref ref-type="bibr" rid="B62">Numan et&#xa0;al., 2018</xref>). Furthermore, the production of bacterial pigments through fermentation is affected by the bioreactors and their design, type of fermentation process, and physicochemical and biological conditions (<xref ref-type="bibr" rid="B94">Venil et&#xa0;al., 2014</xref>). The nature of pigments is highly variable, and as a result, their solubility also differs. Therefore, identifying a suitable solvent system for each pigment is essential. Since different bacteria and microbes produce pigments at varying temperatures, analyzing the thermal stability of these pigments is an important criterion that needs to be explored. Although bacterial pigments possess several properties that make them suitable for use in cosmetic products, studies on their toxicity remain limited. For global acceptance and recognition, extensive toxicological evaluations of these pigments are essential (<xref ref-type="bibr" rid="B61">Nainangu et&#xa0;al., 2025</xref>). Such studies will also help enhance their industrial demand and applications (<xref ref-type="bibr" rid="B74">Ramesh et&#xa0;al., 2019</xref>). All cosmetic pigments, including microbial pigments, fall under the U.S. FDA&#x2019;s Federal Food, Drug, and Cosmetic Act (FD&amp;C Act) and the Modernization of Cosmetics Regulation Act (MoCRA) (<xref ref-type="bibr" rid="B6">Alshehrei, 2024</xref>). According to these acts and regulations, it is mandatory to conduct toxicity testing, heavy metal analysis, microbiological analysis, and particle size evaluation without pre-market approval for cosmetic products. Color additives must receive specific FDA approval before their use in cosmetic products, and the microbial limit is set at 10,000 CFU/g for general topical cosmetics with the complete absence of pathogens like <italic>P. aeruginosa</italic>, <italic>S. aureus</italic>, and <italic>C. albicans</italic>. For each pigment batch, it is important to obtain the certificates of analysis from designated officials (<xref ref-type="bibr" rid="B32">Huang et&#xa0;al., 2024a</xref>). Furthermore, the product should contain a safety data sheet to provide detailed hazard information in the Globally Harmonized System format. Cosmetic products should also contain a technical specification sheet describing the physical and chemical properties of the pigments, such as color index numbers, chemical structure, solubility, and stability.</p>
<p>The yield of pigment produced by native bacterial strains is often very low; however, this can be enhanced by optimizing the growth conditions of bacteria. Furthermore, this yield can be improved by employing the response surface methodology in combination with artificial neural network like statistical approaches (<xref ref-type="bibr" rid="B94">Venil et&#xa0;al., 2014</xref>). Recently, transformations in technology have opened the doors for artificial intelligence and machine learning technologies for their promising use to optimize the fermentation processes (<xref ref-type="bibr" rid="B18">Chong et&#xa0;al., 2024</xref>). Short-term goals can be high-throughput strain screening from diverse environments and habitats to identify hyperproducer strains with stable, vibrant hues suitable for product development. Furthermore, exploration of extremophilic conditions to identify novel bacterial species can account for new pigments with peculiar features such as stability, bioactivity, and multifunctionality. In addition to these approaches, metabolic engineering and synthetic biology can enhance the production of bacterial pigments along with enabling the bacteria&#xa0;for the heterologous production of pigments. Metabolic engineering&#xa0;is useful in improving the precursor supply, reducing/eliminating the competing metabolic pathways, and increasing the availability of cofactors. This approach can also expand the diversity and scale of bacterial pigment production by integrating pigment-producing genes from fungi, plants, and animals. Improving bacterial strains using CRISPR/Cas and other genome editing tools can help eliminate undesired genes and enhance strain efficiency for pigment production. The adoption of a circular bioeconomy model for pigment production, emphasizing the valorization of agricultural and food-industry wastes and other low-cost substrates for bacterial pigment fermentation, can make the process more cost-effective and sustainable (<xref ref-type="bibr" rid="B73">Rajendran et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s6" sec-type="conclusions">
<label>6</label>
<title>Conclusion</title>
<p>The demand for natural cosmetic products has definitely provided a strong foundation for the utilization of bacteria as a source of natural pigments. The multiple health benefits, along with the desirable properties of bacterial pigments for cosmetic products, have already been reported by several research groups. The range of colors produced by these bacteria is also quite broad. However, most of these reports are from research laboratories and are at the very initial stages of investigation. Furthermore, the yield obtained from these microorganisms is also very low. The standardization of process parameters, strain improvement, and the adoption of a sustainable approach for pigment production may provide a boost for industrial-scale recognition and global acceptance of these pigments. The research and development sector is still in a nascent stage for exploring these pigment-producing bacteria.</p>
</sec>
</body>
<back>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>PC: Writing &#x2013; original draft, Formal analysis, Visualization. MK: Writing &#x2013; original draft, Visualization, Formal analysis. BT: Writing &#x2013; original draft, Visualization, Formal analysis. JT: Formal analysis, Writing &#x2013; original draft, Visualization. HS: Formal analysis, Visualization, Writing &#x2013; original draft. DS: Visualization, Conceptualization, Writing &#x2013; review &amp; editing, Formal analysis, Writing &#x2013; original draft, Methodology, Supervision.</p></sec>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
<p>The handling editor BS declared a past co-authorship with the authors PC.</p></sec>
<sec id="s10" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declared that generative AI was not used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p></sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p></sec>
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<fn id="n1" fn-type="custom" custom-type="edited-by">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/388216">Barkha Singhal</ext-link>, Gautam Buddha University, India</p></fn>
<fn id="n2" fn-type="custom" custom-type="reviewed-by">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1783277">Luqman Jameel Rather</ext-link>, Southwest University, China</p>
<p><ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3291268">Dalia Dasgupta Mandal</ext-link>, National Institute of Technology, Durgapur, India</p></fn>
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