Springer Japan
Pigment production by cold-adapted bacteria and fungi: colorful tale of cryosphere with wide range applications

DOI: 10.1007/s00792-020-01180-2, Volume: 24, Issue: 4, Pages: 447-473
Article Type: review-article, Article History

Pigments are an essential part of everyday life on Earth with rapidly growing industrial and biomedical applications. Synthetic pigments account for a major portion of these pigments that in turn have deleterious effects on public health and environment. Such drawbacks of synthetic pigments have shifted the trend to use natural pigments that are considered as the best alternative to synthetic pigments due to their significant properties. Natural pigments from microorganisms are of great interest due to their broader applications in the pharmaceutical, food, and textile industry with increasing demand among the consumers opting for natural pigments. To fulfill the market demand of natural pigments new sources should be explored. Cold-adapted bacteria and fungi in the cryosphere produce a variety of pigments as a protective strategy against ecological stresses such as low temperature, oxidative stresses, and ultraviolet radiation making them a potential source for natural pigment production. This review highlights the protective strategies and pigment production by cold-adapted bacteria and fungi, their industrial and biomedical applications, condition optimization for maximum pigment extraction as well as the challenges facing in the exploitation of cryospheric microorganisms for pigment extraction that hopefully will provide valuable information, direction, and progress in forthcoming studies.



Think for a while, without hemoglobin how oxygen molecules can be carried within a living body or without chlorophyll and other necessary pigments; how plants and other primary producers could prepare their food? Surely impossible. It shows life on this planet depends on pigments. Pigments have been used as coloring agents since the prehistoric era. Sir William Henry Perkin prepared the earliest synthetic dye in 1856 named mauvine. The development of mauvine initiated a historical revolution of synthetic dyes (Walford 1980). Initially, the synthetic dyes gained much attraction due to several characteristics such as easy to develop, economical, no undesirable flavors, excellent coloring properties, and needed in a small amount to use. However, most of the synthetic dyes used had never been tested for their toxic nature for health and adverse effects on the environment (Downham and Collins 2000). In general, synthetic dyes are made up of chemical compounds composed of lead, mercury, copper, chromium, sodium chloride, benzene, and toluene that are harmful to human health. Several synthetic colorants previously permitted by the Food and Drug Administration (FDA) to use in medicines, food, and cosmetics development were later found carcinogenic (Rao et al. 2017). Some of these synthetically-derived colorants banned by the FDA include ethyl acrylate, benzophenone, eugenyl methyl ether (methyl eugenol), pulegone, myrcene, and pyridine. Similarly, several synthetic dyes like cochineal red, tartrazine, and sunset yellow initiate allergies directly or indirectly by combining with other colorants. The Center for Science in the Public Interest in Washington appealed to the FDA in 2008 to disallow synthetic food colorants since of their association with behavioral harms among children (Potera 2010). Most synthetic dyes previously used for various purposes are now quit due to apparent hazards such as carbon black (extensively used as printing ink pigment) a potential carcinogen and benzidine a causative agent of bowel cancer (Rao et al. 2017). Moreover, the unethical release of untreated dye effluents from industries persists for a longer time due to higher stability. Due to the aforementioned drawbacks of synthetic dyes, global demand for natural pigments has been increased (Manikprabhu and Lingappa 2013). Being aware of the adverse effects causing by synthetic dyes, now consumers prefer natural ingredients in their food instead of artificial (Downham and Collins 2000). This has raised the demand for naturally derived pigments.

The use of natural colorants is believed to be relatively safe, since their nature is biodegradable, harmless, and non-carcinogenic (Cristea and Vilarem 2006). Globally, the trend is shifting towards the consumption of biodegradable and environmentally friendly commodities; therefore, the demand for natural colorants is increasing in pharmaceutical, dyestuff, foodstuff, and cosmetics. Waste materials that are environmental pollutants have been used for microbial pigment production, which made this process more sustainable (Joshi et al. 2003; Kamla et al. 2012). Natural biocolorants that are an alternative to synthetic pigments could be obtained from plants and microorganisms. Natural pigments obtained from microbes are more ideal over plants due to the solubility, and stability of pigments and the easy availability of microbes for culturing (Rao et al. 2017). The fast growth of microorganisms in inexpensive medium, easy downstream processing, and their constant cultivation independent of the seasonal variations are advantages of microbes over plants as a source of pigment production (Manikprabhu and Lingappa 2013). Moreover, the use of plants on a large scale can cause damage to the precious species; therefore, the practice is not sustainable (Downham and Collins 2000). Additionally, the yearly growth rate of natural dyes has been projected 5–10% compared to synthetically derived dyes having a lower growth rate of 3–5% (Parmar and Phutela 2015). Pigments produced by microorganisms are not merely colors but consist of diverse chemical components having multidimensional biological activities (Kim 2013). Among microorganisms, bacteria, fungi, and microalgae offer an alternate source for natural pigments (Joshi et al. 2003; Choi et al. 2015; Rao et al. 2017; Pandey et al. 2018; Ramesh et al. 2019).

Earth’s cryosphere has been thought too hostile to harbor life for a long time. However, being extreme ecological conditions, diverse microbial communities exist in the glacial ecosystem (Anesio et al. 2017; Hassan et al. 2018; Rafiq et al. 2019) that maintain active biochemical routes (Anesio and Laybourn-Parry 2012). The microbial communities inhabiting the cryosphere are constantly exposed to several stress conditions such as extremely low temperatures, oligotrophic conditions, freeze–thaw cycles, ultraviolet (UV) radiations, and higher salinity. To cope with these hostile conditions, microorganisms adapt several protective strategies including morphological alteration and production of various metabolites. Most of these metabolites have important biotechnological and industrial applications (Cavicchioli et al. 2011). Among these metabolites, pigments are charismatic traits of these cold-adapted microorganisms that are studied to exploit for several industrial applications. Pigments production mostly occurs within cytoplasm as a responsive agent to hostile ecological conditions presenting several ecological functions (Pagano and Dhar 2015). Physiochemically extreme environments such as the cryosphere provide a suitable setup to microorganisms for pigment production with unique qualities and applications. In this review, pigments produced by cold-adapted bacteria and fungi in the cryosphere and their potential applications in various industries are discussed. Moreover, condition optimization for maximum pigment yield as well as the challenges facing in the exploitation of cryospheric microorganisms for pigment extraction are discussed.

Challenges in cryosphere and microbial adaptation mechanisms

Cryosphere has been dominated and colonized by diverse microorganisms collectively termed as psychrophiles or cold-adapted microorganisms. Life spin and activity of several microorganisms significantly reduce below 0 °C (Tribelli and López 2018). However, psychrophilic and psychrotrophic microorganisms withstand the hostile environments of cryosphere and adapt to grow below 5 °C (close to zero). Low temperature reduces thermal energy with increased viscosity of solvents and solubility of gasses such as oxygen molecules and reactive oxygen species (ROS), reduced solubility of nutrients and solutes, decreased diffusion as well as amplify desiccation, osmotic stress, and ice development. Moreover, life in the cryosphere experience several environmental stresses including high salinity, low availability of nutrients, oxidative stress, low water activity, and freeze–thaw cycles. High-pressure stress is subjected to microbial population inhabiting in deep-sea and sub-glacial habitats. Additionally, psychrophilic microbes are exposed to extremes of light, exposure to UV radiations, and bright light at high elevation and low light in frosted lakes, permafrost, and deeper ice sheets. To cope with these life-endanger challenges microorganisms adapt and develop certain sophisticated strategies (Margesin and Collins 2019; Collins and Margesin 2019). Previously, several microbiological, physiological, biochemical, biophysical, and molecular-based approaches have been adapted to identify biogeographical distribution, physiological adaptation, diversity, and ecological role of psychrophiles. However, recent advances in ‘omics’ technologies such as genomics, metagenomics, transcriptomics, and proteomics have uncovered novel adaptation mechanisms and environmental functions of microorganisms in the cryosphere (Singh et al. 2014; Barauna et al. 2017; Tribelli and Lopez 2018). Several mechanisms adapted by microorganisms to cope with the hostile environment of cryosphere are shown in Fig. 1.

Common strategies adapted by psychrophiles to cope with low temperatures and other stresses in the cryosphere
Fig. 1
Common strategies adapted by psychrophiles to cope with low temperatures and other stresses in the cryosphere

At low temperatures, psychrophiles alter their cellular envelope and components to provide shape, support, and protection to cells; regulate nutrient uptake and solute transport; and participate in cell division, adhesion, signaling, and sensing. This alteration is against the low temperature that reduces membrane permeability, fluidity, diffusion rates, and cause cell rupturing due to freeze–thaw cycles and ice formation. Psychrophiles modify the composition of fatty acid in lipid bilayer present in the cell membranes and become homeoviscous (D’Amico et al. 2001; Siddiqui et al. 2013). Contents of unsaturated fatty acids, short-chain, methyl branched, and/or cis-isomeric can manage the low temperature stress (Hassan et al. 2020) by lowering the liquid phase phospholipid bilayer to gel phase and sustain their function. For this adaptation, upregulation and over-representation of genes occur that encode proteins involved in membrane biogenesis and biosynthesis and desaturation of fatty acids (desaturases that also guard against ROS), biosynthesis of branched-chain fatty acids (KAS-II, KAS-III) and cis-isomerization (fatty acid cis/trans isomerases) (Medigue et al. 2005; De Maayer et al. 2014; He et al. 2015; Goordial et al. 2016). Furthermore, psychrophiles upregulate the membrane transport proteins that could counter the reduced diffusion rates at lower temperatures (Bakermans et al. 2007; De Maayer et al. 2014). Long-chain polyunsaturated fatty acids (LC-PUFAs) (Yoshida et al. 2016) also shield the membrane by generating hydrophobic edges between the lipid bilayers, which prevents the entrance of ROS augmented at low temperatures. Polyunsaturated fatty acids (PUFAs) can act as chaperons for proteins in the membrane, which could be functional in cell division and the efflux process (Okuyama et al. 2008; Yoshida et al. 2016). Turk et al. (2011) reported higher membrane fluidity in psychrotolerant yeast Rhodosporidium diobovatum suggesting that LC-PUFAs maintain the integrity and functionality of the plasma membrane. Similarly, psychrophiles upregulate certain genes encoding peptidoglycan biosynthesis and thicken their peptidoglycan (Mykytczuk et al. 2013), which can protect psychrophiles against freeze–thaw cycle, ice formation, and osmotic pressure. The lipopolysaccharides (LPS) layer in psychrophiles are shortened in length lacking the O-chain component (Corsaro et al. 2017) that may enhance the flexibility and stability of the outer membrane. Genes encoding for the synthesis of outer membrane components, proteins, and LPS (largely glycosyl transferases) was reported upregulated at lower temperatures (Frank et al. 2011; De Maayer et al. 2014). Benforte et al. (2018) reported that how a mutation in the glycosyl transferase gene (wapH) reduced the growth pattern of the Antarctic bacterium at lower temperatures. Several psychrophiles have higher genome contents encoding for compatible solute biosynthesis and uptake genes that could accumulate compatible solutes in molar concentrations, with glycine betaine, glycerol, mannitol, trehalose, sarcosine, sucrose, and sorbitol (Mykytczuk et al. 2013; Ghobakhlou et al. 2015; Goordial et al. 2016) that could restore osmotic balance and cell shrinkage. Fungi synthesize certain compatible solutes to cope with osmotic pressure and dehydration in the cryosphere such as mannitol that acts as a cryoprotective (Feofilova et al. 1994). Psychrophiles produce antifreeze proteins (AFPs) that bind to ice and inhibit ice development and recrystallization. AFPs preserve the liquid present in the close vicinity of cells (Raymond et al. 2008). These AFPs also stabilize cell membranes, structural integrity, and position the cell for increase contact towards nutrients and oxygen in the phototrophic region (Lorv et al. 2014; Bar Dolev et al. 2016; Voets 2017). Another protein called ice-nucleating proteins (INPs) reported in psychrophiles prevent cryoinjury of the cell by inhibiting intracellular ice formation (Lorv et al. 2014). Cold-adapted bacteria produce a high concentration of extracellular polymeric substances (EPS) at low temperatures (Mykytczuk et al. 2013; Caruso et al. 2018) which could form a protective shell around the cells. EPS produced by psychrophilic microorganisms could act as osmoprotective, ROS scavenging, and cryoprotective (Ewert and Deming 2013; Deming and Young 2017). Along with low temperature adaptation, biosurfactant (glycolipid) reported from Antarctic yeast showed ice recrystallization inhibition activity (Kitamoto et al. 2001), also such biosurfactants act as osmolytes (Perfumo et al. 2018). The main adverse effect of low temperature is on the cellular reaction rate of mesophilic microorganisms that either halts their cellular growth or cause death due to cellular lysis. However, psychrophiles produce enzymes that perform high catalytic activities in cold environments, therefore, optimally grow at low temperatures (Santiago et al. 2016; Collins and Gerday 2017). Psychrophiles overcome the obstacle of reduced enzyme activity by developing a changed system for the transportation of nutrients and waste products; altered cellular processes of transcription and translation, and cell division cycle, and decrease the fluidity of membrane to grow in low temperature (D'Amico et al. 2006). Cold-adapted organisms produce chaperones that efficiently fold proteins and DNA/RNA and stabilize the secondary structures of these molecules at low temperatures. Psychrophiles upregulate the production of protein and DNA/RNA chaperones as cold acclimation proteins (Lim et al. 2000). Recently by adapting omics approaches have discovered various traits that are common to psychrophiles (Tribelli and Lopez 2018). One such trait is the metabolic adjustment at sub-zero temperature that takes place in psychrophiles such as the downregulation of primary metabolic pathways and replacement with alternate secondary pathways and accumulations and digestion of spare compounds. Cellular metabolic routes like glycolysis, tricarboxylic acid cycle, electron transport chain, and pentose phosphate pathway get downregulated in cold-adapted microorganisms (Medigue et al. 2005; Piette et al. 2011; Tribelli et al. 2015). Such metabolic reprogramming could protect oxidative stress and conserve energy for long-term survival. Another important protective mechanism adapted by psychrophilic microorganisms is the production of pigments. Psychrophiles reported from ice-cores and glaciers were capable to produce pigments (Shen et al. 2018). Melanin pigment reported in fungi protects stresses such as ionizing radiation, desiccation, oxidizing agents, and UV radiations (Butler and Day 1998; Gorbushina 2003). Antarctic fungal species having melanin were reported, which were resistant to UV radiation (Hughes et al. 2003). Several fungal strains produce mycosporines that absorb UV radiations (Sommaruga et al. 2004). Similarly, psychrophilic bacteria produce carotenoid as a protection mechanism in cryospheric environments (Vila et al. 2019). Moreover, these pigments also play role in photoprotection (in combination with other molecules like mycosporine and scytonemin like amino acids, and act as a protector against intense light range and UV irradiation), act as cryoprotectants, antioxidants, and antimicrobials (Dieser et al. 2010; Pandey et al. 2018).

Role of pigments in cold-adapted microbes

In general, pigments are secondary metabolites produced by microbes and are not necessarily produced by all kinds of microorganisms. Cold-adapted bacteria and fungi inhabiting cryospheric environments produce a diverse range of pigments (Table 1). The pigment production by microbes in extremely low temperature environments is a mechanism to survive ecological stresses. They consume pigment molecules as an energy source (Madigan et al. 2012), for the process of photosynthesis (Siefirmann-Harms 1987), as a confrontation tool against stress (Martin-Cerezo et al. 2015), oxidants, extreme temperature and desiccation (Wada et al. 2013), and for safety against UV irradiation (Becker-Hapak et al. 1997). Moreover, pigments also act as an antimicrobial agent against other bacteria (Suresh et al. 2015) and sometimes act as a shield to protect the cell against natural antimicrobial compounds secreted by other bacteria (van Duin et al. 2002). High diversity of pigment molecules are reported from microorganisms inhabiting in glaciers (Foght et al. 2004), ice cores (Zhang et al. 2008), and marine surface waters (Agogué et al. 2005) which shows that this pigment production is essential to cope with the environmental stresses in the cryosphere. Such as carotenoid modulate the cellular membrane fluidity of bacteria as an adaptive strategy in the cryosphere. Polar microbial communities are concurrently exposed to UV radiations, highly active photosynthetic radiations, and extremely low temperatures which make them more prone to photo-damage (Roos and Vincent 1998). Therefore, the adaptation of protective strategies by microorganisms is crucial in these habitats. In such harsh environmental conditions, pigment production efficiently decreases photo-damage and could provide resources for osmoregulation, thus provide tolerance against salinity, desiccation, and freeze–thaw cycles and prevent serious damages such as inhibition of metabolic processes and cellular repair mechanisms (Mueller et al. 2005). Depend upon the composition and concentration of pigments, the production could visibly provide coloration to snow such as green due to chlorophylls, various yellow shades due to xantophylls, and orange to red due to carotenoids (Anesio et al. 2017).

Table 1
Pigments produced by cold-adapted bacteria and fungi, characterization, and biological activities
Type of microorganism Name of organism Habitat Pigment Color Biological function References
Bacteria Micrococcus roseus Antarctic soil Carotenoid Yellow Regulation of membrane fluidity Chattopadhyay et al. (1997)
Arthrobacter agilis Sea ice, Antarctica Carotenoid Yellow Membrane stabilization at low temperature Fong et al. (2001)
Microbial mat; multiple bacteria isolated Ward Hunt Ice Shelf, Nunavut, Canada Carotenoid, Scytonemins Yellow Protection against reactive oxygen species, coping radiations, and other stresses Mueller et al. (2005)
Multiple bacteria isolated Antarctic sites: Pony Lake and Cotton Glacier Carotenoid Yellow, orange, and dark rose red Prevent against freeze–thaw cycles. Protection against UV radiation Dieser et al. 2010
Sanguibacter suarezii KK6, Kocuria turfanensis KK7, Kocuria rosea KK12, and Planococcus maritimus KK21 Leh and Ladakh soil, Himalayas Carotenoid (Lycopene) Yellow and orange Survival strategy in a cold environment Kushwaha et al. (2014)
Arthrobacter isolates Scott Base, Granite Harbour, Minna Bluff and Marble Point (Antarctic) Carotenoid Genes reported Cold adaptation by stabilizing cell membrane, resistance to oxidative stress Dsouza et al. (2015)
30 different heterotrophic pigmented bacteria were isolates Fildes Peninsula, King George Island, Antarctica Carotenoid Yellow and orange Survival strategy in a cold environment Vila et al. (2019)
Novel species very close to Janthinobacterium lividum Organic residue of a water tank keeping rainbow trout Mixture of violacein and deoxyviolacein Violet Antibacterial activity and enhance competitive ability Nakamura et al. (2002)
Corynebacterium glutamicum ATCC ATCC


(Carotenoid); Lycopene


Yellow and red Photoprotection and light-harvesting Heider et al. (2012)
Sphingomonas faeni and other Anthropogenic cold environments Astaxanthin (Carotenoid) Yellow Antagonistic effect against competitors Mageswari et al. (2015)
Janthinobacterium sp. HHS7 Xinjiang glacier, China Violacein Violet Adaptation to extremely low temperature Lu et al. (2009)
11 psychrophilic Flavobacterium strains Laigu, Zepu, Renlongba, and Gawalong glaciers in Bome County, Tibetan Autonomous Region, P.R. China Flexirubin-type pigment Yellow Adaptation to extremely low temperature Liu et al. (2019)
Pseudomonas. Cryobacterium, Psychrobacter, Flavobacterium, and many others Yuzhufeng Glacier, Tibetan Plateau, China carotenoids such as α- and β-carotene, 19′-butanoyloxy-fucoxanthin, fucoxanthin, diatoxanthin, peridinin and zea/lutein, with α-carotene as the dominant carotenoid (orange, reddish-orange, yellow, pink, brown, and white Survival strategy in a cold environment Shen et al. (2018)
Genus Arthrobacter; Arthrobacter psychrophenolicus sp. nov Alpine ice cave, Austria Unidentified Yellow Survival strategy in a cold environment Margesin et al. (2004)
Thirty-seven heterotrophic bacterial isolates

Franz Josef and Fox

Glaciers, New Zealand

Unidentified Yellow and orange Survival strategy in a cold environment Foght et al. (2004)
Multiple bacteria isolated Prydz Bay, Eastern, Antarctica Unidentified Yellow, red, orange, pink Survival strategy in a cold environment Bowman et al. (1997)
Multiple bacterial isolates are reported Gulkana Glacier, Alaska Unidentified Pink, orange, white, cream, and yellow Resistance and adaptation to cold temperatures Segawa et al. (2011)
Multiple bacterial isolates are reported Siachen Glacier, Pakistan Unidentified Several colors pigments Adaptation against cold temperature Rafiq et al. (2017)





Himalayan Glaciers in Uttarakhand, India Unidentified Lemon yellow, orange, brown, violet, pinkish-red and pale yellow Adaptation against cold temperature Panwar et al. (2019)
Janthinobacterium sp. strain Ant5-2 Lake Podprudnoye, Schirmacher Oasis, East Antarctica Violacein Purple violet photoprotective role. UV and cold tolerance Mojib et al. (2013)
total of 31 g-positive bacteria was reported King George Island, Antarctica Peninsula) Unidentified Yellow, red, orange, and amber inhibit potential surface competitors Leiva et al. (2015)
Lysobacter oligotrophicus Antarctica (Lo-melanin) melanin Dark-brown Protection against UV-B irradiation. Scavenge ROS Kimura et al. (2015)
Hymenobacter actinosclerus and Chryseobacterium chaponense King George Island, Antarctica Carotenoids Yellow and red tolerate excessive UV irradiation Órdenes-Aenishanslins et al. (2016)
Pedobacter terrae Doumer Island, Antarctica Different pigments belong to carotenoids group Red Antioxidant activity, protection against UV radiation Correa-Llantén et al. (2012)
Multiple isolates were reported coastal Antarctic Station Dumont d’Urville, European alpine snow pits, Nevado Illimani, Bolivia, and Antarctic Station Artigas, King George Island Unidentified Multiple colors Uv radiation protection González-Toril et al. (2008)
Leeuwenhoekiella aequorea, Pseudomonas pelagia, Halomonas boliviensis, Rhodococcus yunnanensis, Algoriphagus ratkowskyi Kongsfjorden, a glacial fjord on the west coast of Svalbard, (Zeaxanthin) carotenoid Yellow and red cryoprotective agent in regulating membrane fluidity Singh et al. (2017)
Kocuria polaris sp. nov McMurdo Dry Valley, Antarctica Carotenoid Orange Reddy et al. (2003)
Arthrobacter sp. G20 Caspian Sea Carotenoid Red Afra et al. (2017)
Cyanobacterial mats Antarctic stromatolitic and endolithic cyanobacterial communities Scytonemin Photoprotection Wynn-Williams et al. (1999)
Lichen and cyanobacterial samples Antarctic desert habitats, β-carotene, Scytonemin, phycocyanin, etc UV protection Wynn-Williams and Edwards (2002)
Cyanobacteria-dominated microbial mats (several isolates) McMurdo Sound region, Antarctica Scytonemin, carotenoids, Multiple colors Vincent et al. (1993)
Fungi Several yeasts were isolated Antarctica (Admiralty Bay, King George Island, and Port Foster Bay and Deception Island)

Carotenoids and

Other pigments

Several colors Protection against UV radiation Vaz et al. (2011)

Dioszegia sp.,

Rhodotorula mucilaginosa, Rhodotorula laryngis

Italian alpine glacier (Tonale Pass, TN, Italy) Carotenoids Orange to salmon-colored Protection against UV radiation Amaretti et al. (2014)
Thelebolus microsporus McLeod Island, Larsemann Hills, in the Prydz Bay area of East Antarctica



Bright orange to yellow-orange Stress tolerance of UV radiation and extreme cold Singh et al. (2014)



King George Island, Antarctic Peninsula Astaxanthin, phoenicoxanthin, β-carotene, (Carotenoid) red, pale-yellow and yellow photoprotective role Contreras et al. (2015)
Penicillium sp. (GBPI_P155) Himalayan region of India



Dark orange pigment Survival of the cold temperature Pandey et al. (2018)
Multiple fungal isolates were reported Batura Glacier, Karakoram, Pakistan Unidentified Multiple colors Adaptation against cold temperature Hassan et al. (2018)
Multiple endophytic fungal isolates were reported Admiralty Bay, King George Island, South Shetland Islands, Antarctica Melanin Dark blackish Protection against environmental stresses in cold habitats Rosa et al. (2009)
Antarctomyces pellizariae sp. nov., Coppermine Peninsula, Robert Island, in the South Shetland Islands, Antarctica Unidentified Blue Cryo-protection of fungus de Menezes et al. (2017)
Cystofilobasidium capitatum and Sporobolomyces ruberrimus Zooplankton (Antarctica) β-Carotenoid Resistance to ultraviolet B (UVB Moliné et al. (2009)
Sporobolomyces salmonicolor, Cryptococcus albidus, and Cryptococcus laurentii Livingston Island, Antarctica β-carotene Resistance to ultraviolet A (UVA) Dimitrova et al. (2010)
Sporobolomyces salmonicolor Livingston Island, Antarctica β-Carotene, torularhodin, and torulene Antioxidant activity Dimitrova et al. (2013)
Rhodotorula mucilaginosa, Rhodotorula larynges, Dioszegia sp. (King George Island, Antarctica 2-γ-Carotene, Torulene, γ-carotene, and lycopene, Torulene, and lycopene, OHK torulene Pink, pale red, red, orange UV-C radiation tolerance Villarreal et al. 2016
Nadsoniella nigra Antarctica Melanin Black Increase resistance in piglets and reduction of morbidity and mortality Chyizhanska and Beregova (2009)
Several yeasts strains Antarctic biotopes Melanin Coal-black Tashirev et al. (2010)
Cryptococcus sp. and Torrubiella sp Sedimentary rocks (Union Glacier) Antarctic Mycosporine Pink and cream Protection against UV radiations Barahona et al. (2016)
Dioszegia patagonica (King George Island Carotenoid Orange Trochine et al. (2017)
Arthrobotrys ferox Moss samples (Wood Bay, Victoria Land Antarctica Carotenoid and mycosporines Resistance to ultraviolet-B (UVB Arcangeli et al. (1997, 2000); Arcangeli and Cannistraro (2000)
6 yeast strains and 11 bacterial strains Galindez Island in the Argentine Islands in the north-western Antarctic Peninsula Carotenoids and other pigments Multiple colors UV resistance Vasileva-Tonkova et al. (2014)

Pigments produced by cold-adapted bacteria and fungi


Carotenoids are the main natural pigments widely produced by plants and microorganisms and initially were reported by H.W.F Wackenroder (Wackenroder 1831). Currently, carotenoids signify the largest and highly diverse known group of natural pigments, and 1183 carotenoid structures are compiled from 702 source organisms by Carotenoid Database Japan (https://carotenoiddb.jp). Carotenoid is the family of tetraterpenoids consists of an extensively conjugated polyene, which absorbs blue light and appears from yellow to red in shade (Fraser and Bramley 2004). The structure of carotenoids acquires the form of a polyene hydrocarbon chain that is occasionally terminated by rings with or without extra oxygen atoms. Carotenoids are classified into carotenes, which are hydrocarbons (torulene, lycopene, and α- to ε-carotenes) or xanthophylls that possess keto, hydroxyl, and/or carboxyl groups (lutein, zeaxanthin, torularhodin, and astaxanthin,) (Table 2). Naturally, carotenoids are extensively produced by plants and microorganisms as a photo-protectants. The color of carotenoids ranging from yellow to deep red depending on the structure (Alija et al. 2005; Rao and Rao 2007). The maximum absorption capacity of carotenoids is ranged from 440 to 520 nm, having a stronger molar absorption coefficient (ca. 105 L mol−1 cm−1 ) (Marizcurrena et al. 2019). Cryospheric habitats especially glacial environment provide suitable conditions for pigment production by microorganisms. These pigments also protect the bacterial cells either screening or absorbing UV radiation (Dieser et al. 2010). Carotenoids safeguard microbial cells from photo-oxidative injury and other environmental stresses at low temperatures either by inhibiting the ROS generation through thermal degeneracy of the extra energy or satisfying the excited states of singlet oxygen and chlorophyll (Tian and Hua 2010). Moreover, these pigments play a crucial role in the physiological plasticity of Antarctic microorganisms, efficiently respond to a lower temperature and freeze–thaw cycles (Singh et al. 2017). Additionally, carotenoids play a role in cell differentiation and regulation during cell cycles.

Table 2
Types of pigments from cold-adapted microorganisms, their applications, and chemical structures
Pigments Applications Structures



Food additives, Antioxidant activities, Antimicrobial activities, and as Sun protector

C40H56 (psi,psi-Carotene)



Antioxidant activity, Cosmetics additives, Anti-cancerous activities, and Antimicrobial activities

C40H54 (3′,4′-Didehydro-beta,psi-carotene)



Food colorant, Antioxidant activity, and Precursor of vitamin A in food

C40H56 (beta-Carotene)



Feed additives, Protection against

free radicals, and use in the pharmaceutical industry



Prodigiosin Dyeing agent in the textile industry, Coloring agents in the food industry, Antibacterial antiviral, and anticancer activities

C20H25N3O (4-Methoxy-5-[(Z)-(5-methyl-4-pentyl-2H-pyrrol-2-yliden)methyl]-1H,1′H-2,2′-bipyrrol)

Melanin Antibacterial activity against antibiotic-resistant pathogens, Antioxidant activity, and cytotoxic activity

C18H10N2O4 (6,14-dimethyl-4,12-diazapentacyclo[,5.013,17.09,18]octadeca-1(17),2,5,9(18),10,13-hexaene-7,8,15,16-tetrone)

Pheomelanin Markers in fossils, Antioxidant activity, Photoprotective effects

C34H29N6O12S3 (Pheomelanin)

Eumelanin Antioxidant activity, Biomedical activities

C25H13N3O13 (Eumelanin)

Violacein Dyeing agent in the textile industry, Food industry, Antibacterial, fungicidal and antiviral activities, Antioxidant and cytotoxic activities, Use in cosmetics and medicine

C20H13N3O3 (3-[2-hydroxy-5-(5-hydroxy-1H-indol-3-yl)-1H-pyrrol-3-yl]indol-2-one)

Indigoidine As bioindicator, Antimicrobial activities, and the Textile industry as a dyeing agent

C10H8N4O4 (3-(5-amino-2-hydroxy-6-oxo-1H-pyridin-3-yl)-5-iminopyridine-2,6-dione)

Scytonemin Anti-inflammatory and Anti-proliferative drugs, Antioxidant agents, Use in cosmetics

C36H20N2O4 (3E,3′E)-3,3′-Bis(4-hydroxybenzyliden)-1,1′-bicyclopenta[b]indol-2,2′(3H,3′H)-dion)

Several studies reported carotenoid producing bacteria and fungi from low temperature environments (Table 1). Vila et al. (2019) reported 30 bacterial strains capable of carotenoids production from Fildes Peninsula, King George Island, Antarctica. Kim et al. (2015) isolated carotenoid producing Planococcus faecalis from stools of Antarctic penguins. Carotenoid production was reported in heterotrophic bacteria from Antarctica as a strategic tool against ecological stresses (Dieser et al. 2010). The accumulation of C-50 carotenoid (Bacterioruberin and its glycosylated derivatives) observed in psychrotrophic Arthrobacter agilis from an Antarctic sea ice sample. This accumulation of pigments might be responsible for the regulation of cellular membrane fluidity at low temperatures (Fong et al. 2001). Moreover, several polar and non-polar carotenoid pigments produced by Antarctic strains of Sphingobacterium antarcticus and Micrococcus roseus (Chattopadhyay and Jagannadham 2001). Silva et al. (2019) reported carotenoid producing bacterial strains isolated from the Antarctic continent. Three different carotenoids such as zeaxanthin, β-cryptoxanthin, and β-carotene were reported from Antarctic bacterium Sphingobacterium antarcticus. Similarly, carotenoid producing cold-adapted Penicillium sp was reported in the Himalaya region of India (Pandey et al. 2018). Several carotenoid-producing yeasts were reported in Italian alpine glaciers (Amaretti et al. 2014). Vaz et al. (2011) reported multiple yeasts isolates from Antarctica capable of carotenoid production.


Prodigiosin is a red linear tripyrrole pigment primarily reported from Serratia marcescens (Boger and Patel 1987). Prodigiosin was named after its extraction from Bacillus prodigious later given the name of S. marcescens (Gerber 1975). Prodigiosin produces only in later growth stages of bacteria (Harris et al. 2004). Biosynthesis of prodigiosin is controlled by quorum sensing (Thomson et al. 2000). Certain proposed eco-physiological roles of prodigiosin are; air diaspora of bacteria (Burger and Bennett 1985), metabolic precursor for NAD(P)H or proline (Hood et al. 1992), light energy storage (Ryazantseva et al. 1995), ion exchange (Seganish and Davis 2005), energy spilling function in S. marcescens (Haddix et al. 2008) and acts as an antimicrobial agent to provide a competitive advantage within communities (Starič et al. 2010). Borić et al. (2011) studied that prodigiosin is protective against UV radiation in Vibrio sp. DSM 14379. Prodigiosin was extracted from psychrotrophic bacterial strain Janthinobacterium lividum isolated from Alaskan soil (Schloss et al. 2010). Shen et al. (2018) reported several red pigment-producing bacterial strains inside deep ice core collected from the Yuzhufeng Glacier, Tibetan Plateau. Centurion et al. (2019) reported genes in Antarctic volcanic island sediments responsible for the biosynthesis of 3-oxoacyl-[acyl-carrying protein] reductase (K00059) enzyme that belongs to the fatty acid synthesis pathway type II associated to the production of prodigiosin.


Melanin is dark in color (brown to dark green, or fully black) and higher molecular weight biological pigment found in hair, feather, skin, eyes, scales, and some interior membranes. Melanin is chemically a polymerized product of phenolic and/or indolic compounds (Tarangini and Mishra 2014). Melanin is further classified into three groups based on structure and color (i) pheomelanins (red or yellow), (ii) eumelanins (black-brown), and (iii) allomelanins (black to dark brown) (Table 2). Due to variations in the occurrence and structure of melanin, its biosynthesis is not from a single route (Solano 2014). Melanin synthesis has been associated in providing resistance against UV- and visible light-irradiations, confrontation the attack of cell wall enzymes, fortification against oxidizing and reducing agents, and acts as an antiviral agent to enhanced the competitive and survival abilities in environmental stresses (Castro-Sowinski et al. 2007; Solano 2014). Melanin is a strong absorber of UV radiations and provides strong protective functions to microbes that’s why extreme low temperature ecosystems are suitable habitats for microbial biosynthesis of melanin (Gessler et al. 2014).

In general, melanin pigment is insoluble both in organic and aqueous solvents, however, Kimura et al. (2015) isolated bacteria strain Lysobacter oligotrophicus from the Antarctic environment that produced water-soluble heteropolymer (Lo-melanin). Lo-melanin protects against UV radiation and scavenges ROS. Melanin is commonly present in polar dark septate hyphae, and guards hyphae against low temperatures and play a crucial role in their persistence (Robinson 2001). Chyizhanska and Beregova (2009) isolated melanin from Antarctic yeasts. An Antarctic fungus, Friedmanniomyces endolithicus produces highly melanized thick-walled cells that protected cells from UV radiation (Onofri et al. 2004). Similarly, Rosa et al. (2009) isolated melanin-producing endophytic fungal strains from the leaves of Deschampsia antarctica and about 80% were black molds. In the Antarctic rocky deserts, rock-inhabiting fungi produce melanin pigments that protect the cells from extreme cold and heat, polychromatic UV radiations, extreme pH and osmotic conditions, and provides tolerance towards potentially toxic metals (Selbmann et al. 2015). Melanin was purified from cold-tolerant and oligotrophic bacterium L. oligotrophicus isolated from the Skarvsnes region, East Antarctica (Fukuda et al. 2013). UV-B radiations induced melanogenesis in L. oligotrophicus cells, that protected the suspensions of E. coli DH5α cells from UV radiations by melanin solutions. In short, melanin biosynthesis by microorganisms can play crucial roles in low temperature environments such as to cope with low temperature, protection against UV radiations, freeze–thaw cycling and desiccation, scavenging ROS. These microbes could be a microbial model for exobiological/astrobiological studies.


Violacein is violet or purple bisindole water-insoluble pigment and first reported from Chromobacterium violaceum from the Amazon River in Brazil (Durán et al. 1953). In nature, violacein protects cells from UV radiations (Füller et al. 2016). Several studies have reported the antibacterial, antiviral, anticancer, antiulcerogenic, antileishmanial, and enzymatic modulation activities of violacein pigments (Duran et al. 2007; Soliev et al. 2011). The maximum UV absorption capacity of violacein is λ  = 260 nm, which suggests its crucial role in the protection of cells from visible and UV radiations (Marizcurrena et al. 2019). In nature, violacein is associated with biofilm production (Pantanella et al. 2007) and its production is regulated by quorum sensing thus acts as a marker of quorum sensing molecules (Burt et al. 2014). Violacein plays a vital role in protecting bacterial cells from predation (Choi et al. 2015). Apart from several other habitats, violacein has been reported from several low temperature environments. Lu et al. (2009) reported a novel psychrophilic bacterium from Xinjiang glaciers, China, capable of violacein production. Violacein producing psychrophilic Janthinobacterium svalbardensis bacterium was isolated from Glacier, Spirsbergen, Arctic region (Ambrožič et al. 2013). Kim et al. (2012) reported violacein producing psychrophilic bacterium Janthinobacterium sp. from alpine glacier cryoconite. Hakvag et al. (2009) reported violacein producing Collimonas sp. from Arctic coastal water in Trøndelag, Norway. Similarly, Nakamura et al. (2002) reported violet pigment-producing psychrotrophic bacterium from the water tank keeping rainbow and exploited its pigment for antibacterial potential. Shivaji et al. (1991) isolated the bacterium of Janthinobacterium genus that produced violacein and further Durán et al. (2007) evaluated the pigments for several therapeutic activities. Mojib et al. (2010) extracted violacein-like purple violet pigment from Janthinobacterium sp. reported from Proglacial Lake Podprudnoye, Schirmacher Oasis, Antarctica that inhibited the growth of Mycobacterium smegmatis and Mycobacterium tuberculosis. At low doses, the same pigment showed activity against methicillin-resistant and multiple drug-resistant clinical strains of Staphylococcus aureus (Huang et al. 2012).


Indigoidine is a brilliant blue, water-soluble pigment synthesized by very few microorganisms (Sutthiwong et al. 2014) namely Erwinia chrysanthemi (Reverchon et al. 2002), Phaeobacter sp. (Cude et al. 2012), Streptomyces chromofuscus (Yu et al. 2013) and Vogesella indigofera (Day et al. 2017), and the biological and environmental role of indigoidine is unclear, however, it has been described that this pigment could protect against oxidative stress (Reverchon et al. 2002). Indigoidine also possesses antimicrobial activities as stated in Leisingera isolates (Gromek et al. 2016). Consequently, microbes producing indigoidine could have advantage of competition in the environment due to the antibiotic and antioxidant properties of indigoidine. Furthermore, indigoidine acts as intracellular signaling molecules related to motility (Reverchon et al. 2002; Cude et al. 2012). Additionally, indigoidine provides adaptability to microbial cells such as Vogesella sp. reported from Andean Patagonia in iron-rich environments (Day et al. 2017). Several isolates of Antarctic genus Arthrobacter were reported to produce indigoidine (Sutthiwong et al. 2014). Arthrobacter genus (family Micrococcaceae) are more common in Antarctic surroundings such as sediments and soils (Reddy et al. 2003; Dsouza et al. 2015). Bacteria belong to this genus produce different pigments including blue indigoidine (Sutthiwong et al. 2014). Kobayashi et al. (2007) reported indigoidine production from Shewanella violacea DSS12 a piezophilic and psychrophilic bacterial strain isolated from deep-sea sediments of the Ryukyu Trench. Liao et al. (2019) studied the multipartite genomes and sRNome of Arctic Pseudoalteromonas fuliginea BSW20308 and reported upregulated genes encoding indigoidine biosynthetic.


Scytonemin is a secondary metabolite of small hydrophobic alkaloids having yellowish-brown color (Table 2). Cyanobacteria when get exposed to UVA-blue wavelengths, scytonemin was produced (Fleming and Castenholz 2007). Scytonemin and its methoxylated and methylated derivatives are mostly reported in the upper portion of the microbial mats; that could protect the cells from extreme environments, which makes this pigment a potential biomarker molecule for studying the presumed exobiology habitats. Exposure of cyanobacteria to UV radiation produces scytonemin along with several other protective metabolites (Sinha et al. 1998). Therefore, these molecules might have performed a crucial role in Antarctica during the initial life stages on this planet (Dillon and Castenholz 1999). The palaeolimnological investigation that was carried out in 62 east Antarctic lakes revealed that the composition of pigments was different in different depths of lakes and higher scytonemin was reported in shallow lakes. Since cyanobacteria regulate their pigment production, this difference of pigment contents could be due to the light-harvesting ability of pigments at different depths (Sabbe et al. 2004). Several Antarctic cyanobacterial genera that are capable to produce scytonemin are reported in Table 1.

Applications of microbial pigments

Versatile pigments produced by microorganisms are more focused during present research due to their extensively observed applications in textile industries, food industries, and biomedical purposes. Global trade of naturally derived pigments has been increased by 29% and reached 600 million USD from 2007 to 2011 (Tuli et al. 2015). This indicates that the pigments produced by microorganisms would dominate the organic market and pigment industries very soon (Ramesh et al. 2019). Such economic growth and vast applications of pigments rationalize the exploration of new sources of natural pigments. Cryospheric environments might provide the best source for the exploration of novel pigments producing microorganisms. Biotechnological applications of psychrophilic microorganisms remained a dream for decades. However, in the early 1990s researchers started focusing their possible usefulness in genetic bioengineering, food and mining industries (Gounot 1991). Currently, several of these expectations are being confirmed, since psychrophilic microorganisms and their secondary metabolites are now acting as sustainable and priceless resources for biotechnological development. Among other invaluable products, pigments produced by these psychrophilic microorganisms are the center of focus in modern research. Table 1 illustrates the microorganisms isolated from cryospheric environments producing pigments of potential applications in industries (Fig. 2). General applications of microbial pigments are as following.

Schematic representation of the pigments applications obtained from cold-adapted microbes
Fig. 2
Schematic representation of the pigments applications obtained from cold-adapted microbes

Industrial applications

Textile industry

Textile industries use 1.3 million tons of synthetic dyes and precursors (worth of 23 billion USD) 15% of which leak as effluents during their use (Venil et al. 2013; Chequer et al. 2013). Unluckily, enough portion of these dyes escape the conventional processes of wastewater treatment and exist as a potentially toxic environmental pollutant with adversities on public health and environment (Ogugbue and Sawidis 2011). Consequently, a great concern exists to replace the synthetic dyes in the textile industry with environmentally friendly dyes. Pigments obtained from microorganisms are environmentally friendly that is considered appropriate for the textile industry (Chadni et al. 2017). Violacein, a multipurpose pigment extracted from C. violaceum is capable of dyeing both natural and synthetic fibers and has gained increasing importance in textiles (Wan et al. 2014; Venil et al. 2016). Prodigiosin a bright red pigment from Vibrio sp. is suitable for dyeing a range of fibers including nylon, acrylics, wool, and silk (Alihosseini et al. 2008). Similarly, prodigiosin reported from Serratia sp. BTW J8 was demonstrated to color various fabric types such as chiffon, cotton, pure silk, poplene, century cotton, organdi, dupoil silk, terrycotton, polyester, and nylon (Krishna 2008). The blue pigment indigoidine produced by fungal host Rhodosporidium toruloides possesses potential applications in the dye industry (Wehrs et al. 2019). Pigment extracted from J. lividum shows a bluish-purple color tone on cotton, silk, and wool, while with vinylon and nylon it appears dark blue (Shirata et al. 2000). Likewise, the yellow pigment extracted from Thermomyces exhibited a higher dyeing affinity for silk fabric compared to other fabrics (Poorniammal et al. 2013). Similarly, red and deep blue pigments reported from Streptomyces strains NP2 and NP4 exhibited substantial variations in dyeing ability depending upon fabric materials. For instance, acrylic and polyamide fibers were vibrantly stained, however, cellulose and cotton fibers were weakly stained (Kramar et al. 2014). In the view of such extensive use of microbial pigments in the textile industry could increase their market value; therefore, new microbial sources are crucial to explore for pigment production and cryosphere inhabit broad microbial diversity capable of pigment production.

Food industry

One important goal in the food industry is to develop foods having an attractive appearance that has been achieved with synthetic dyes. Progressively, food industries are now using natural food colorants, since certain synthetic food colorants have demonstrated adverse health problems after consumption. Due to the shortage of natural food additives resources, their demand is increasing particularly in the food industry (Aberoumand 2011). This demand can be fulfilled by providing a more natural and healthy means of food colors offering a clean and safe tag declaration. Scientists have isolated food-grade pigments from bacterial strains that might provide a natural food colorant having tremendous stability, health safety profile and also act as antioxidants and preservatives (Nigam and Luke 2016). Pigment approval as food color and nutritional supplements greatly depend upon the health safety of consumers and product freshness. Thus, the use of pigmented molecules in food, cosmetics, medicine, and other medical devices is under the control of the Federal Food, Drug, and Cosmetic Act (Chapter VII, Sect. 721) and authorization should be taken before use for the specific time (https://www.fda.gov/).

Several microbial pigments like astaxanthin from Xanthophyllomyces dendrorhous, a red pigment obtained from Monascus sp., β-carotene produced by Blakeslea trispora, lycopene from Fusarium sporotrichioides and Erwinia uredovora, riboflavin reported from Ashbya gossypii, and Arpink red™ from Penicillium oxalicum were supplemented in different food items to enhance its appeal (Dharmaraj et al. 2009). For instance, carotenoids obtained from bacteria are used in feed as pro-vitamins and coloring agent in food industries (Nelis and De Leenheer 1991). Carotenoids are studied to be used as coloring agents for soft drinks, cooked sausages, and baked items (Konuray and Erginkaya 2015). It can also protect food from intense light and maintain the food quality by acting as a sunscreen (Chattopadhyay et al. 2008). Spray-dried prodigiosin from S. marcescens has been applied effectively as a coloring agent in milk, yogurt, and carbonated drinks (Namazkar and Ahmed 2013). Violacein obtained from bacterial sources has been successfully used in food industries (Dufosse 2018). Canthaxanthin is used in food items such as fish, candy, cheese, fruits, meat, snacks, beverages, wine, and beer. Currently, several pigments obtained from microorganisms are approved and used in foods for several purposes (Nigam and Luke 2016).

Biomedical applications

Antimicrobial activities

Infectious diseases are the second major reason for global human deaths and third in developed countries after non-contagious diseases (WHO 2018). In the recent few decades, new drug entry in the market has been decreased and microbial resistance against antibiotics is increasing. This increasing trend of microbial antibiotic resistance amplified the demand for novel antimicrobial agents. As an alternative to antibiotics, several microbial pigments have been successfully evaluated for antimicrobial activities. Umadevi and Krishnaveni (2013) studied pigments extracted from Micrococcus luteus that exhibited inhibitory potential against wound pathogens such as Pseudomonas sp. Klebsiella sp. and Staphylococcus sp. Carotenoids extracted from Holomonas sp. exhibited antimicrobial potential against antibiotic-resistant S. aureus, Klebsiella sp., and Pseudomonas aeruginosa, and ophthalmic S. aureus, Streptococcus pyogens, and E. coli (Ravikumar et al. 2016). Vasanthabharathi et al. (2011) studied the inhibitory potential of crude melanin obtained from Streptomyces against E. coli. Pigment extracted from Streptomyces hygroscopicus was effective against pathogens such as vancomycin and methicillin-resistant S. aureus, β-lactamase producing E. coli, P. aeruginosa and Klebsiella sp. (Berlanga et al. 2000; Selvameenal et al. 2009). Violacein obtained from C. violaceum and J. lividum showed promising antibacterial activities (Durán et al. 2007). Violacein reported from C. violaceum ATCC 12472 showed tremendous antimicrobial activity against S. aureus and inhibition of its biofilm formation (Batista and da Silva Neto 2017). Similarly, Antarctic bacterial violacein in lower concentrations showed activity against avirulent M. tuberculosis (Mojib et al. 2010). A combination of violacein with antibiotics such as kanamycin and cefadroxil minimize the inhibitory concentration of drugs against S. aureus (Subramaniam et al. 2014). Additionally, violacein extracted from J. lividum exhibited fungicidal activity against white root rot causing Rosellinia necatrix (Shirata et al. 1997). Prodigiosin obtained from Vibrio ruber DSM 14,379 exhibited bacteriostatic activity against E. coli and Bacillus subtilis (Danevčič et al. 2016). Furthermore, prodigiosin intracellularly produced by S. marcescens IBRL USM 84 inhibited the growth of tested bacteria (Ibrahim 2008). Prodigiosin revealed promising antibacterial activity against pathogens such as S. aureus, S. pyogenes, P. aeruginosa and Klebsiella pneumoniae (Nwankwo et al. 2017). Pigments extracted from endophytic fungi Monodictysc astaneae demonstrated antibacterial activity against Vibrio cholera, K. pneumoniae, and S. aureus (Visalakchi and Muthumary 2010). Suryawanshi et al. (2017) studied the strong antimicrobial activity of prodigiosin against E. coli, S. aureus, and Candida albicans. Similarly, several prodiginine compounds are reported having fungicidal activities against fungi such as Penicillium, Aspergillus, Candida, and Cryptococcus sp. (Stankovic et al. 2014).

Throughout human history, several viral outbreaks occurred and still occurring such as the Western African Ebola virus epidemic and the recent Coronavirus pandemic COVID-19 accompanied with moderate to high mortality rate. This urge that the discovery of novel antiviral drugs is critically important as many viral infections lack vaccines and effective treatments. Researches showed some initial successes using microbial pigments as antiviral agents. Violacein possesses antiviral potentials against poliovirus, simian rotavirus SA II, and herpes simplex virus (HSV) (Durán et al. 2007). In vitro effects of violacein are also evaluated against Acquired Immuno Deficiency Syndrome (AIDS) related lumphoma (Duran et al. 1996). Andrighetti-Fröhner et al. (2003) studied the antiviral activities of bacterial extracted violacein and inhibition of HSV-1 and Poliovirus-2 (PV-2) replication was observed. Similarly, an in-silico examination of prodigiosin by targeting proteins of Human Immunodeficiency Virus (HIV), Hepatitis B virus (HBV), influenza A virus (H1N1) and Hepatitis C virus (HCV) was performed (Suba et al. 2013). The outcome proposed potential antiviral activities of prodigiosin pigment against all tested viruses excluding HCV, where no binding interactions with active sites were reported. Moreover, compounds of quinone such as naphthoquinones, anthraquinones, and benzoquinones possess strong antiviral potential (Koyama 2006; Gessler et al. 2013). Several compounds obtained from Phoma species exhibited the inhibition of HIV integrase (Rai 2009). Phenazine compounds extracted from Streptomyces and Pseudomonas species have been demonstrated promising antiviral activities (Schneemann et al. 2011). Therefore, microbial pigments are potential agents to administered as a novel source of medications against pathogens.

Antioxidants activity

The rise of free radicals inside the body increases the risks of chronic diseases such as diabetes, autoimmune disorders, and cancer (Phaniendra et al. 2015). To evade this, antioxidant compounds are used which donate electrons to free radicals and neutralize them to protect cellular damage (Lobo et al. 2010). Antioxidants are obtained either from natural or synthetic sources, however, synthetic antioxidants are losing demand due to possible side effects on the human body (Ahmed et al. 2013). Therefore, microbial-based antioxidants are gaining ground in the pharmaceutical industry. Pigments from microorganisms such as carotenoid, xanthomonadin, and naphthaquinone showed antioxidant potential (Tuli et al. 2015). Carotenoids extracted from Antarctic bacterium Pedobacter exhibited solid antioxidant activity with protection against oxidative harm (Correa-Llantén et al. 2012). Osawa et al. (2010) demonstrated promising antioxidant abilities of rare C-50 carotenoids such as sarcinaxanthin monoglucoside, sarcinaxanthin diglucoside, and sarcinaxanthin, reported from a halophilic bacterium Micrococcus yunnanensis . Moreover, Shindo and Misawa. (2014) extracted rare and new carotenoids from the novel bacterial isolates and these pigments exhibited strong antioxidant activities. Similarly, flexirubin (red carotenoid), obtained from Fontibacter flavus showed antioxidant activity against 2, 2-diphenyl-1-picryl-hydrazyl-hydrate (DPPH), nitric oxide, hydroxyl radical, and inhibition of lipid peroxidation (Prabhu et al. 2013). Additionally, melanin reported from fungal isolate Streptomyces glaucescens (El-Naggar and El-Ewasy 2017) and anthraquinones obtained from endophytic Stemphylium lycopersici (Li et al. 2017) were characterized as antioxidants. Another valuable bacterial pigment violacein obtained from C. violaceum exhibited antioxidant protection in gastric ulceration (Antonisamy and Ignacimuthu 2010). Violacein can also safeguard the cellular lipid membranes from peroxidation due to hydroxyl radicals (Stafsnes and Bruheim 2013). Several studies reported melanin obtained from Pseudomonas sp, S. glaucescens NEAE-H, and Bifidobacterium infantis were found antioxidant agents (Huang et al. 2011; Tarangini and Mishra 2014; Zerrad et al. 2014). These reports recommend that pigments obtained from microorganisms could be used as antioxidant agents to prevent several chronic diseases.

Anticancer activity

Cancer is one of the most lethal diseases in human history. Till now, several anticancer medicines have been designed and are in the stage of the clinical trial. However, success limitations, adverse effects, and resistance towards treatments are the major challenges in cancer treatment (Foo and Michor 2014). Therefore, search for novel and effective anticancer agents with least or no side effects is of great interest. Several kinds of research conducted on microbial pigments as anticancer agents exhibited promising results. One such study on novel red pigment extracted from Athrobacter sp. G20 exhibited anticancer potential against the oesophageal cancer cell line (KYSE30) (Afra et al. 2017). A yellow pigment reported from Streptomyces griseoaurantiacus demonstrated strong cytotoxic activity against cervical cancer cells (HeLa) and HepG2 and resulted in a lower number of viable cells (Prashanthi et al. 2015). Carotenoid extracted from Kocuria sp. QWT-12 revealed anticancer potential against breast cancer cell lines and lung cancer cells (Rezaeeyan et al. 2017). Similarly, carotenoids from Haloferax volcanii killed 53.52% of human liver carcinoma cell lines (Sikkandar et al. 2013). Black melanin extracted from S. glaucescens exhibited significant cytotoxic activity against the HFB4 skin cancer cell line (El-Naggar et al. 2017). Dihydroxyphenylalanine melanin obtained from Streptomyces sp. MVCS6 exhibited dose–response anticancer activity against the cervical cancer cell line (Sivaperumal et al. 2015). Wang et al. (2012) studied prodigiosin obtained from Pseudoalteromonas sp. having a cytotoxic effect against U937 leukemia cells. Strong anticancer activity of prodigiosin obtained from S. marcescens is reported against human cervical cancer cells and laryngeal cancer cells (Maheswarappa et al. 2013). Cheng et al. (2017) evaluated the influence of prodigiosin reported from Vibrio sp. against human oral squamous carcinoma cells (OSCC) and found that the prodigiosin arrests their cell cycle. Recently, prodigiosin showed a reduction of the intracellular signaling pathway during the cell cycle and induced apoptosis in lung cancer cells and also showed in vivo tumoricidal activity (Chiu et al. 2018). Interestingly, a combination of natural compounds with chemotherapy drugs enhance its efficacy and reduce their toxicity. Prodigiosin when combined with paclitaxel against MCF7 breast cancer cells (Ho et al. 2009) and doxorubicin (Dox) against OSCC cell lines (Lin and Weng 2018) an efficient synergistic effect was reported. Violacein obtained from C. violaceum induced cytotoxicity and apoptosis activity in Chinese hamster lung fibroblast V79 cells (Melo et al. 2000) and leukemia cell lines (Melo et al. 2003; Ferreira et al. 2004). Additionally, violacein also increased apoptosis in colon cancer cells (Kodach et al. 2017) and human breast cancer cells (Alshatwi et al. 2016). Several anthraquinone derivatives from marine-derived fungi are capable of tumor cell inhibition (Fouillaud et al. 2016). Anthraquinone derivatives from fungus Alternaria sp. have been studied for anticancer activity against human breast cancer cell lines (Huang et al. 2011). In light of the above studies, the microbial pigment could be potential chemotherapeutic agents for cancer treatment.


Bioindicator bacteria are bacteria that can monitor environmental health and reveal the qualitative status of the environment by changing their behavior and specific physiological characteristics (Parmar et al. 2016). One such change is the production and/or alteration of certain pigments in pigmented bacteria that could be effectively used as bioindicators. For instance, violet pigmented bacteria along with Sporocytophaga and Flexibacter species were bioindicators of polluted drinking water (Schindler and Metz 1989). Indigoidine production by Vogesella indigofera is suppressed by Cr6+ in a concentration-dependent manner which is an indication of chromium concentration and toxicity in the environment (Gu and Cheung 2001). The existence of carotenoid in Lecanoraceae lichens has been confirmed to depend upon the pollution level in the atmosphere of the surrounding environment, where they reside by analyzing the carotenoid content of fungi acting as bioindicator (Ibarrondo et al. 2016). Using the gene clusters encoding prodigiosin biosynthesis in S. marcescens and violacein biosynthesis from C. violaceum demonstrate the implementation of reporter systems for the signal of biosynthetic gene expression (Domröse et al. 2017). The participation of melanin pigments in protection from environmental stress like UV radiation and potentially toxic metals is regarded as a bioindicator due to its overproduction in adverse conditions (Egorova et al. 2011). Similarly, cyanobacteria naturally present in water sources could act as excellent bioindicators for heavy metals, since in the presence of heavy metals, the carotenoids content in these cyanobacteria reduces (Wong and Teo 2014).

Miscellaneous applications

Pigment-producing microbes in cryospheric environments could act as a potential source of electrons and may be employed for developing dye-sensitized solar cells (DSSC) that would be a potential alternative of conventional photovoltaic-silicon cells based. Consequently, DSSC might signify a remarkable alternative that could somewhat solve the energy requirement at Antarctic regions. For instance, the orange-xanthophyll pigment extracted from UVC-resistant Hymenobacter sp. (Marizcurrena et al. 2017) was exploited for DSSC development (Órdenes-Aenishanslins et al. 2016; Montagni et al. 2018). Similarly, Pezzella et al. (2019) studied the eumelanin and graphene integration and observed improved electrical conductivity which shows the scope of eumelanin in bioelectronics. Bacterial pigments can be used as biodegradable ink on plastic materials. The hue and chroma values are observed in red and violet color suggestive of prodigiosin and violacein pigments isolated from S. marcescens and C. violaceum, respectively (Venil et al. 2017). Astaxanthin from radioresistant Deinococcus sp. exhibiting radio-protective and antioxidant activities can be incorporated in cosmetics including sunblock and sunscreen (Sajjad et al. 2017). The bacterial pigments prodigiosin and violacein exhibiting antioxidant and antimicrobial activities represent a new paradigm for sunscreens that utilize substances of biological origin (Suryawanshi et al. 2015). These pigments can be a potential ingredient in a range of commercial sunscreen products. Indigoidine can be used as an organic semiconductor with numerous applications in carbon dioxide capture devices, electrochemical cells, super capacitors, batteries, etc. (Yumusak et al. 2019).

Conditions optimization for pigment production

Microbial survival depends upon a wide range of nutrients and physicochemical factors that further control the production of metabolites illustrated in Fig. 3. In general, microbes could adapt three routes for metabolites production. These routes are; natural production of metabolites; metabolites production under strained environmental conditions; and stimulation of metabolites production by supplementing certain nutritional requirements along with physiological parameters (Ramesh et al. 2019). Detail description of pigment extraction has been reviewed from microalgae (Amaro et al. 2016), yeast (Yurkova et al. 2008), fungi (Rai 2009; Ahmad et al. 2015), and bacteria (Stafsnes and Bruheim 2013). Psychrophilic microorganisms produce several pigments to cope with the life-endanger challenges in the cryosphere. Cold-adapted algae are extensively studied for pigment production, however, little is known about the pigment production potential of cold-adapted bacteria and fungi due to several reasons such as their unique physiological and nutritional requirements, slow and unseen growth in the environment, and less exploration in axenic form. To obtain pigments from cold-adapted microorganisms identifying suitable substrates and evaluation of physicochemical factors are essential for better growth and higher pigment yield.

Schematic representation of pigment extraction from cold-adapted microbes at low temperature
Fig. 3
Schematic representation of pigment extraction from cold-adapted microbes at low temperature

Pigment production takes place intracellular or/and extracellular depending on pH, temperature, light, several media constituents (Buck et al. 1974), sampling sites and cultivation conditions (Stafsnes and Bruheim 2013). Under laboratory conditions, the microbial pigment production is ephemeral in nature. However, higher yield could be obtained if several factors such as medium components and environmental parameters are optimized in an understandable way (Ramesh et al. 2019). Such as a study conducted by Pandey et al. (2018) stated that additional supplementation of maltose as a source of carbon in potato dextrose broth boosted the orange pigment production by a psychrophilic strain of Penicillium sp. followed by fructose and glucose, however, supplementation of lactose inhibited this production. Pandey et al. (2018) found that extra nitrogen source could not enhance the pigment production, however, minerals salts like MgSO4 and KH2PO4 can enhance the production of pigment. Maximum pigment production was observed in Monascus sp. supplemented with maltose (Omamor et al. 2008). Pradeep and Pradeep (2013) reported maximum pigment production from Fusarium moniliforme by supplementing glucose in the medium. In the case of filamentous fungi, Carbon/Nitrogen (C/N) ratio is extremely essential in pigment production (Gmoser et al. 2017). A decrease in pigment production was reported by Cho et al. (2002) and Gunasekaran and Poorniammal (2008) during a higher C/N ratio. Similarly, high phosphate concentration and acidity caused the reduction of pigment production and a trace amount of sulfate vitiate pigment production (Reichenbach et al. 1974). Singgih et al. (2015) studied Neurospora intermedia N-1 for carotenoid production in the presence of maltose (2% w/v) as a carbon source. Moreover, minerals salts having different cations were found to improve the production of pigments (Pradeep and Pradeep 2013). Organic acid produced by Monascus ruber inhibits pigment production (Hajjaj et al. 2000). The addition of certain substrates such as wheat, rice meals, and light stimulation enhanced carotenoid production in yeasts and fungi (De Carvalho et al. 2014). ZoBell and Upham (1944) reported an increase in pigment production when bacteria were cultured in sea-water supplemented with neopeptone, bacto-tryptone, and beef extract at 4 °C. Ghosh et al. (2007) reported that methanol acted as a sole carbon source induced the production of pink pigment in Acinetobacter wofii. The yellow–green pigment was produced by Pseudomonas fluorescens supplemented with succinate as a sole carbon source, while this production was stopped with the addition of malic and citric acids as substrate (Margalith 1992). Moreover, the influence of light and dark on pigment production was studied and Myxococcus xanthus, Mycobacterium marinum, Rhodotorula glutinis, and Dacryopinax spathularia produced carotenoids during the presence of light. S. marcescens produced red pigment on solid peptone-glycerol agar plates, and unable to produce pigment in liquid medium except when supplemented with silica gel (Yamashita et al. 2001). Chen et al. (2013) reported that the prodigiosin production was enhanced when peptone and starch were supplemented as a source of carbon and calcium alginate beads were added as a porous carrier. Similarly, Lu et al. (2009) reported sucrose and casein as best carbon and nitrogen sources, respectively, for violacein production by J. lividum XT1 bacteria reported from the glacier in Xinjiang China.

To obtained higher pigment yield, pH and temperature optimization are essential as they affect the metabolic growth and physical activities of microorganisms. Being cold-adapted organisms, the temperature is a vital factor to be optimized for pigment production. Pandey et al. (2018) reported maximum pigment production at low temperature (15 °C) and acidic condition (pH 5) by Penicillium sp. Chattopadhyay et al. (1997) studied Antarctic psychrotrophic bacterium M. roseus that produced higher carotenoid pigment at 5 °C compared to 25 °C. Similarly, Lu et al. (2009) reported an optimal temperature of 15 °C for violacein production by psychrotrophic bacteria. M. tuberculosis produced carotenoid pigments under acidic stress (pH 5–6) (Saviola 2014), and different pigment production was reported in other Mycobacterium species (Robledo et al. 2011). Overproduction of pigment at lower temperatures shows the phenomenon of ecological resilience among cold-adapted microorganisms that combat the deadly environmental conditions. Similarly, enhancement of pigment production below optimum temperature range could be an adapting reaction that compensates the downregulation of metabolic processes at low temperatures (Gmoser et al. 2017). Several genes are responsible for the biosynthesis of pigments in microorganisms. To enhance the pigment production mutagens such as UV light, 1-methyl-3-nitro-1-nitrosoguanidine, and ethyl methanesulfonate were used in Haematococcus pluvialis and microwave in S. marcescens (Nigam and Luke 2016), which shows these mutagens could enhance the pigment yield when used precisely. In addition, alteration in genes (knockout or promotion of genes) and mutagenesis practices can increase the pigment production (Venil et al. 2014). Different substrates such as tryptophan, phenylalanine, tyrosine, and several physicochemical parameters can efficiently stimulate pigment production by microorganisms.

Essential pigments from fungal strains can be obtained under optimum growth conditions by solid, semi-solid, and submerged fermentation (Vendruscolo et al. 2010; Akilandeswari and Pradeep 2016) using a diverse range of sustainable substrates (Gupta and Aggarwal. 2014). However, parameters such as pH, temperature, agitation, aeration, and culture medium directly influence the production of pigment and should be kept at an optimized level (Medentsev et al. 2005). Moreover, several solvents such as acetone, chloroform, cyclohexane, acetonitrile, ethanol, dichloromethane, pyridine, hexane, and water are used to extract fungal pigments. The solvent choice is crucial for pigment extraction and depends upon the polarity of the studied molecules. After extraction and purification of pigments, it should be handled carefully and must be kept under optimum temperature, and pH to preserve its original colorimetric characteristics. Pigment storage in dry powder form is more preferable because of its higher stability and low water activity (De Carvalho et al. 2014).

Challenges in pigments obtained from psychrophiles

Antarctic region and the adjacent Southern Ocean is the rich source of psychrophilic microorganisms capable of pigment production. However, in 2014 the bioprospection is defined during XXXVII Antarctic Treaty Consultative Meeting held in Brasilia. According to this definition, bioprospection is “any activity of search, identification, description, collection, survey, monitoring, cultivation, replication, or any other scientific investigation processes, performed on indigenous biological species with the initial intention to consider potential industrial or commercial derived products or applications, notably through the development of patentable material or process.” According to this definition, research curiosity on the biological resources of Antarctica is growing for commercial use. However, several reports highlighted that due to the lack of clear governing and ownership rules for these resources causes appropriation by those who discover it and subsequently develop patent (UNU-UNEP 2009; Jabour 2010; UNEP 2012; Puig-Marcó 2014). In 2008, Antarctic Treaty Committee launched an online database for bioprospecting in Antarctica that provides up-to-date information about bioprospecting in Antarctica. This database also provides information about commercialized products obtained from biological samples indigenous to Antarctica. The records in the database contained many patents related to microbial metabolites having potential biotechnological applications (UNEP Report 2012). The increase in patents shows that Antarctic microorganisms are a precious source of marvelous potential profits. However, one of the leading challenges is the assurance of sustainable exploitation of indigenous biological resources of Antarctica which is yet to decide that how positive benefits of bioprospecting could achieve without inducing major damage to the indigenous environment. Since the uncontrolled prospecting actions that would include logistical labors to reach the biological resources and their definite outcomes could make the process unsustainable and compromise the indigenous microbial diversity (Hughes et al. 2015). In addition, the commercial based research activities and their secrecy could compromise the main pillar of the Antarctic Treaty System (Article III), viz., the unrestricted cooperation and free exchange of information among all parties (Yarzábal et al. 2016). Furthermore, the emerging profitable interest in biological resources raised several key policies, moral and ethical queries: Who would be the owner of these biological resources? How should they be exploited and used? And whom should be benefited and to what extent? Although, few of these questions are included in the 2014 SCAR1-sponsored Antarctic and Southern Ocean Horizon Scan (Kennicutt et al. 2014, 2015). However, several current uncertainty and absence of policies for these biological resources should be addressed to the territorial and commercial sensitivities to agree upon common agreement to protect the local environment. For instance, the Nagoya protocol that entered into force in October 2014 by over 70 countries under international law for the fair and equitable sharing of benefits derived from the utilization of genetic resources (Smith et al. 2017). Like Nagoya protocol, countries should be placed under an obligation to establish legislative, administrative and/or political measures regarding the access to common cold reservoirs such as Arctic and Antarctic. This will aim to provide equal chances of obtaining common cold resources for commercial or academic development by a country or organization. Moreover, to commercialize any kind of microbial pigments, huge investment along with research work for characterization, optimization processes, possible toxicological testing, regulatory approval, and penchant by the consumers are greatly essential (Ahmad et al. 2015; Dufossé et al. 2016; Carvalho et al. 2016). Similarly, optimized fermentation conditions, design and type of bioreactor to achieve desired productivity of microbial pigments (Venil et al. 2014) along with the downstream processing of product purification are quite essential to be considered especially for microorganisms from unusual environments such as cryosphere. Therefore, widespread research is essential to convey the microbial pigments from laboratories to markets, since its demand is higher than supply.


The demand for natural pigments obtained from microorganisms is now increasing due to the adverse effects causing by synthetic dyes. Therefore, new potential sources are crucial to explore for pigment-producing microorganisms. One such environment is the cryosphere which inhabits microorganisms that produce natural pigments as a protective shield against life-endangered ecological stresses. Exploration of these microorganisms would certainly provide promising sources of novel pigmented molecules having wider biotechnological applications. These unexplored microbes especially novel species could be the potential source for pigments that could be used to develop novel drug compounds as well as in the textile and food industries. Efforts should be done to produce cost-effective production of pigments from these microbes by optimization, and strain improvement through genetic engineering to get rid of toxic synthetic dyes. Sustainable exploitation of these biological resources from the cryosphere should be focused to prevent environmental disruption. Several cryospheric environments such as Antarctica should be considered a common legacy for mankind and the international community should declare a clear ethical and moral policy for certain questions including possession of these resources. It is obvious that due to unique qualities, the applications of these pigments obtained from cold-adapted microorganisms would be broader and will start many newer domains in biotechnology.

Publisher's Note:

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.


This study was supported by the second Tibetan Plateau Scientific Expedition and Research Program (STEP) (2019QZKK0605), the Strategic Priority Research Programme of the Chinese Academy of Sciences, Pan-Third Pole Environment Study for a Green Silk Road (Pan-TPE) (XDA20040501), the National Natural Science Foundation of China (Grant 41630754). Wasim Sajjad is supported by a PIFI Fellowship from the Chinese Academy of Sciences (2020PC0052).

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.


    Aberoumand   A . A review article on edible pigments properties and sources as natural biocolorants in foodstuff and food industry. WJDFS 2011. 6: 1 71-78
    Afra   S , Makhdoumi   A , Matin   MM , Feizy   J . A novel red pigment from marine Arthrobacter sp. G20 with specific anticancer activity. J Appl Microbiol 2017. 123: 5 1228-1236 doi: 10.1111/jam.13576
    Agogué   H , Joux   F , Obernosterer   I , Lebaron   P . Resistance of marine bacterioneuston to solar radiation. Appl Environ Microbiol 2005. 71: 5282-5289 doi: 10.1128/AEM.71.9.5282-5289.2005
    Ahmad   WA , Venil   CK , Aruldass   CA Liong   MT . Production of violacein by Chromobacterium violaceum grown in liquid pineapple waste: current scenario. Beneficial microorganisms in agriculture, aquaculture and other areas, microbiology monographs, 29 2015. Berlin Springer 45-58
    Ahmed   N , Singh   J , Mudasir   MA , Kour   H , Gupta   P , Chauhan   H . Naturally occurring preservatives in food and their role in food preservation. Int J Pharm Bio Sci 2013. 4: 22-30
    Akilandeswari   P , Pradeep   BV . Exploration of industrially important pigments from soil fungi. Appl Microbiol Biotechnol 2016. 100: 1631-1643 doi: 10.1007/s00253-015-7231-8
    Alihosseini   F , Ju   KS , Lango   J , Hammock   BD , Sun   G . Antibacterial colorants: characterization of prodiginines and their applications on textile materials. Biotechnol Prog 2008. 24: 3 742-747 doi: 10.1021/bp070481r
    Alija   AJ , Sommerburg   O , Langhans   CD , Siems   W , Eckl   PM . Cyto- and genotoxic potential of beta-carotene and cleavage products under oxidative stress. BioFactors 2005. 24: 159-163 doi: 10.1002/biof.5520240119
    Alshatwi   AA , Subash-Babu   P , Antonisamy   P . Violacein induces apoptosis in human breast cancer cells through up regulation of BAX, p53 and down regulation of MDM2. Exp Toxicol Pathol 2016. 68: 89-97 doi: 10.1016/j.etp.2015.10.002
    Amaretti   A , Simone   M , Quartieri   A , . Isolation of Carotenoid-producing Yeasts from an Alpine Glacier. Chem Eng Trans 2014. 38: 217-222
    Amaro   HM , Sousa-Pinto   I , Malcata   FX , Guedes   AC Leo   MLN . Microalgae as a source of pigments extraction and purification methods. Marine microorganisms extraction and analysis of bioactive compounds 2016. Boca Raton CRC Press 99-128
    Ambrožič Avguštin   J , Žgur Bertok   D , Kostanjšek   R , Avguštin   G . Isolation and characterization of a novel violacein-like pigment producing psychrotrophic bacterial species Janthinobacterium svalbardensis sp. nov. Anton Van Leeuwen 2013. 103: 763-769 doi: 10.1007/s10482-012-9858-0
    Andrighetti-Fröhner   CR , Antonio   RV , Creczynski-Pasa   TB , . Cytotoxicity and potential antiviral evaluation of violacein produced by Chromobacterium violaceum. The Memórias do Instituto Oswaldo Cruz 2003. 98: 843-848 doi: 10.1590/S0074-02762003000600023
    Anesio   AM , Laybourn-Parry   J . Glaciers and ice sheets as a biome. Trends Ecol Evol 2012. 27: 219-225 doi: 10.1016/j.tree.2011.09.012
    Anesio   AM , Lutz   S , Chrismas   NA , Benning   LG . The microbiome of glaciers and ice sheets. NPJ Biofilms Microb 2017. 3: 1 1-11 doi: 10.1038/s41522-017-0019-0
    Antonisamy   P , Ignacimuthu   S . Immunomodulatory, analgesic and antipyretic effects of violacein isolated from Chromobacterium violaceum. Phytomedicine 2010. 17: 300-304 doi: 10.1016/j.phymed.2009.05.018
    Arcangeli   C , Zucconi   L , Onofri   S , Cannistraro   S . Fluorescence study on whole Antarctic fungal spores under enhanced UV irradiation. J Photochem Photobiol B 1997. 39: 3 258-264 doi: 10.1016/S1011-1344(97)00019-5
    Arcangeli   C , Yu   W , Cannistraro   S , Gratton   E . Two-photon autofluorescence microscopy and spectroscopy of an Antarctic fungus: a new approach for studying the effects of UV-B irradiation. Biospectroscopy 2000. 57: 218-225
    Bakermans   C , Tollaksen   SL , Giometti   CS , . Proteomic analysis of Psychrobacter cryohalolentis K5 during growth at subzero temperatures. Extremophiles 2007. 11: 343-354 doi: 10.1007/s00792-006-0042-1
    Bar Dolev   M , Bernheim   R , Guo   S , . Putting life on ice: bacteria that bind to frozen water. J R Soc Interface 2016. doi: 10.1098/rsif.2016.0210
    Barahona   S , Yuivar   Y , Socias   G , . Identification and characterization of yeasts isolated from sedimentary rocks of Union Glacier at the Antarctica. Extremophiles 2016. 20: 479-491 doi: 10.1007/s00792-016-0838-6
    Barauna   RA , Freitas   DY , Pinheiro   JC , . A proteomic perspective on the bacterial adaptation to cold: integrating OMICs data of the psychrotrophic bacterium Exiguobacterium antarcticum B7. Proteomes 2017. doi: 10.3390/proteomes5010009
    Batista   JH , da Silva Neto   JF . Chromobacterium violaceum pathogenicity: updates and insights from genome sequencing of novel Chromobacterium species. Front Microbiol 2017. 8: 2213 doi: 10.3389/fmicb.2017.02213
    Becker-Hapak   M , Troxtel   E , Hoerter   J , Eisenstark   A . RpoS dependent overexpression of carotenoids from Erwinia herbicola in OXYR-deficient Escherichia coli. Biochem Bioph Res Co 1997. 239: 305-309 doi: 10.1006/bbrc.1997.7469
    Benforte   FC , Colonnella   MA , Ricardi   MM , . Novel role of the LPS core glycosyltransferase WapH for cold adaptation in the Antarctic bacterium Pseudomonas extremaustralis. PLoS ONE 2018. 13: 2 e0192559 doi: 10.1371/journal.pone.0192559
    Berlanga   M , Ruiz   N , Hernandez-Borrell   J , . Role of the outer membrane in the accumulation of quinolones by Serratia marcescens. Can J Microbiol 2000. 46: 716-722 doi: 10.1139/w00-052
    Borić   M , Danevčič   T , Stopar   D . Prodigiosin from Vibrio sp. DSM 14379; a new UV-protective pigment. Microb Ecol 2011. 62: 3 528 doi: 10.1007/s00248-011-9857-0
    Bowman   JP , Brown   MV , Nichols   DS . Biodiversity and ecophysiology of bacteria associated with Antarctic sea ice. Antarct Sci 1997. 9: 2 134-142 doi: 10.1017/S0954102097000175
    Buck   JD . Effects of medium composition on the recovery of bacteria from sea water. J Exp Mar Biol Ecol 1974. 15: 25-34 doi: 10.1016/0022-0981(74)90060-4
    Burger   SR , Bennett   JW . Droplet enrichment factors of pigmented and nonpigmented Serratia marcescens: possible selective function for prodigiosin. Appl Environ Microbiol 1985. 50: 487-490 doi: 10.1128/AEM.50.2.487-490.1985
    Burt   SA , Ojo-Fakunle   VT , Woertman   J , Veldhuizen   EJ . The natural antimicrobial carvacrol inhibits quorum sensing in Chromobacterium violaceum and reduces bacterial biofilm formation at sub-lethal concentrations. PLoSOne 2014. 9: 4 e93414 doi: 10.1371/journal.pone.0093414
    Butler   MJ , Day   AW . Fungal melanins. Can J Microbiol 1998. 44: 1115-1136 doi: 10.1139/w98-119
    Caruso   C , Rizzo   C , Mangano   S , . Production and biotechnological potential of extracellular polymeric substances from sponge-associated Antarctic bacteria. Appl Environ Microbiol 2018. doi: 10.1128/AEM.01624-17
    Carvalho   JC , Bicas   JL , Fernández   DER , Bicas   JL , Maróstica   MR , Pastore   GM , . Natural colorants from microorganisms. Biotechnological production of natural ingredients for food industry 2016. 1Sharjah Bentham Science Publishers 288-321
    Castro-Sowinski   S , Matan   O , Bonafede   P , Okon   Y . A thioredoxin of Sinorhizobium meliloti CE52G is required for melanin production and symbiotic nitrogen fixation. Mol Plant Microbe In 2007. 20: 986-993 doi: 10.1094/MPMI-20-8-0986
    Cavicchioli   R , Charlton   T , Ertan   H , . Biotechnological uses of enzymes from psychrophiles. Microb Biotechnol 2011. 4: 4 449-460 doi: 10.1111/j.1751-7915.2011.00258.x
    Centurion   VB , Delforno   TP , Lacerda-Júnior   GV , . Unveiling resistome profiles in the sediments of an Antarctic volcanic island. Environ Pollut 2019. 255: 113240 doi: 10.1016/j.envpol.2019.113240
    Chadni   Z , Rahaman   MH , Jerin   I , . Extraction and optimisation of red pigment production as secondary metabolites from Talaromyces verruculosus and its potential use in textile industries. Mycology 2017. 8: 48-57 doi: 10.1080/21501203.2017.1302013
    Chattopadhyay   M , Jagannadham   M . Maintenance of membrane fluidity in Antarctic bacteria. Polar Biol 2001. 24: 5 386-388 doi: 10.1007/s003000100232
    Chattopadhyay   MK , Jagannadham   MV , Vairamani   M , Shivaji   S . Carotenoid pigments of an antarctic psychrotrophic bacteriumMicrococcus roseus: temperature dependent biosynthesis, structure, and interaction with synthetic membranes. Biochem Biophys Res Commun 1997. 239: 1 85-90 doi: 10.1006/bbrc.1997.7433
    Chattopadhyay   P , Chatterjee   S , Sen   SK . Biotechnological potential of natural food grade biocolorants. Afr J Biotechnol 2008. 7: 17 2972-2985
    Chen   WC , Yu   WJ , Chang   CC , . Enhancing production of prodigiosin from Serratia marcescens C3 by statistical experimental design and porous carrier addition strategy. Biochem Eng J 2013. 78: 93-100 doi: 10.1016/j.bej.2013.02.001
    Cheng   MF , Lin   CS , Chen   YH , . Inhibitory growth of oral squamous cell carcinoma cancer via bacterial prodigiosin. Mar Drugs 2017. 15: 224 doi: 10.3390/md15070224
    Chequer   FD , de Oliveira   GAR , Ferraz   ERA , . Textile dyes: dyeing process and environmental impact. Eco-friendly Textile Dye Finish 2013. 6: 151-176
    Chiu   WJ , Lin   SR , Chen   YH , . Prodigiosin-emerged PI3K/Beclin-1-independent pathway elicits autophagic cell death in Doxorubin-sensitive and resistant lung cancer. J Clin Med 2018. 7: 321 doi: 10.3390/jcm7100321
    Cho   YJ , Park   JP , Hwang   HJ , . Production of red pigment by submerged culture of Paecilomyces sinclairii. Lett Appl Microbiol 2002. 35: 195-202 doi: 10.1046/j.1472-765X.2002.01168.x
    Choi   SY , Yoon   KH , Lee   JI , Mitchell   RJ . Violacein: properties and production of a versatile bacterial pigment. Biomed Res Int 2015. doi: 10.1155/2015/465056
Chyizhanska N, Beregova T (2009) Effect of melanin isolated from Antarctic yeasts on preservation of pig livestock after ablactation. Укpaїнcький aнтapктичний жypнaл.
    Collins   T , Gerday   C Margesin   R . Enzyme catalysis in psychrophiles. Psychrophiles: from biodiversity to biotechnology 2017. 2Cham Springer 209-235
    Collins   T , Margesin   R . Psychrophilic lifestyles: mechanisms of adaptation and biotechnological tools. Appl Microbiol Biotechnol 2019. 103: 7 2857-2871 doi: 10.1007/s00253-019-09659-5
    Contreras   G , Barahona   S , Sepúlveda   D , . Identification and analysis of metabolite production with biotechnological potential in Xanthophyllomyces dendrorhous isolates. World J Microbiol Biotechnol 2015. 31: 3 517-526 doi: 10.1007/s11274-015-1808-3
    Correa-Llantén   DN , Amenabar   MJ , Blamey   JM . Antioxidant capacity of novel pigments from an Antarctic bacterium. J Microbiol 2012. 50: 374-379 doi: 10.1007/s12275-012-2029-1
    Corsaro   MM , Casillo   A , Parrilli   E , Tutino   ML Margesin   R . Molecular structure of lipopolysaccharides of cold-adapted bacteria. Psychrophiles: from biodiversity to biotechnology 2017. 2Cham Springer 285-303
    Cristea   D , Vilarem   G . Improving light fastness of natural dyes on cotton yarn. Dyes Pigm 2006. 70: 3 238-245 doi: 10.1016/j.dyepig.2005.03.006
    Cude   WN , Mooney   J , Tavanaei   AA , . Production of the antimicrobial secondary metabolite indigoidine contributes to competitive surface colonization by the marine roseobacter Phaeobacter sp. strain Y4I. Appl Environ Microbiol 2012. 78: 14 4771-4780 doi: 10.1128/AEM.00297-12
    D'Amico   S , Gerday   C , Feller   G . Structural determinants of cold adaptation and stability in a large protein. J Biol Chem 2001. 276: 28 25791-25796 doi: 10.1074/jbc.M102741200
    D’Amico   S , Collins   T , Marx   JC , . Psychrophilic microorganisms: challenges for life. EMBO Rep 2006. 7: 4 385-389 doi: 10.1038/sj.embor.7400662
    Danevčič   T , Borić Vezjak   M , Tabor   M , . Prodigiosin induces autolysis in actively grown Bacillus subtilis cells. Front Microbiol 2016. 7: 27 doi: 10.3389/fmicb.2016.00027
    Day   PA , Villalba   MS , Herrero   OM , . Formation of indigoidine derived-pigments contributes to the adaptation of Vogesella sp. strain EB to cold aquatic iron-oxidizing environments. Antonie Van Leeuwenhoek 2017. 110: 3 415-428 doi: 10.1007/s10482-016-0814-2
    De Carvalho   JC , Cardoso   LC , Ghiggi   V , Woiciechowski   AL , de Souza Vandenberghe   LP , Soccol   CR Brar   S , Dhillon   G , Soccol   C . Microbial Pigments. Biotransformation of Waste Biomass into High Value Biochemicals 2014. New York, NY, USA Springer 73-98
    De Maayer   P , Anderson   D , Cary   C , Cowan   DA . Some like it cold: understanding the survival strategies of psychrophiles. EMBO Rep 2014. 15: 5 508-517 doi: 10.1002/embr.201338170
    de Menezes   GC , Godinho   VM , Porto   BA , . Antarctomyces pellizariae sp. nov., a new, endemic, blue, snow resident psychrophilic ascomycete fungus from Antarctica. Extremophiles 2017. 21: 259-269 doi: 10.1007/s00792-016-0895-x
    Deming   JW , Young   JN Margesin   R . The role of exopolysaccharides in microbial adaptation to cold habitats. Psychrophiles: from biodiversity to biotechnology 2017. 2Cham Springer 259-284
    Dharmaraj   S , Ashokkumar   B , Dhevendaran   K . Food-gradepigments from Streptomyces sp. isolated from the marine sponge Callyspongia diffusa. Food Res Int 2009. 42: 487-492 doi: 10.1016/j.foodres.2009.02.006
    Dieser   M , Greenwood   M , Foreman   CM . Carotenoid pigmentation in Antarctic heterotrophic bacteria as a strategy to withstand environmental stresses. Arct Antarct Alp Res 2010. 42: 4 396-405 doi: 10.1657/1938-4246-42.4.396
    Dillon   JG , Castenholz   RW . Scytonemin, a cyanobacterial sheath pigment, protects against UVC radiation: implications for early photosynthetic life. J Phycol 1999. 35: 4 673-681 doi: 10.1046/j.1529-8817.1999.3540673.x
    Dimitrova   S , Pavlova   K , Lukanov   L , Zagorchev   P . Synthesis of coenzyme q10 and β-carotene by yeasts isolated from Antarctic soil and lichen in response to ultraviolet and visible radiations. Appl Biochem Biotechnol 2010. 162: 795-804 doi: 10.1007/s12010-009-8845-z
    Dimitrova   S , Pavlova   K , Lukanov   L , . Production of metabolites with antioxidant and emulsifying properties by Antarctic strain Sporobolomyces salmonicolor AL1. Appl Biochem Biotechnol 2013. 169: 301-311 doi: 10.1007/s12010-012-9983-2
    Domröse   A , Weihmann   R , Thies   S , . Rapid generation of recombinant Pseudomonas putida secondary metabolite producers using yTREX. Synth Syst Biotechnol 2017. 2: 4 310-319 doi: 10.1016/j.synbio.2017.11.001
    Dsouza   M , Taylor   MW , Turner   SJ , Aislabie   J . Genomic and phenotypic insights into the ecology of Arthrobacter from Antarctic soils. BMC Genomics 2015. 16: 1 36 doi: 10.1186/s12864-015-1220-2
    Dufossé   L Carle   R , Schweiggert   RM . Current and potential natural pigments from microorganisms (bacteria, yeasts, fungi, microalgae). Handbook on natural pigments in food and beverages 2016. Cambridge Elsevier Ltd 337-354
    Dufosse   L . Microbial pigments from bacteria, yeasts, fungi, and microalgae for the food and feed industries. Natural and artificial flavoring agents and food dyes 2018. USA Academic Press 113-132
    Durán   N , Erazo   S , Campos   V . Bacterial chemistry-II: antimicrobial photoproduct from pigment of Chromobacterium violaceum. An Acad Bras Cienc 1953. 55: 231-234
Duran N, Melo PS, Haun M (1996) In Vitro evaluation of violacein on AIDS-related lumphoma and human tumor cell lines. In: Proceedings of the 25th Annual Meetings of the Brazilian Society of Biochemistry and Molecular Biology, Sociedade Brasi-leira de Bioquimica e Biologia Molecular (SBBq), Caxambu, Brazil
    Duran   N , Justo   GZ , Ferreira   CV , . Violacein: properties and biological activities. Biotechnol Appl Biochem 2007. 48: 127-133 doi: 10.1042/BA20070115
    Egorova   AS , Gessler   NN , Belozerskaya   TA . Melanin pigments in the fungus Paecilomyces lilacinus (thom) samson. Dokl Biochem Biophys 2011. 437: 1 84 doi: 10.1134/S1607672911020086
    El-Naggar   NE , El-Ewasy   SM . Bioproduction, characterization, anticancer and antioxidant activities of extracellular melanin pigment produced by newly isolated microbial cell factories Streptomyces glaucescens NEAE-H. Sci Rep 2017. 14: 42129 doi: 10.1038/srep42129
    Ewert   M , Deming   JW . Sea ice microorganisms: environmental constraints and extracellular responses. Biology 2013. 2: 2 603-628 doi: 10.3390/biology2020603
    Feofilova   EP , Tereshina   VM , Gorova   IB . Changes in carbohydrate composition of fungi during adaptation to thermostress. Microbiology 1994. 63: 442-445
    Ferreira   CV , Bos   CL , Versteeg   HH , . Molecular mechanism of violacein-mediated human leukemia cell death. Blood 2004. 104: 14464-14591 doi: 10.1182/blood-2004-02-0594
    Fleming   ED , Castenholz   RW . Effects of periodic desiccation on the synthesis of the UV-screening compound, scytonemin, in cyanobacteria. Environ Microbiol 2007. 9: 6 1448-1455 doi: 10.1111/j.1462-2920.2007.01261.x
    Foght   J , Aislabie   J , Turner   S , . Culturable bacteria in subglacial sediments and ice from two southern hemisphere glaciers. Microb Ecol 2004. 47: 4 329-340 doi: 10.1007/s00248-003-1036-5
    Fong   NJC , Burgess   ML , Barrow   KD , Glenn   DR . Carotenoid accumulation in the psychrotrophic bacterium Arthrobacter agilis in response to thermal and salt stress. Appl Microbiol Biotechnol 2001. 56: 750-756 doi: 10.1007/s002530100739
    Foo   J , Michor   F . Evolution of acquired resistance to anti-cancer therapy. J Theor Biol 2014. 355: 10-20 doi: 10.1016/j.jtbi.2014.02.025
    Fouillaud   M , Venkatachalam   M , Girard-Valenciennes   E , . Anthraquinones and derivatives from marine-derived fungi: structural diversity and selected biological activities. Mar Drugs 2016. 14: 64 doi: 10.3390/md14040064
    Frank   S , Schmidt   F , Klockgether   J , . Functional genomics of the initial phase of cold adaptation of Pseudomonas putida KT2440. FEMS Microbiol Lett 2011. 318: 47-54 doi: 10.1111/j.1574-6968.2011.02237.x
    Fraser   PD , Bramley   PM . The biosynthesis and nutritional uses of carotenoids. Prog Lipid Res 2004. 43: 3 228-265 doi: 10.1016/j.plipres.2003.10.002
    Fukuda   W , Kimura   T , Araki   S , . Lysobacteroligotrophicus sp. Nov., isolated from an Antarctic freshwater lake in Antarctica. Int J Syst Evol Microbiol 2013. 63: 9 3313-3318 doi: 10.1099/ijs.0.051805-0
    Füller   JJ , Röpke   R , Krausze   J , . Biosynthesis of violacein, structure and function of l-Tryptophan oxidase VioA from Chromobacterium violaceum. J Biol Chem 2016. 291: 38 20068-20084 doi: 10.1074/jbc.M116.741561
    Gessler   NN , Egorova   AS , Belozerskaya   TA . Fungal anthraquinones. Appl Biochem Microbiol 2013. 49: 85-99 doi: 10.1134/S000368381302004X
    Gessler   NN , Egorova   AS , Belozerskaya   TA . Melanin pigments of fungi under extreme environmental conditions. Appl Biochem Micro 2014. 50: 2 105-113 doi: 10.1134/S0003683814020094
    Ghobakhlou   AF , Johnston   A , Harris   L , . Microarray transcriptional profiling of Arctic Mesorhizobium strain N33 at low temperature provides insights into cold adaption strategies. BMC Genomics 2015. 16: 1 383 doi: 10.1186/s12864015-1611-4
    Ghosh   A , Goyal   A , Jain   RK . Study of methanol-induced phenotypic changes in a novel strain of Acinetobacter lwoffii. Arch Microbiol 2007. 188: 533-539 doi: 10.1007/s00203-007-0268-z
    Gmoser   R , Ferreira   JA , Lennartsson   PR , Taherzadeh   MJ . Filamentous ascomycetes fungi as a source of natural pigments. Fungal Biol Biotechnol 2017. 4: 1 4 doi: 10.1186/s40694-017-0033-2
    González-Toril   E , Amils   R , Delmas   RJ , . Diversity of bacteria producing pigmented colonies in aerosol, snow and soil samples from remote glacial areas (Antarctica, Alps and Andes). Biogeosci Discuss 2008. 5: 1607-1630 doi: 10.5194/bgd-5-1607-2008
    Goordial   J , Raymond-Bouchard   I , Zolotarov   Y , . Cold adaptive traits revealed by comparative genomic analysis of the eurypsychrophile Rhodococcus sp. JG3 isolated from high elevation McMurdo Dry Valley permafrost, Antartica. FEMS Microbiol Ecol 2016. doi: 10.1093/femsec/fiv154
    Gorbushina   A . Methodologies and techniques for detecting extraterrestrial (microbial) life microcolonial fungi: survival potential of terrestrial vegetative structures. Astrobiology 2003. 3: 543-554 doi: 10.1089/153110703322610636
    Gounot   AM . Bacterial life at low temperature: physiological aspects and biotechnological implications. J Appl Bacteriol 1991. 71: 386-397 doi: 10.1111/j.1365-2672.1991.tb03806.x
    Gromek   SM , Suria   AM , Fullmer   MS , . Leisingera sp JC1, a bacterial isolate from Hawaiian bobtail squid eggs, produces indigoidine and differentially inhibits vibrios. Front Microbiol 2016. 7: 1342 doi: 10.3389/fmicb.2016.01342
    Gu   JD , Cheung   KH . Phenotypic expression of Vogesella indigofera upon exposure to hexavalent chromium, Cr6+. World J Microb Biot 2001. 17: 475-480 doi: 10.1023/A:1011966725978
    Gunasekaran   S , Poorniammal   R . Optimization of fermentation conditions for red pigment production from Penicillium sp. under submerged cultivation. Afr J Biotechnol 2008. 7: 1894-1898 doi: 10.5897/AJB2008.000-5037
    Gupta   S , Aggarwal   S . Novel bio-colorants for textile application from fungi. J Textile Assoc 2014. 74: 282-287
    Haddix   PL , Jones   S , Patel   P , . Kinetic analysis of growth rate, ATP, and pigmentation suggests an energy-spilling function for the pigment prodigiosin of Serratia marcescens. J Bacteriol 2008. 190: 22 7453-7463 doi: 10.1128/JB.00909-08
    Hajjaj   H , Blanc   P , Groussac   E , . Kinetic analysis of red pigment and citrinin by Monascus rubber as a function of organic acid accumulation. Enzym Microb Technol 2000. 27: 619-625 doi: 10.1016/S0141-0229(00)00260-X
    Hakvag   S , Fjaervik   E , Klinkenberg   G , . Violacein-producing Collimonas sp. from the sea surface microlayer of costal waters in Trondelag, Norway. Mar Drugs 2009. 7: 576-588 doi: 10.3390/md7040576
    Harris   AK , Williamson   NR , Slater   H , . The Serratia gene cluster encoding biosynthesis of the red antibiotic, prodigiosin, shows species and strain-dependent genome context variation. Microbiology 2004. 150: 3547-3560 doi: 10.1099/mic.0.27222-0
    Hasan   N , Hasan   F , Nadeem   S , . Community analysis and characterization of fungi from Batura glacier, Karakoram mountain range, Pakistan. Appl Ecol Environ Res 2018. 16: 5 5323-5341 doi: 10.15666/aeer/1605_53235341
    Hassan   N , Anesio   AM , Rafiq   M , . Cell membrane fatty acid and pigment composition of the psychrotolerant cyanobacterium Nodularia spumigena CHS1 isolated from Hopar glacier, Pakistan. Extremophiles 2020. 24: 1 135-145 doi: 10.1007/s00792-019-01141-4
    He   J , Yang   Z , Hu   B , . Correlation of polyunsaturated fatty acids with the cold adaptation of Rhodotorula glutinis. Yeast 2015. 32: 11 683-690 doi: 10.1002/yea.3095
    Heider   SA , Peters-Wendisch   P , Wendisch   VF . Carotenoid biosynthesis and overproduction in Corynebacterium glutamicum. BMC Microbiol 2012. 12: 1 198 doi: 10.1186/1471-2180-12-198
    Ho   TF , Peng   YT , Chuang   SM , . Prodigiosin down-regulates survivin to facilitate paclitaxel sensitization in human breast carcinoma cell lines. Toxicol Appl Pharm 2009. 235: 253-260 doi: 10.1016/j.taap.2008.12.009
    Hood   DW , Heidstra   R , Swoboda   UK , Hodgson   DA . Molecular genetic analysis of proline and tryptophan biosynthesis in Streptomyces coelicolor A3(2): interaction between primary and secondary metabolism—a review. Gene 1992. 115: 5-12 doi: 10.1016/0378-1119(92)90533-U
    Huang   HC , Chiu   SH , Ke   HJ , . Antimelanogenic and antioxidant activities of Bifidobacterium infantis. Afr J Microbiol Res 2011. 5: 3150-3156 doi: 10.5897/AJMR11.021
    Huang   JP , Mojib   N , Goli   RR , . Antimicrobial activity of PVP from an Antarctic bacterium, Janthinobacterium sp. Ant5–2, on multi-drug and methicillin resistant Staphylococcus aureus. Nat Products Bioprospect 2012. 2: 3 104-110 doi: 10.1007/s13659-012-0021-4
    Hughes   KA , Lawley   B , Newsham   KK . Solar UV-B radiation inhibits the growth of Antarctic terrestrial fungi. Appl Environ Microbiol 2003. 69: 1488-1491 doi: 10.1128/AEM.69.3.1488-1491.2003
    Hughes   KA , Cowan   DA , Wilmotte   A . Protection of Antarctic microbial communities—‘out of sight, out of mind’. Front Microbiol 2015. 6: 151 doi: 10.3389/fmicb.2015.00151
    Ibarrondo   I , Prieto-Taboada   N , Martínez-Arkarazo   I , Madariaga   JM . Resonance Raman imaging as a tool to assess the atmospheric pollution level: carotenoids in Lecanoraceae lichens as bioindicators. Environ Sci Pollut 2016. 23: 7 6390-6399 doi: 10.1007/s11356-015-5849-9
    Ibrahim   A . Production of carotenoids by a newly isolated marine Micrococcus sp.. Biotechnology 2008. 7: 469-474 doi: 10.3923/biotech.2008.469.474
    Jabour   J . Biological prospecting: the ethics of exclusive reward from Antarctic activity. Ethics Sci Environ Polit 2010. 10: 19-29 doi: 10.3354/esep00105
    Joshi   VK , Attri   D , Bala   A , Bhushan   S . Microbial pigments. Indian J Biotechnol 2003. 2: 362-369
    Kamla   M , Jayanti   T , Sneh   G . A review on microbial pigment. Int J Microbial Res Technol 2012. 1: 4 361-365
    Kennicutt   MC , Chown   SL , Cassano   JJ , . Polar research: six priorities for Antarctic science. Nature 2014. 512: 7512 23-25 doi: 10.1038/512023a
    Kim   SK Marine biomaterials: characterization, isolation and applications 2013. Boca Raton CRC Press/Taylor & Francis
    Kim   SJ , Shin   SC , Hong   SG , Lee   YM , Lee   H , Lee   J , Choi   IG , Park   H . Genome sequence of Janthinobacterium sp. strain PAMC 25724, isolated from alpine glacier cryoconite. J Bacteriol 2012. 194: 8 2096-2096 doi: 10.1128/JB.00096-12
    Kim   JH , Kang   HJ , Yu   BJ , . Planococcus faecalis sp. Nov., a carotenoid-producing species isolated from stools of Antarctic penguins. Int J Syst Evol Microbiol 2015. 65: 10 3373-3378 doi: 10.1099/ijsem.0.000423
    Kimura   T , Fukuda   W , Sanada   T , Imanaka   T . Characterization of water-soluble dark-brown pigment from Antarctic bacterium, Lysobacter oligotrophicus. J Biosci Bioeng 2015. 120: 1 58-61 doi: 10.1016/j.jbiosc.2014.11.020
    Kitamoto   D , Yanagishita   H , Endo   A , . Remarkable antiagglomeration effect of a yeast biosurfactant, diacylmannosylerythritol, on ice-water slurry forcold thermal storage. Biotechnol Prog 2001. 17: 2 362-365 doi: 10.1021/bp000159f
    Kobayashi   H , Nogi   Y , Horikoshi   K . New violet 3, 3′-bipyridyl pigment purified from deep-sea microorganism Shewanella violacea DSS12. Extremophiles 2007. 11: 2 245-250 doi: 10.1007/s00792-006-0032-3
    Kodach   LL , Bos   CL , Durán   N , . Violacein synergistically increases 5-fluoro-uracil cytotoxicity, induces apoptosis and inhibits Akt-mediated signal transduction in human colorectal cancer cells. Carcinogenesis 2017. 27: 508-516 doi: 10.1093/carcin/bgi307
    Konuray   G , Erginkaya   Z Méndez-Vilas   A . Antimicrobial and antioxidant properties of pigments synthesized from microorganisms. The battle against microbial pathogens: basic science, technological advances and educational programs 2015. Spain Formatex Research Center 27-33
    Koyama   J . Anti-infective quinone derivatives of recent patents. Recent Pat Anti-Infect Drug Discov 2006. 1: 113-125 doi: 10.2174/157489106775244073
    Kramar   A , Llic Tomic   T , Petkovic   M , . Crude bacterial extracts of two new Streptomyces sp. isolates as bio-colorants for textile dyeing. World J Microbiol Biotechnol 2014. 30: 2231-2240 doi: 10.1007/s11274-014-1644-x
Krishna JG (2008) Pigment production by marine Serratia sp. BTW J8. Doctoral dissertation, Ph. D. Thesis. Department of Biotechnology, Cochin University of Science and Technology, Cochin, Kerala, India
    Kushwaha   K , Saxena   J , Tripathi   BK , Agarwal   MK . Detection of carotenoids in psychrotrophic bacteria by spectroscopic approach. J BioSci Biotechnol 2014. 3: 3 253-260
    Leiva   S , Alvarado   P , Huang   Y , Wang   J , Garrido   I . Diversity of pigmented Gram-positive bacteria associated with marine macroalgae from Antarctica. FEMS Microbiol Lett 2015. 362: 24 fnv206 doi: 10.1093/femsle/fnv206
    Li   F , Xue   F , Yu   X . GC–MS, FTIR and Raman analysis of antioxidant components of red pigments from Stemphylium lycopersici. Curr Microbiol 2017. 74: 532-539 doi: 10.1007/s00284-017-1220-3
    Liao   L , Liu   C , Zeng   Y , Zhao   B , Zhang   J , Chen   B . Multipartite genomes and the sRNome in response to temperature stress of an Arctic Pseudoalteromonas fuliginea BSW20308. Environ Microbiol 2019. 21: 1 272-285 doi: 10.1111/1462-2920.14455
    Lim   J , Thomas   T , Cavicchioli   R . Low temperature regulated DEAD-box RNA helicase from the Antarctic archaeon Methanococcoides burtonii. J Mol Biol 2000. 297: 3 553-567 doi: 10.1006/jmbi.2000.3585
    Lin   SR , Weng   CF . PG-priming enhances doxorubin influx to trigger necrotic and autophagic cell death in oral squamous cell carcinoma. J Clin Med 2018. 7: 375 doi: 10.3390/jcm7100375
    Liu   Q , Liu   HC , Zhou   YG , Xin   YH . Microevolution and adaptive strategy of psychrophilic species Flavobacterium bomense sp. Nov. isolated from glaciers. Front Microbiol 2019. 10: 1069 doi: 10.3389/fmicb.2019.01069
    Lobo   V , Patil   A , Phatak   A , Chandra   N . Free radicals, antioxidants and functional foods: impact on human health. Pharmacogn Rev 2010. 4: 118-126 doi: 10.4103/0973-7847.70902
    Lorv   JS , Rose   DR , Glick   BR . Bacterial ice crystal controlling proteins. Scientifica 2014. doi: 10.1155/2014/976895
    Lu   Y , Wang   L , Xue   Y , . Production of violet pigment by a newly isolated psychrotrophic bacterium from a glacier in Xinjiang, China. Biochem Eng J 2009. 43: 2 135-141 doi: 10.1016/j.bej.2008.09.009
    Madigan   MT , Martinko   JM , Stahl   DA , Clark   DP Brock biology of microorganisms 2012. 13San Francisco Pearson Education, Inc
    Mageswari   A , Subramanian   P , Srinivasan   R , . Astaxanthin from psychrotrophic Sphingomonas faeni exhibits antagonism against food-spoilage bacteria at low temperatures. Microbiol Res 2015. 179: 38-44 doi: 10.1016/j.micres.2015.06.010
    Maheswarappa   G , Kavitha   D , Vijayarani   K , Kumanan   K . Prodigiosin as anticancer drug produced from bacteria of termite gut. Indian J Basic Appl Med Res 2013. 3: 257-266
    Manikprabhu   D , Lingappa   K . Actinorhodin a natural and attorney source for synthetic dye to detect acid production of fungi. Saudi J Biol Sci 2013. 20: 163-168 doi: 10.1016/j.sjbs.2013.01.004
    Margalith   PZ Pigment microbiology 1992. London Chapman & Hall 1-156
    Margesin   R , Collins   T . Microbial ecology of the cryosphere (glacial and permafrost habitats): current knowledge. Appl Microbiol Biotechnol 2019. doi: 10.1007/s00253-019-09631-3
    Margesin   R , Schumann   P , Spröer   C , Gounot   AM . Arthrobacter psychrophenolicus sp. Nov., isolated from an alpine ice cave. Int J Syst Evol Microbiol 2004. 54: 6 2067-2072 doi: 10.1099/ijs.0.63124-0
    Marizcurrena   JJ , Morel   MA , Braña   V , . Searching for novel photolyases in UVC-resistant Antarctic bacteria. Extremophiles 2017. 21: 2 409-418 doi: 10.1007/s00792-016-0914-y
    Marizcurrena   JJ , Cerdá   MF , Alem   D , Castro-Sowinski   S . Living with pigments: the colour palette of Antarctic life. The ecological role of micro-organisms in the Antarctic environment 2019. Cham Springer 65-82
    Martin-Cerezo   ML , Garcia-Lopez   E , Cid   C . Isolation and identification of a red pigment from the Antarctic bacterium Shewanella frigidimarina. Protein Peptide Lett 2015. 22: 1076-1082 doi: 10.2174/0929866522666150915122247
    Medentsev   AG , Arinbasarova   AY , Akimenko   VK . Biosynthesis of naphthoquinone pigments by fungi of the genus Fusarium. Appl Biochem Microbiol 2005. 41: 503-5076 doi: 10.1007/s10438-005-0091-8
    Medigue   C , Krin   E , Pascal   G , . Coping with cold: the genome of the versatile marine Antarctica bacterium Pseudoalteromonas haloplanktis TAC125. Genome Res 2005. 15: 10 1325-1335 doi: 10.1101/gr.4126905
    Melo   PS , Justo   GZ , de Azevedo   MB , Duran   N , Haun   M . Violacein and its beta-cyclodextrin complexes induce apoptosis and differentiation in HL60 cells. Toxicology 2003. 186: 217-225 doi: 10.1016/S0300-483X(02)00751-5
    Mojib   N , Philpott   R , Huang   JP , Niederweis   M , Bej   AK . Antimycobacterial activity in vitro of pigment isolated from Antartica bacteria. Anton Van Leeuwen 2010. 98: 531-540 doi: 10.1007/s10482-010-9470-0
    Mojib   N , Farhoomand   A , Anderson   DT , Bej   AK . UV and cold tolerance of a pigment- producing Antarctic Janthinobacterium sp. Ant5-2. Extremophiles 2013. 17: 367-378 doi: 10.1007/s00792-013-0525-9
    Moliné   M , Libkind   D , Dieguez   MC , van Broock   M . Photoprotective role of carotenoid pigments in yeasts: experimental study contrasting naturally occurring pigmented and albino strains. J Photochem Photobiol B 2009. 95: 156-161 doi: 10.1016/j.jphotobiol.2009.02.006
    Montagni   T , Enciso   P , Marizcurrena   JJ , . Dye sensitized solar cells based on Antarctic Hymenobacter sp. UV11 dyes. J. Environ Sustain 2018. 1: 1 89-97 doi: 10.1007/s42398-018-0007-1
    Mueller   DR , Vincent   WF , Bonilla   S , Laurion   I . Extremotrophs, extremophiles and broadband pigmentation strategies in a high arctic ice shelf ecosystem. FEMS Microbiol Ecol 2005. 53: 1 73-87 doi: 10.1016/j.femsec.2004.11.001
    Mykytczuk   NCS , Foote   SJ , Omelon   CR , Southam   G , Greer   CW , Whyte   LG . Bacterial growth at −15°C; molecular insights from the permafrost bacterium Planococcus halocryophilus Or1. ISME J 2013. 7: 1211-1226 doi: 10.1038/ismej.2013.8
    Nakamura   Y , Sawada   T , Morita   Y , . Isolation of a psychrotrophic bacterium from the organic residue of a water tank keeping rainbow trout and antibacterial effect of violet pigment produced from the strain. Biochem Eng J 2002. 12: 1 79-86 doi: 10.1016/S1369-703X(02)00079-7
    Namazkar   S , Ahmad   WA . Spray-dried prodigiosin from Serratia marcescens as a colorant. Biosci Biotech Res Asia 2013. 10: 1 69-76 doi: 10.13005/bbra/1094
    Nelis   HJ , De Leenheer   AP . Microbial sources of carotenoid pigments used in foods and feeds. J Appl Bacteriol 1991. 70: 3 181-191 doi: 10.1111/j.1365-2672.1991.tb02922.x
    Nigam   PS , Luke   JS . Food additives: production of microbial pigments and their antioxidant properties. Curr Opin Food Sci 2016. 7: 93-100 doi: 10.1016/j.cofs.2016.02.004
    Nwankwo   IU , Itaman   VO , Chidiebere   OL , Nwachukwu   MP . Evaluation of antimicrobial activity of prodigiosin produced from Serratia marcescens against some pathogenic bacteria. Futo J Ser 2017. 3: 93-102
    Ogugbue   CJ , Sawidis   T . Bioremediation and detoxification of synthetic waste water containing triarylmethane dyes by Aeromonas hydrophila isolated from industrial effluent. Biotechnol Res Int 2011. doi: 10.4061/2011/967925
    Okuyama   H , Orikasa   Y , Nishida   T . Significance of antioxidative functions of eicosapentaenoic and docosahexaenoic acids in marine microorganisms. Appl Environ Microbiol 2008. 74: 3 570-574 doi: 10.1128/AEM.02256-07
    Omamor   IB , Eziashi   EI , Adekunle   AA . Carbon nutrition in relation to growth of three Monascus species isolated from decaying date fruits. Afr J Microbiol Res 2008. 2: 153-155
    Onofri   S , Selbmann   L , Zucconi   L , Pagano   S . Antarctic microfungi as models for exobiology. Planet Space Sci 2004. 52: 229-237 doi: 10.1016/j.pss.2003.08.019
    Órdenes-Aenishanslins   N , Anziani-Ostuni   G , Vargas-Reyes   M , . Pigments from UV-resistant Antarctic bacteria as photosensitizers in dye sensitized solar cells. J Photochem Photobiol B 2016. 162: 707-714 doi: 10.1016/j.jphotobiol.2016.08.004
    Osawa   A , Ishii   Y , Sasamura   N , . Characterization and antioxidative activities of rare C50 carotenoids-sarcinaxanthin, sarcinaxanthin monoglucoside, and sarcinaxanthin diglucoside obtained from Micrococcus yunnanensis. J Oleo Sci 2010. 59: 653-659 doi: 10.5650/jos.59.653
Pagano MC, Dhar PP (2015) Fungal pigments: an overview. In: Gupta VK, Mach RL, Sreenivasaprasad S (eds) Fungal biomolecules: sources, applications and recent developments. 1st edn., Wiley, London. ISBN: 978-1-118-95829-2
    Pandey   N , Rahul   J , Anita   P , Sushma   T . Optimisation and characterisation of the orange pigment produced by a cold adapted strain of Penicillium sp. (GBPI_P155) isolated from mountain ecosystem. Mycology 2018. 9: 2 81-92 doi: 10.1080/21501203.2017.1423127
    Pantanella   F , Berlutti   F , Passariello   C , . Violacein and biofilm production in Janthinobacterium lividum. J Appl Microbiol 2007. 102: 4 992-999
    Panwar   AS , Molpa   D , Joshi   GK . Biotechnological potential of some cold-adapted bacteria isolated from North-Western Himalaya. Microbiology 2019. 88: 3 343-352 doi: 10.1134/S002626171903007X
    Parmar   M , Phutela   UG . Biocolors: the new generation additives. Int J Curr Microbiol Appl Sci 2015. 4: 7 688-694
    Parmar   TK , Rawtani   D , Agrawal   YK . Bioindicators: the natural indicator of environmental pollution. Front Life Sci 2016. 9: 2 110-118 doi: 10.1080/21553769.2016.1162753
    Perfumo   A , Banat   IM , Marchant   R . Going green and cold: biosurfactants from low-temperature environments to biotechnology applications. Trends Biotechnol 2018. 36: 3 277-289 doi: 10.1016/j.tibtech.2017.10.016
    Pezzella   A , Gargiulo   V , Alfè   M , . Eumelanin graphene-like integration: the impact on physical properties and electrical conductivity. Front Chem 2019. 7: 121 doi: 10.3389/fchem.2019.00121
    Phaniendra   A , Jestadi   DB , Periyasamy   L . Free radicals; properties, sources, targets, and their implication in various diseases. Indian J Clin Biochem 2015. 30: 11-26 doi: 10.1007/s12291-014-0446-0
    Piette   F , D’Amico   S , Mazzucchelli   G , . Life in the cold: a proteomic study of cold-repressed proteins in the Antarctic bacterium Pseudoalteromonas haloplanktis TAC125. Appl Environ Microbiol 2011. 77: 11 3881-3883 doi: 10.1128/aem.02757-10
    Poorniammal   R , Parthiban   M , Gunasekaran   S , . Natural dye production from Thermomyces sp. fungi for textile application. Indian J Fiber Textile Res 2013. 38: 276-279
    Potera   C . Diet and nutrition: the artificial food dye blues. Environ Health Perspect 2010. 118: 10 A428-A428 doi: 10.1289/ehp.118-a428
    Prabhu   S , Rekha   PD , Young   CC , . Zeaxanthin production by novel marine isolates from coastal sand of India and its antioxidant properties. Appl Biochem Biotech 2013. 171: 817-831 doi: 10.1007/s12010-013-0397-6
    Pradeep   FS , Pradeep   BV . Optimization of pigment and biomass production from Fusarium moniliforme under submerged fermentation conditions. Int J Pharm Pharm Sci 2013. 5: 526-535
    Prashanthi   K , Suryan   S , Varalakshmi   KN . In vitro anticancer property of yellow pigment from Streptomyces griseoaurantiacus JUACT 01. Braz Arch Biol Technol 2015. 58: 869-876 doi: 10.1590/S1516-89132015060271
    Puig-Marcó   R . Access and benefit sharing of Antarctica’s biological material. Mar Genomics 2014. 17: 73-78 doi: 10.1016/j.margen.2014.04.008
    Rafiq   M , Hayat   M , Anesio   AM , . Recovery of metallo-tolerant and antibiotic resistant psychrophilic bacteria from Siachen glacier, Pakistan . PloS one 2017. 12: 7 e0178180 doi: 10.1371/journal.pone.0178180
    Rafiq   M , Hayat   M , Zada   S , . Geochemistry and bacterial recovery from Hindu Kush Range glacier and their potential for metal resistance and antibiotic production. Geomicrobiology J 2019. 36: 4 326-338 doi: 10.1080/01490451.2018.1551947
    Rai   M Advances in fungal biotechnology 2009. New Delhi I. K. International Pvt Ltd. 545
    Ramesh   C , Vinithkumar   NV , Kirubagaran   R , . Multifaceted applications of microbial pigments: current knowledge, challenges and future directions for public health implications. Microorganisms 2019. 7: 7 186 doi: 10.3390/microorganisms7070186
    Rao   N , Prabhu   M , Xiao   M , Li   WJ . Fungal and bacterial pigments: secondary metabolites with wide applications. Front Microbiol 2017. 8: 1113 doi: 10.3389/fmicb.2017.01113
    Ravikumar   S , Uma   G , Gokulakrishnan   R . Antibacterial property of Halobacterial carotenoids against human bacterial pathogens. J Sci Ind (India) 2016. 75: 253-257
    Raymond   JA , Christner   BC , Schuster   SC . A bacterial ice-binding protein from the Vostok ice core. Extremophiles 2008. 12: 5 713-717 doi: 10.1007/s00792-008-0178-2
    Reddy   GS , Prakash   JS , Prabahar   V , . Kocuria polaris sp nov, an orange-pigmented psychrophilic bacterium isolated from an Antarctic cyanobacterial mat sample. Int J Syst Evol Microbiol 2003. 53: 1 183-187 doi: 10.1099/ijs.0.02336-0
    Reichenbach   H , Kleinig   H , Achenbach   H . The pigments of Flexibacter elegans: novel and chemosystematically useful compounds. Arch Microbiol 1974. 101: 131-144 doi: 10.1007/BF00455933
    Reverchon   S , Rouanet   C , Expert   D , Nasser   W . Characterization of indigoidine biosynthetic genes in Erwinia chrysanthemi and role of this blue pigment in pathogenicity. J Bacteriol 2002. 184: 3 654-665 doi: 10.1128/JB.184.3.654-665.2002
    Rezaeeyan   Z , Safarpour   A , Amoozegar   MA , . High carotenoids production by a halotolerant bacterium, Kocuria sp. strain QWT-12 and anticancer activity of its carotenoids. EXCLI J 2017. 16: 840-851
    Robledo   JA , Murillo   AM , Rouzaud   F . Physiological role and potential clinical interest of mycobacterial pigments. IUBMB Life 2011. 63: 71-78 doi: 10.1002/iub.424
    Roos   JC , Vincent   WF . Temperature dependence of UV radiation effects on Antarctic cyanobacteria. J Phycol 1998. 34: 118-125 doi: 10.1046/j.1529-8817.1998.340118.x
    Rosa   LH , Vaz   AB , Caligiorne   RB , Campolina   S , Rosa   CA . Endophytic fungi associated with the Antarctic grass Deschampsia antarctica Desv. (Poaceae). Polar Biol 2009. 32: 2 161-167 doi: 10.1007/s00300-008-0515-z
    Ryazantseva   IN , Andreyeva   IN , Klementyeva   GS , . Pigment-dependent light influence on the energetics of Serratia marcescens. Thermochim Acta 1995. 251: 63-67 doi: 10.1016/0040-6031(94)02148-H
    Sabbe   K , Hodgson   DA , Verleyen   E , . Salinity, depth and the structure and composition of microbial mats in continental Antarctic lakes. Freshw Biol 2004. 49: 3 296-319 doi: 10.1111/j.1365-2427.2004.01186.x
    Sajjad   W , Ahmad   M , Khan   S , . Radio-protective and antioxidative activities of astaxanthin from newly isolated radio-resistant bacterium Deinococcus sp. strain WMA-LM9. Ann Microbiol 2017. 67: 7 443-455 doi: 10.1007/s13213-017-1269-z
Santiago M, Ramírez-Sarmiento CA, Zamora RA, Parra LP (2016) Discovery, molecular mechanisms, and industrial applications of cold-active enzymes. Front Microbiol 7(1408). 10.3389/fmicb.2016.01408
    Saviola   B . Pigments and pathogenesis. J Mycobact Dis 2014. 4: 1068-2161
    Schindler   PR , Metz   H . Bacteria of the Flexibacter/Sporocytophaga group and violet-pigmented bacteria as indicators for hygienically doubtful drinking water. Zentralblatt fur Hygiene und Umweltmedizin. Int J Hyg Envir Med 1989. 189: 1 29-36
    Schloss   PD , Allen   HK , Klimowicz   AK , . Psychrotrophic strain of Janthinobacterium lividum from a cold Alaskan soil produces prodigiosin. DNA Cell Biol 2010. 29: 9 533-541 doi: 10.1089/dna.2010.1020
    Schneemann   I , Wiese   J , Kunz   AL , Imhoff   JF . Genetic approach for the fast discovery of phenazine producing bacteria. Mar Drugs 2011. 9: 772-789 doi: 10.3390/md9050772
    Seganish   JL , Davis   JT . Prodigiosin is a chloride carrier that can function as an anion exchanger. Chem Commun 2005. 46: 5781-5783 doi: 10.1039/b511847f
    Segawa   T , Yoshimura   Y , Watanabe   K , . Community structure of culturable bacteria on surface of Gulkana Glacier. Alaska Polar Sci 2011. 5: 1 41-51 doi: 10.1016/j.polar.2010.12.002
    Selbmann   L , Zucconi   L , Isola   D , Onofri   S . Rock black fungi: excellence in the extremes, from the Antarctic to space. Curr Genet 2015. 61: 335-345 doi: 10.1007/s00294-014-0457-7
    Selvameenal   L , Radhakrishnan   M , Balagurunathan   R . Antibiotic pigment from desert soil actinomycetes; biological activity, purification and chemical screening. Indian J Pharm Sci 2009. 71: 499-504 doi: 10.4103/0250-474X.58174
    Shen   L , Liu   Y , Wang   N , . Variation with depth of the abundance, diversity and pigmentation of culturable bacteria in a deep ice core from the Yuzhufeng Glacier, Tibetan Plateau. Extremophiles 2018. 22: 1 29-38 doi: 10.1007/s00792-0170973-8
    Shindo   K , Misawa   N . New and rare carotenoids isolated from marine bacteria and their antioxidant activities. Mar Drugs 2014. 12: 1690-1698 doi: 10.3390/md12031690
    Shirata   A , Tsukamoto   T , Yasui   H , . Production of bluish-purple pigments by Janthinobacterium lividum isolated from the raw silk and dyeing with them. Nihon Sanshigaku Zasshi 1997. 66: 377-385
    Shirata   A , Tsukamoto   T , Yasui   H , . Isolation of bacteria producing bluish-purple pigment and use for dyeing. Jpn Agric Res Q 2000. 34: 131-140
    Shivaji   S , Ray   MK , Kumar   GS , . Identification of Janthinobacterium lividum from the soils of the islands of Scotia Ridge and from Antarctic peninsula. Polar Biol 1991. 11: 4 267-271 doi: 10.1007/BF00238461
    Siefirmann-Harms   D . The light-harvesting and protective functions of carotenoids in photosynthetic membranes. Physiol Plant 1987. 69: 501-568
    Sikkandar   S , Murugan   K , Al-Sohaibani   S , . Halophilic bacteria—a potent source of carotenoids with antioxidant and anticancer potentials. J Pure Appl Microbiol 2013. 7: 2825-2830
    Silva   TR , Tavares   RS , Canela-Garayoa   R , . Chemical characterization and biotechnological applicability of pigments isolated from Antarctic bacteria. Mar Biotechnol 2019. 21: 3 416-429 doi: 10.1007/s10126-019-09892-z
    Singgih   M , Andriatna   W , Damayanti   S , Priatni   S . Carotenogenesis study of Neurospora intermedia N-1 in liquid. J Chem Pharm Res 2015. 7: 842-847
    Singh   AK , Sad   K , Singh   SK , Shivaji   S . Regulation of gene expression at low temperature: role of cold-inducible promoters. Microbiology 2014. 160: 7 1291-1296 doi: 10.1099/mic.0.077594-0
    Singh   A , Krishnan   KP , Prabaharan   D , Sinha   RK . Lipid membrane modulation and pigmentation: a cryoprotection mechanism in Arctic pigmented bacteria. J Basic Microbiol 2017. 57: 9 770-780 doi: 10.1002/jobm.201700182
    Sinha   RP , Klisch   M , Gröniger   A , Häder   DP . Ultraviolet-absorbing/screening substances in cyanobacteria, phytoplankton and macroalgae. J Photochem Photobiol B 1998. 47: 2–3 83-94
    Sivaperumal   P , Kamala   K , Rajaram   R . Bioactive DOPA melanin isolated and characterized from a marine actinobacterium Streptomyces sp. MVCS6 from Versova coast. Nat Prod Res 2015. 29: 2117-2121 doi: 10.1080/14786419.2014.988712
    Smith   D , da Silva   M , Jackson   J , Lyal   C . Explanation of the Nagoya Protocol on access and benefit sharing and its implication for microbiology. Microbiology 2017. 163: 3 289-296 doi: 10.1099/mic.0.000425
    Solano   F . Melanin: skin pigments and much more-types, structural models, biological functions, and formation routes. New J Sci 2014. 2014: 1-28 doi: 10.1155/2014/498276
    Soliev   AB , Hosokawa   K , Enomoto   K . Bioactive pigments from marine bacteria: applications and physiological roles. Evid Based Complement Alternat Med 2011. doi: 10.1155/2011/670349
    Sommaruga   R , Libkind   D , Broock   M , Whitehead   K . Mycosporine-glutaminol-glucoside, a UV-absorbing compound of two Rhodotorula yeast species. Yeast 2004. 21: 1077-1081 doi: 10.1002/yea.1148
Stafsnes MH, Bruheim P (2013) Pigmented marine heterotrophic bacteria. In: Kim S (ed) Marine biomaterials: characterization, isolation and applications, CRC Press/Taylor & Francis Group, London, pp 117–148
    Stankovic   N , Senerovic   L , Ilic-Tomic   T , . Properties and applications of undecylprodigiosin and other bacterial prodigiosins. Appl Microbiol Biotechnol 2014. 98: 3841-3858 doi: 10.1007/s00253-014-5590-1
    Starič   N , Danevčič   T , Stopar   D . Vibrio sp. DSM 14379 pigment production—a competitive advantage in the environment?. Microb Ecol 2010. 60: 592-598 doi: 10.1007/s00248-010-9671-0
    Suba   KP , Stalin   A , Girija   AS , Raguraman   R . Homology modelling and docking analysis of prodigiosin from Serratia marcescens. Biotechnology 2013. 55: 12897-12902
    Subramaniam   S , Ravi   V , Sivasubramanian   A . Synergistic antimicrobial profiling of violacein with commercial antibiotics against pathogenic micro-organisms. Pharm Biol 2014. 52: 86-90 doi: 10.3109/13880209.2013.815634
    Suresh   M , Renugadevi   B , Brammavidhya   S , . Antibacterial activity of red pigment produced by Halolactibacillus alkaliphilus MSRD1—an isolate from seaweed. Appl Biochem Biotech 2015. 176: 185-195 doi: 10.1007/s12010-015-1566-6
    Suryawanshi   RK , Patil   CD , Borase   HP , . Towards an understanding of bacterial metabolites prodigiosin and violacein and their potential for use in commercial sunscreens. Int J Cosmet Sci 2015. 37: 1 98-107 doi: 10.1111/ics.12175
    Suryawanshi   RK , Patil   CD , Koli   SH , Hallsworth   JE , Patil   SV . Antimicrobial activity of prodigiosin is attributable to plasma-membrane damage. Nat Prod Res 2017. 31: 572-577 doi: 10.1080/14786419.2016.1195380
    Sutthiwong   N , Fouillaud   M , Valla   A , . Bacteria belonging to the extremely versatile genus Arthrobacter as novel source of natural pigments with extended hue range. Food Res Int 2014. 65: 156-162 doi: 10.1016/j.foodres.2014.06.024
    Tarangini   K , Mishra   S . Production of melanin by soil microbial isolate on fruit waste extract: two step optimizations of key parameters. Biotechnol Rep 2014. 4: 139-146 doi: 10.1016/j.btre.2014.10.001
    Tashirev   AB , Romanovskaia   VA , Shilin   SO , Chernaia   NA . Screening of yeast-producers of melanin in the Antarctic terrestrial biotopes. Mikrobiol Zh 2010. 72: 3-8
    Thomson   NR , Crow   MA , McGowan   SJ , . Biosynthesis of carbapenem antibiotic and prodigiosin pigment in Serratia is under quorum sensing control. Mol Microbiol 2000. 36: 539-556 doi: 10.1046/j.1365-2958.2000.01872.x
    Tian   B , Hua   Y . Carotenoid biosynthesis in extremophilic Deinococcus-Thermus bacteria. Trends Microbiol 2010. 18: 11 512-520 doi: 10.1016/j.tim.2010.07.007
    Tribelli   PM , Lopez   NI . Reporting key features in cold-adapted bacteria. Life (Basel) 2018. doi: 10.3390/life8010008
    Tribelli   PM , Solar Venero   EC , Ricardi   MM , . Novel essential role of ethanol oxidation genes at low temperature revealed by transcriptome analysis in the Antarctic bacterium Pseudomonas extremaustralis. PLoS ONE 2015. 10: 12 e0145353 doi: 10.1371/journal.pone.0145353
    Trochine   A , Turchetti   B , Vaz   ABM , . Description of Dioszegia patagonica sp. nov., a novel carotenogenic yeast isolated from cold environments. Int J Syst Evol Microbiol 2017. 67: 4332-4339 doi: 10.1099/ijsem.0.002211
    Tuli   HS , Chaudhary   P , Beniwal   V , Sharma   AK . Microbial pigments as natural color sources: current trends and future perspectives. J Food Sci Technol 2015. 52: 4669-4678 doi: 10.1007/s13197-014-1601-6
    Turk   M , Plemenitaš   A , Gunde-Cimerman   N . Extremophilic yeasts: plasma-membrane fluidity as determinant of stress tolerance. Fungal Biol 2011. 115: 10 950-958 doi: 10.1016/j.funbio.2011.04.006
    Umadevi   K , Krishnaveni   M . Antibacterial activity of pigment produced from Micrococcus luteus KF532949. Int J Chem Anal Sci 2013. 4: 149-152 doi: 10.1016/j.ijcas.2013.08.008
UNEP (2012) An update on biological prospecting in Antarctica and recent policy developments at the international level. ATCM XXXV Meeting. https://www.unep.org/dewa/Portals/67/pdf/ATCM35_ip063_e.pdf
UNU UNEP (2009) Biological prospecting: an update on recent policy developments at the international level. ATCM XXXII meeting. https://www.unep.org/dewa/Portals/67/pdf/Atcm32_ip091_e.pdf
    Van Duin   D , Casadevall   A , Nosanchuk   JD . Melanization of Cryptococcus neoformans and Histoplasma capsulatum reduces their susceptibilities to amphotericin B and caspofungin. Antimicrob Agents Chemother 2002. 46: 11 3394-3400 doi: 10.1128/AAC.46.11.3394-3400.2002
    Vasanthabharathi   V , Lakshminarayanan   R , Jayalakhshmi   S . Melanin production from marine Streptomyces. Afr J Biotechnol 2011. 10: 11224-11234 doi: 10.5897/AJB11.296
    Vasileva-Tonkova   E , Romanovskaya   V , Gladka   G , . Ecophysiological properties of cultivable heterotrophic bacteria and yeasts dominating in phytocenoses of Galindez Island, Maritime Antarctica. World J Microbiol Biotechnol 2014. 30: 1387-1398 doi: 10.1007/s11274-013-1555-2
    Vaz   AB , Rosa   LH , Vieira   ML , . The diversity, extracellular enzymatic activities and photoprotective compounds of yeasts isolated in Antarctica. Braz J Microbiol 2011. 42: 3 937-947 doi: 10.1590/S1517-83822011000300012
    Vendruscolo   F , Pitol   LO , Carciofi   BAM , . Construction and application a vanesystem in a rotational rheometer for determination of the rheological properties of Monascus ruber CCT 3802. J Biorheol 2010. 24: 29-35 doi: 10.1007/s12573-010-0019-7
    Venil   CK , Aruldass   CA , Dufosse   L , . Current perspective on bacterial pigments: emerging sustainable compounds with coloring and biological properties for the industry-an incisive evaluation. RSC Adv 2014. 4: 39523-39529 doi: 10.1039/c4ra06162d
    Venil   CK , Zakaria   ZA , Ahmad   WA . Bacterial pigments and their applications. Process Biochem 2013. 48: 1065-1079 doi: 10.1016/j.procbio.2013.06.006
    Venil   CK , Yusof   NZ , Aruldass   CA , . Application of violet pigment from Chromobacterium violaceum UTM5 in textile dyeing. Biologia 2016. 71: 2 121-127 doi: 10.1515/biolog-2016-0031
    Venil   CK , Wahidin   MAB , Aruldass   CA , Ahmad   WA . Production of bacterial pigments in low cost medium and formulation of biodegradable ink. Indian J Exp Biol 2017. 55: 7 441-447
    Vila   E , Hornero-Méndez   D , Azziz   G , . Carotenoids from heterotrophic bacteria isolated from Fildes Peninsula, King George Island, Antarctica. Biotechnol Rep 2019. 21: e00306 doi: 10.1016/j.btre.2019.e00306
    Villarreal   P , Carrasco   M , Barahona   S , . Tolerance to ultraviolet radiation of psychrotolerant yeasts and analysis of their carotenoid, mycosporine, and ergosterol content. Curr Microbiol 2016. 72: 94-101 doi: 10.1007/s00284-015-0928-1
    Vincent   WF , Downes   MT , Castenholz   RW , Howard-Williams   C . Community structure and pigment organisation of cyanobacteria-dominated microbial mats in Antarctica. Eur J Phycol 1993. 28: 4 213-221 doi: 10.1080/09670269300650321
    Visalakchi   S , Muthumary   J . Antimicrobial activity of the new endophytic Monodictys castaneae SVJM139 pigment and its optimization. Afr J Microbiol Res 2010. 4: 38-44
    Voets   IK . From ice-binding proteins to bio-inspiredantifreeze materials. Soft Matter 2017. 13: 28 4808-4823 doi: 10.1039/C6SM02867E
    Wackenroder   H . Ueber das Oleum radicis Dauci aetherum, das Carotin, den Carotenzucker und den officinellen succus Dauci; so wie auch über das Mannit, welches in dem Möhrensafte durch eine besondere Art der Gahrung gebildetwird. Geigers Magazin Pharmazie 1831. 33: 144-172
    Wada   N , Sakamoto   T , Matsugo   S . Multiple roles of photosynthetic and sunscreen pigments in cyanobacteria focusing on the oxidative stress. Metabolites 2013. 3: 2 463-483 doi: 10.3390/metabo3020463
    Walford   J “Historical development of food colouration”, in evelopments in Food Colours 1980. London Applied Science publishers
Wan A, Rozi AM, Bin MI et al (2014) A method for dyeing polyester fabrics using colorant extracted from Chromobacterium violaceum. MY150969 (A) (Patent).
    Wang   Y , Nakajima   A , Hosokawa   K , . Cytotoxic prodigiosin family pigments from Pseudoalteromonas sp. 1020R isolated from the Pacific coast of Japan. Biosci Biotechnol Biochem 2012. 76: 1229-1232 doi: 10.1271/bbb.110984
    Wehrs   M , Gladden   JM , Liu   Y , . Sustainable bioproduction of the blue pigment indigoidine: expanding the range of heterologous products in R. toruloides to include non-ribosomal peptides. Green Chem 2019. 21: 12 3394-3406 doi: 10.1039/C9GC00920E
Wong LS, Teo SC (2014) Naturally occurring carotenoids in cyanobacteria as a bioindicator for heavy metals detection. In Proceedings of the international Conference on Advances. In: Applied Science and Environmental Engineering–ASEE.
World Health Organization (WHO) (2018) The top 10 causes of death. https://www.who.int/news-room/fact-sheets/detail/the-top10-causes-of-death. Accessed 1 Jun 2018
    Wynn-Williams   DD , Edwards   HGM Horneck   G , Baumstark-Khan   C . Environmental UV radiation: biological strategies for protection and avoidance. Astrobiology 2002. Heidelberg Springer
    Wynn-Williams   DD , Edwards   HGM , Garcia-Pichel   F . Functional biomolecules of Antarctic stromatolitic and endolithic cyanobacterial communities. Eur J Phycol 1999. 34: 4 381-391 doi: 10.1080/09670269910001736442
    Yamashita   M , Nakagawa   Y , Li   H , Matsuyama   T . Silica gel-dependent production of prodigiosin and serrawettins by Serratia marcescens in a liquid culture. Microbes Environ 2001. 16: 250-254 doi: 10.1264/jsme2.2001.250
    Yarzábal   LA . Antarctic psychrophilic microorganisms and biotechnology: history, current trends, applications, and challenges. Microbial models: from environmental to industrial sustainability 2016. Singapore Springer 83-118
    Yoshida   K , Hashimoto   M , Hori   R , . Bacterial long-chain polyunsaturated fatty acids: their biosynthetic genes, functions, and practical use. Mar Drugs 2016. 14: 5 94 doi: 10.3390/md14050094
    Yu   D , Xu   F , Valiente   J , . An indigoidine biosynthetic gene cluster from Streptomyces chromofuscus ATCC 49982 contains an unusual IndB homologue. J Ind Microbiol Biot 2013. 40: 1 159-168 doi: 10.1007/s10295-012-1207-9
    Yumusak   C , Prochazkova   AJ , Apaydin   DH , . Indigoidine-biosynthesized organic semiconductor. Dyes Pigm 2019. 171: 107768 doi: 10.1016/j.dyepig.2019.107768
    Yurkova   AM , Vustin   MM , Tyaglov   BV , . Pigmented basidiomycetous yeasts are a promising source of carotenoids and ubiquinone Q10. Microbiology 2008. 77: 1-6 doi: 10.1134/S0026261708010013
    Zerrad   A , Anissi   J , Ghanam   J , . Antioxidant and antimicrobial activities of melanin produced by a Pseudomonas balearica strain. J Biotechnol Lett 2014. 5: 87-94
    Zhang   X , Ma   X , Wang   N , Yao   T . New subgroup of Bacteroidetes and diverse microorganisms in Tibetan plateau glacial ice provide a biological record of environmental conditions. FEMS Microbiol Ecol 2008. 67: 21-29 doi: 10.1111/j.1574-6941.2008.00604.x
    Zobell   C , Upham   HC . A list of marine bacteria including descriptions of new species. BidL. ScrippsIns£. Oceanogr 1944. 5: n9-292
This article is made available via the PMC Open Access Subset for unrestricted research re-use and secondary analysis in any form or by any means with acknowledgement of the original source. These permissions are granted for the duration of the World Health Organization (WHO) declaration of COVID-19 as a global pandemic.
https://www.researchpad.co/tools/openurl?pubtype=article&doi=10.1007/s00792-020-01180-2&title=Pigment production by cold-adapted bacteria and fungi: colorful tale of cryosphere with wide range applications&author=Wasim Sajjad,Ghufranud Din,Muhammad Rafiq,Awais Iqbal,Suliman Khan,Sahib Zada,Barkat Ali,Shichang Kang,&keyword=Cryosphere,Microbial pigments,Cold-adapted microbes,Carotenoid,Melanin,&subject=Review,