5---------------

color affects every bit of life, including the clothes we wear, the furniture in our home, and the allure of
food (Downham and Collins, 2000; Manikprabhu and Lingappa, 2013). Just think, for instance, how
plants could prepare their own food without chlorophyll or how oxygen could be carried in the body
without hemoglobin. It can be said that life on earth depends on pigments (Britton, 1995).

(Prabhu et al. 2017).

Every aspect of Life is affected by colors ranging from clothes we wear to the furniture in our homes and
the food we eat. Have you ever thought for a second how would it look if plants synthesize their food
without chlorophyll or how oxygen can be supplied to the body without the requisite hemoglobin. So,
suffice to say life on earth depends on pigments in one way or other (Prabhu et al. 2017) including food ,
clothes, painting, cosmetics, paintings and pharmaceuticals etc. but due to unrestricted use of synthetic
pigments , severe hazards have fell upon human health along with being a threat to environment. Hence
the need to find an alternative, biodegradable , natural source of pigemnts is important to find. (Ahmad
et al. 2011). A lot efforts have been made in this regard to replace artificially made pigments with
naturally produced pigments (Tuli, Chaudhary, and Beniwal 2015). As nature is a great source organisms
producing colourful pigments. Recently, research have also shown the markert value of natural pigments
over syntheric one. Approximately $600 millions equal to 29% up from 2007 was announced in 2011 by
global sale of natural coloures (Tuli, Chaudhary, and Beniwal 2015).

17---------------

Pigments are widely used in food, cloth, painting, cosmetics, pharmaceuticals, and plastics. However,
uncontrolled industrial applications of synthetic pigments may result in serious hazard to human health
and a potential environmental disaster. Therefore, it is imperative to find an alternative pigment which
can be biodegraded and easily available at a minimal production cost. Natural pigments are
biodegradable [1] (Ahmad et al. 2011)

13--------------

Many research efforts have been made to replace synthetic pigments with natural pigments (Tuli,
Chaudhary, and Beniwal 2015)

13----------------

because nature is a rich source of colored pigment producing organisms including plants, animals and
microorganisms. Recent Research has prominently projected the value of natural colors over that of
artificial/syntheticcolors.In2011,global sales of natural colors amounted to an estimated $600 millions,
up by almost 29 % from 2007, depicting annual growth in excess of 7 %.(Tuli, Chaudhary, and Beniwal
2015)

19--------------

Conventiona Sources of Pigments l

Conventional sources of pigments range from inorganic metals and metal oxides to organic molecules.
Paleolithic humans made paint 30,000 years ago by grinding up earth pigments such as red ochre, which
gets its color from hematite (Fe2O3), yellow ochre (Fe2O3?H2O), and green malachite (Cu2CO3(OH)2)
and suspending these in their own saliva as a binder [2]. These prehistoric pigments carved out a place
on the modern artist’s palette and have been joined by other common inorganic pig- ments including
titanium white (TiO2), cobalt blue (CuO + Al2O3), Egyptian blue (CuONCaONSiO2), cadmium red (CdSe),
and black oxide of iron (FeO). The source ofdelocalized electrons in these pigments is the metal–ligand
complex.

Traditional sources of organic pigments and dyes include natural products such as flavinoids (Figure 1,
compound A) and anthraquinones (Figure 1, compound B) produced by plants and animals. For example,
carminic acid, a deep red anthraquinone produced by scale insects, is now used as a pigment in paints,
crimson ink, cosmetics, and food colors [3]. Over the course of the 20th century, naturally occurring
organic pigments have been almost completely displaced by synthetic molecules such as phthalocya-
nines that range from blue to green (Figure 1, compound C), arylides that are yellow to greenish- or
reddish-yellow (Figure 1, compound D), and quinacri- dones, ranging from orange to violet (Figure 1,
compound E) [4]. Advances in organic chemistry enabled mass produc- tion of these compounds
relatively cheap- ly, thereby allowing them to displace natural product pigments, whose procurement is
often more challenging.
 Figure 1. Structures of natural and synthetic pigments. (A) flavone, a yellow pigment produced
by paints; (B) carminic acid, a red pigment produced by scale insects; (C) copper phthalocyanine-
blue, a blue pigment of the synthetic phthalocyanine family; (D) benzidine yellow, an azo
pigment of the synthetic arylide family; (E) PV19, a red pigment of the synthetic quinacridone
family. Each molecule is colored to match its perceived color.
doi:10.1371/journal.pbio.1000510.g001
(Charkoudian et al. 2010)

28---------------
In nature, a great number of microorganisms e.g. yeast, fungi, algae, protozoa and bacteria
produce pigments (Fig. 1). Nevertheless, appropriate species should satisfy some criteria: i)
ability to use a variety of C and N sources, ii) tolerance to growth conditions (e.g. pH,
temperature, nutrient concentration), iii) providing expected color at reasonable yield, iv) non -
toxic and non-pathogenic product, and v) simple extraction of pigment from cell biomass if not
excreted [2]
(Pankaj and Kumar 2016)
20-------------------
Current sources of opigments
There are different sources of natural pigments (Pigments and Heer 2017)

4-----------
Plants, animals, and microbes are the sources of natural pigments.

(Masi et al. 2014)

20-------------------
Plants Pigments:
Any type of colored substance produced by the plants is called as “Plant Pigment”.
Photosynthesis is the primary function of pigments in plant, which uses the green pigment
chlorophyll. Carotenoids are also essential for their functions in photosynthesis 12. There are
different types of plant pigments, found in different classes of organic compounds. Plant
 pigments can give color to leaves, flowers and fruits. Some of the naturally derived pigments
from plant sources have been summarized in Table 1. (Pigments and Heer 2017)

5---------------

The use of plant pigments has many drawbacks such as non-availability throughout the year and pigment
stability and solubility. Large scale plant use may lead to loss of valuable species. For these reasons, the
process may not consider viable (Downham and Collins).

(Prabhu et al. 2017).

20----------------

? Animals Pigments: Even animals produce bichroms or biological pigments, as mentioned

earlier. In most of the plants chlorophyll is the primary biological pigment, and in mammals
melanin is the main biochrome that is found. However, melanin is responsible for the color of
hair and fur of an animal. A small parasitic insect i.e Cochineal beetle, Latin name Dactylopius
Confusus, that lives and dines on the prickly pear cactus, or nopal, in spanish. Cactus is a plant
from which the insects or, grana are harvested and ground into a red pulp color. A brilliant red
liquid pigment produced from their blood is used by the ancient Meso- Americans to make a
beautiful steadfast dye. Through the world, Cochineal is still used in many products. It is
commonly used in red lipsticks, and is one of the few red pigments allowed to be used in eye
shadow. Further, the color additive used in Cherry Coke is also made from cochineal 13.
? Microbial Pigments: The natural pigments
extracted from microorganism are termed as “microbial pigments”. Table 2 shows the naturally
derived pigments from microorganism. Microorganism’s bacteria, algae and fungi produce
variety of pigments and therefore, are the promising source of food colorants 14, 15. These
pigments from biological or microbial sources have desirable properties like stability to light heat
and pH 11. Microbial pigments possesses anti-cancer properties and are a source of pro-Vitamin
A. Microbial production has various benefits as their production is independent to weather
condition, easy and fast with different colors uses. Hence, microbial production of colorant has
many advantages over the other colorants as they can be produced under controlled condition
in a very less time.
 (Pigments and Heer 2017)

4-----------

Microbial pigment production is a recent phenomenon.They display all the colors from rainbow including
light or dark tinges and unusual colors like black, white, brown, golden, silver and fluorescent green,
yellow or blue. (Masi et al. 2014)

21------------

History

Addition of colour to the processed foods is an old practice but microbial pigments production is a recent
phenomenon (Table 1). Saffron, turmeric and vegetable dyes had been used to colour food (Singhal &
Kulkarni, 1999). Nata, a microbial coloured food, is one of the traditional holiday foods in Philippines and
is also popular in Indonesia, Japan and Taiwan.
 (Joshi and Bhushan 2014)

Addition of colour to the processed foods is an old practice but microbial pigments production is a recent
phenomenon (Table 1).

11------------

Microbial pigments or biopigments are multitude of chemical structures capable of absorbing light in the
visible range (400–700 nm). There is an ever-growing number of biopigment

(Carvalho et al. 2014)

9---------------
Chemical composition of pigments and growth conditions
Pigments diversity depends on differences in their chemical structure and also the presence of
special chromophores (10). An unsaturated organic molecule contains a group of single or
double bonds that serves the changes of light absorption to lower frequencies (longer wave
lengths). The presence of these substances, results in the creation of the desired color. Indeed,
the assortment of these materials, which is commonly called root color, creates the extended
spectrum colors like Beta Carotene, Lycopene and Anthocyanin (64). Pigment production and
biosynthesis are associated with a variety of factors like the environmental and culture
conditions, PH, inorganic phosphate concentration, temperature, etc. (75). For instance
production of prodigiosin pigment will increase in Peptone Glycerol Broth and Nutrient Broth
from 28°C up to 30°C (20,37).
(Sabbagh and Ebrahimzadeh Namvar 2017)
8------------------

Bacterial pigment production is now one of the emerging field of research to demonstrate its potential
for various industrial applications.8 Most of the bacterial pigment production is still at the R&D stage.
Hence, work on the bacterial pigments should be intensified especially in finding cheap and suitable
growth medium which can reduce the cost and increase its applicability for industrial production.
(Usman et al. 2017)

8---------------

Bacterial pigments offer the following benefits and advantages as follows:

 . Increasingly attractive to science because of broad ranging ac- tivities.

 Easy propagation and wide strain selection.
 . High versatile and productive over other sources.
 Fermentation is inherently faster and more productive compa- red to any other chemical
process.
 Easy to manipulate genes.
 Simple and fast culturing techniques allowing continuous bio- reactor operation.
 Structural complexity suits for industrial needs.
 Bacterial pigments extracted using simple liquid–liquid extrac- tion technique minimizing
operation cost.
 Cheap substrates used for bulk production
(Usman et al. 2017)

28--------------------

Pigment producing microorganisms like, bacteria offers certain distinctive advantages, owing to their
short life cycle, non-sensibility to season and climate, potentiality to produce pigments of different colors
and shades, easier scale-up production of pigments, etc.

(Pankaj and Kumar 2016)

5----------------
Bacterial Pigments The use of bacteria for pigment production has several
advantagesoverfungi,suchasshortlifecycleandeaseforgenetic modification (Venil et al., 2013,
2014). However, compared with fungal pigments, most of bacterial pigments are still at the
research and development stage (Table 2); hence, work on bacterial pigments production should
be intensified to make them available on the market. Pigment producing bacteria are ubiquitous
and present in various ecological niches, such as soil (Zhu et al., 2007), rhizospheric soil (Peix et
al., 2005), desert sand (Liu et al., 2009), fresh water (Asker et al., 2008), and marine samples
(Franks et al., 2005). They were reported in low (Nakamura et al., 2003) and high (Manachini et
al., 1985) temperatureregions,canpersistinsaltregions(AskerandOhta,
1999),andevenasendophytes(Dengetal.,2011).
Compared with other bacterial groups, the pigment production is more likely to be present in
actinobacteria (Marroquin and Zapata, 1954). Various genera such as Streptomyces, Nocardia,
Micromonospora, Thermomonospora, Actinoplanes, Microbispora, Streptosporangium,
Actinomadura, Rhodococcus, and Kitasatospora (Rana and Salam, 2014) produce a wide variety
of pigments. The genus Streptomyces was reported for highest pigment production (Conn and
Jean, 1941). Many species of this genus, like Streptomyces griseus, Streptomyces griseoviridis,
Streptomyces coelicolor (Darshan and Manonmani, 2015), Streptomyces cyaneus (Petinate et al.,
1999),Streptomycesvietnamensis(Zhuetal.,2007),Streptomyces peucetius (Arcamone, 1998),
Streptomyces echinoruber (Gupta et al., 2011), Streptomyces shaanxiensis (Lin et al., 2012), and
Streptomyces caeruleatus (Zhu et al., 2011) were reported to producepigments.
(Prabhu et al. 2017)