Open access peer-reviewed chapter

Studies on Colored Cotton: Biochemical and Genetic Aspects

Written By

Sathees Nagarajan, Yazhni Purushothaman, Monika Selvavinayagam, Pandidurai Govindharaj and Aasif Musthafa

Submitted: 04 April 2022 Reviewed: 12 April 2022 Published: 05 October 2022

DOI: 10.5772/intechopen.104898

From the Edited Volume

Cotton

Edited by Ibrokhim Y. Abdurakhmonov

Chapter metrics overview

170 Chapter Downloads

View Full Metrics

Abstract

Cotton (Gossypium hirsutum L.) is a commercially important fiber crop used as the primary raw material in the textile industry and is cultivated throughout the world. Normally cotton fiber is white color and various dyes are used to color the fiber. In textile industry, the process of artificial dying is a major source of pollution to the environment and the cost of dying is also higher. Apart from the white fiber, several cotton species have colored fiber which can be used to reduce the dying process and its ill effects to the environment. The cotton fiber color inheritance pattern is an urgent problem. The physical and chemical properties of colored cotton are determined by its chemical composition. The naturally colored cotton contain some important properties such as, greater hygiene, hypoallergenic properties, lower flammability and higher ultraviolet protection value compared to traditional white cotton. The natural colored cotton loss their market value due to the poor fiber quality. Understanding of the colored cotton pigment composition, biochemical and genetic prospects of colored cotton will be useful for the development of high quality of colored cotton.

Keywords

  • colored cotton
  • fiber quality
  • colored pigment
  • biochemical and genetic property

1. Introduction

In world, cotton is an important cash crop and it is a most traded commodity [1]. China, India, United States, Pakistan, and Brazil are the largest cotton producers [2]. With 312 lakh bales, India has the world’s largest cotton area of around 12.7 million hectares and is now the world’s second largest cotton producer (each of 170 kg) [3]. The cotton is a dicotyledon comes under the malvaceae family and Gossypium genus. Globally, Gossypium genus is spread in 5 continents. It contains 50 species in the world which are woody and herbaceous form [4]. The 50 species contain 45 diploid (2 n = 2x = 26) and five allotetraploid (2 n = 4x = 52) species [4]. Among these two diploid species (G. arboreum L. and Gossypium herbaceum L.) and two tetraploid species (G. hirsutum L. and Gossypium barbadense L.) comes under cultivated species. The 95 percentage of world cotton production was fulfilled by two tetraploid cotton cultivars such as G. hirsutum L. (upland cotton) and G. barbadense L. (Sea Island or Egyptian cotton) because it contains good fiber yield and broad adaptation to several environments [5]. In global fiber market, the polyester and other synthetic fibers have the robust competition during 1990 s which increases the competition in beginning of 2000 s [1]. Most of textile products are manufactured by cotton fiber and lint [6]. In Gossypium genus, the formation of fibers is abnormal. The surface of the ovule has the outward elongated cells growing. Additionally, as the fiber matures, the protoplast dies, and the cell wall collapses inward to form a convoluted ribbon [5]. One of the most significant raw materials for the textile industry is cotton (Gossypium hirsutum L.). Nearly all cotton fiber used for textiles is white, dyes are necessary throughout the fiber processing process to color the cloth. The massive usage of dyes has resulted in pollution, which has had a significant impact on human health [7]. Cotton that is naturally colored is made up of pigmented fibers with color embedded in the lumen [8]. The colorful strain of G. hirsutum is crossed with a white linted strain to create hybrids that outperform the color parents in terms of fiber length, strength, and color fastness. Natural colored cotton will be the next big thing in the market as the world shifts toward pollution-free organic fabrics and products. This is due to the fact that the production of naturally colored cotton avoids the most polluting activity of textile product manufacturing (dyeing) [9]. In ecology textiles, naturally colored cotton is an important raw material which eliminates the dying during the processing. It would significantly decrease the processing cost, environmental pollution and chemical residue [8, 10]. Furthermore, when compared to standard white cotton, naturally colored cotton may have a reduced flammability and a greater UV protection rating [11]. One of the most efficient solutions is to breed cotton varieties that naturally contain colored cotton fibers (CCFs), which are environmentally safe. Cotton plants with colorful fibers have been cultivated for a long time [12]. However, for the following two reasons, their development has been slower than that of white fiber cotton. To begin with, colored fiber cotton yields far less than white fiber cotton [13]. Various dye products that have been employed in the textile industry since the industrial revolution are available, as well as the negative consequences of their use has long been forgotten [14]. Cotton fiber is the most important fabric material on the planet, with almost all the industrially used cotton coming from white cotton fiber (WCF). However, with rising environmental concerns and improved human life quality, interest in naturally colored cotton (NCC, G. hirsutum) fiber has steadily increased over the previous decade. Natural colors are used in NCC fibers. The use of NCC fiber in fabrics would cut textile processing and the generation of harmful chemical wastes significantly [15]. Nonetheless, poor fiber quality and drab colors have hampered NCC fiber adoption on a big scale [16]. The genetics and plant breeding of cotton goal is improving fiber quality while increasing the cotton yield [17]. Fiber quality is a complex trait which includes fiber length, strength, and fineness. The following traits determine the cotton yield such as number of bolls per plant, number of plants per unit area, lint percentage, and single boll weight [18]. The fiber quality and yield have the negative correlation, so the synchronous improvement of that traits through conventional breeding techniques is difficult. So, understanding of the colored cotton biochemical and molecular aspects will be used to improve that cotton.

Advertisement

2. Genes regulating pigmentation and its inheritance pattern

Knowledge on the genes responsible for pigmentation can help the breeders to develop colored cotton and to overcome the barriers in developing it. The natural colored lint of cotton shows great variability for lint color of which brown and green are the most stable and predominant types. The pigments are accumulated in the lumen of the lint [19] only when exposed to sunlight [20]. The developed color fades out when exposed to sunlight for a longer time and moisture content also affects the developed color [15]. The poor fiber quality viz., fiber length, fiber percent and resistance is attributed to the pleiotropic effects of genes controlling fiber quality [21]. Presence of modifying genes also affects the color and quality of pigments. By expression analysis, GhF3’5’H and GhCHS3 genes were found to be higher in colored cottons than in white cotton [22].

Green color results due to the deposition of caffeic acid in the suberin layer. Brown pigmentation results due to the pro anthocyanidin (PA) (condensed tannin) accumulation in cell vacuoles [23]. The prime sequences in PA biosynthesis were highly conserved in both white (G. arboreum) and brown (G. raimondii or G. stocksii) fibers [24]. Earlier it was reported that both the green and brown pigments are governed by single gene with incomplete dominance [25]. Later, in 1944, conventional approaches revealed the existence of six loci (Lc1- Lc6) for brown pigment, out of which Lc1 was located on chromosome 7, Lc2 on chromosome 6 and one locus (Lg) for green pigment. Further, colored cotton is dominant over white fiber cotton [26]. But in texas green and brown lint, they are controlled by single incompletely dominant gene [25]. Generally, these genes are pleiotropic in nature [10].

Genetically, green color genes are dominant over both brown and white colors in cotton. From the findings of several researchers, it is evident that, the brown color is controlled monogenically with incomplete dominance [27]. Also, the brown lint and brown fuzz color was found to be correlated and controlled by single gene with incomplete dominance [28, 29]. Further investigation advocated that the GhTT2-AO7 gene of LC 1 controls the brown color fiber trait [30]. Additionally, in brown color cottons, the flavanoid genes (GhCH1, GhF3H, GhDFR, GhANS & GhANR) were involved in proanthocyanidin flavanoid production which is responsible for the brown pigmentation [22]. The gene Gh3GT coding flavonoid 3- glucosyltransferase leads to green color even in brown fiber cotton [31]. Flavonoid biosynthesis is regulated by several transcription factors such as R2R3-MYB type factors, basic helix- loop- helix and WD40 repeats [32]. The genes GhMYB10 and GhMYB36, homologous to genes which encode the R2R3-MYB type transcription factors were noticed in cotton and they enhance the PA synthesis [33]. GhTT2-A07 and GhTT2-3A also involve in the production of brown pigment. GhTT2-3A and GhbHLH130D drive the structural genes GhANR and GhLAR to accumulate PA in fiber [30]. Also, the genes GhTTG1 and GhTT3 genes play an inevitable role in PA synthesis and fiber development [34]. The genes which are responsible for anthocyanin pigment were also found to act in the regulation of PA synthesis [35].

Caffeic acid biosynthesis in green fiber takes place via phenyl propanoid pathway. The expression of the gene GhPAL was found higher at the initiation stage of secondary cell wall thickening. The genes Gh4CL1- GhCL4 convert the caffeic acid to its ester form. Out of these four above mentioned genes, Gh4CL2 showed higher expression in green fiber, which was confirmed by its expression level and enzymatic activity studies [36].

Advertisement

3. Biosynthetic pathway/molecular basis of pigment synthesis and deposition

In brown fiber cotton, the amount of oxidized pro anthocyanidin increases with the maturation of bolls and its structure was observed to be modified by a galloyl group [37]. MALDI- TOF MS proteomic analysis [38] in brown fiber revealed that out of 21 proteins responsible for pigmentation, 15 were the members of flavonoid biosynthesis process. PAs are polymers of polyhydroxy flavan-3-ol units and addition of leucoanthocyanidin (flavan-3, 4-diol) molecules. Digital gene expression (DGE) analysis showed that 34 PA synthase genes are involved out of which only 24 were upregulated [39]. These upregulated genes coding for the enzymes involved in the synthesis of PA including, 3- phenylalanine ammonia lyases (PAL), cinnamic acid-4-hydroxylase (C4H), 1, 4- coumarate CoA ligase (4CL), chalcone synthase (CHS), chalcone isomerase (CHI), flavone 3- hydroxylase (F3H), flavonoid 3′ hydroxylase (F3’H) and flavonoid 3′ 5′- hydroxylase (F3’5’H), dihydroflavonol 4- reductase (DFR), anthocyanidin synthase (ANS) and anthocyanidin reductase (ANR) [39]. Out of these, CHI plays a major role in the coloration of brown fiber [31]. Individual biochemical pathways were identified in brown fiber cotton some involving the major role of ANR [23] and some with major role of leucoanthocyanin reductase (LAR) [39]. Also, the activity of PAL was found to be higher in brown fiber than white ones [40]. The accumulation of PA in the brown fiber was found to be in peak at 30 DAS and decreased due to their oxidation toward maturation [37]. The biosynthetic pathway involved in the synthesis of pro anthocyanins is illustrated in Figure 1.

Figure 1.

Biosynthesis of pro anthocyanins [41].

Green color in fibers is the result of Caffeic acid (CA) accumulation. Nearly 70% ω- hydroxydocosanoic acid and 25% decosanedoic acid, which are the components of caffeic acid, were isolated from green cotton whereas only 0.5% was reported in white fiber. UV and Nuclear magnetic resonance (H-NMR) spectroscopic studies revealed that wax portion of green fiber is mostly composed of glycerol, CA and its esterified form. Isolated fatty acids from green fiber showed the presence of 22-O-caffeol-22- hydroxydocosanoic acid and 22-O-caffeol-22- hydroxydocosanonin. They were responsible for green and yellow pigmentation and increasing the concentration of the latter, leads to deep green color [23]. Another important point to be noted was the suberization of seed coat in the plant, producing green fiber, while it was absent in white and brown cotton [42].

Advertisement

4. Characterization of colored cotton pigments

Comparing to white cotton, naturally colored cotton has more flavonoids, which would reach 1 mg/g at maturity. This forms the major portion of pigments synthesized in colored cotton. pH values also tend to vary in the colored and white cotton, which was 5.60 in white fiber and 5.63 in colored cotton at 30 days post anthesis (DPA). But in brown cotton, it rose to 6.07 at 35 DPA and may reach 6.38. Generally, a drop in pH, favors cell elongation and secondary cell wall thickening during fiber cell development and this rise in pH may lead to poor development of fiber [13]. Cellulose content of colored cotton differs from that of white cotton after 20 DPA and this may be due to the fact that flavonoid synthesis in CCF may make use of the available simple carbohydrates. This in turn may affect the quality and quantity of fiber [43]. Pigment development in green cotton takes more time than that of brown cotton. The fibers of naturally colored cotton give low lint yield, produce short, weak and coarse fiber. Also, the distribution of pigment may not be uniform [10]. The brown pigmented cotton fiber was found to be superior to green. In addition, the green pigment deposits in fiber during 15 to 20 days post anthesis period. While testing the amount of Nitrogen in the colored cotton, it was higher in colored cotton. Potassium level was lower in the colored cotton particularly in green. Correlation studies indicated negative relationship between the pigment with fiber quality parameters. The pleiotropic nature of the color genes inhibits the fiber development and this becomes the reason in the difficulty of developing colored cotton with good fiber quality [13]. Thus, cotton with high cellulose level, low N and P and high K levels with acceptable level of pigment is desirable.

Suberin lamella was present in the cell wall of green cotton fiber cells [44]. Presence of glycerol also has been found in the green cotton fibers. The presence of some yellow green pigments due to the presence of caffeic acid derivatives [23].

Advertisement

5. Evaluation of quality of colored cotton

Colored cotton fibers are currently available and can be combined with conventional white cottons. They are shorter, weaker, and finer than regular upland cotton fibers. Due to smaller bolls and low ginning outturn, color-linted cultivars are often low yielders with low productivity per unit area. Other issues with these cottons include high whiteness per cent, higher wax content, isolation distance requirements, the availability of only a few hues, and unpredictability and non-uniformity of fiber color across seasons and locales. To make these environmentally friendly color cottons commercially viable, researchers must focus their efforts on improving the genetics of agronomic features, fiber quality, and color uniformity [45]. The correct assessment of basic fiber properties and quality classification is a major issue for dealers, spinners, and farmers who are working to improve cotton production characteristics [46]. The degree of reflectance (Rd) and yellowness (+b) as specified by official criteria and measured by the high volume instrument determine the color grade (Figure 2). The equipment specification for Rd. and + b is specified in the Table 1. The brightness or dullness of a sample is determined by its reflectance, while the degree of pigmentation is determined by its yellowness. Finding the intersection of the Rd. and + b values on the color chart for American Upland cotton yields a three-digit color code [47]. The color of cotton fibers can be affected by rainfall, freezes, insects, fungi, and staining through contact with soil, grass, or cotton-plant leaf. Color can also be affected by excessive moisture and temperature levels during storage, both before and after ginning. Color deterioration because of environmental conditions affects the fibers’ ability to absorb and hold dyes and finishes and is likely to reduce processing efficiency [48].

Figure 2.

The cotton color grading instrument [46].

Fiber proppertyEquipment specifications
Precision specificationsCalibration tolerances
Color (Rd) (units)0.7000.400
Color (+b) (units)0.3000.400

Table 1.

Color cotton equipment specifications [47].

Several tools and programs are in place to manage quality. These include laboratory conditioning, sample conditioning, equipment performance specifications, instrument calibration, in-house monitoring, and USDA’s Quality Management Program.

5.1 Laboratory conditioning

The measurement of cotton fiber characteristics is influenced by atmospheric conditions. As a result, the classing laboratory’s temperature and humidity must be strictly controlled. The temperature is kept at 70 degrees Fahrenheit plus or minus 1 degree Fahrenheit (about 21 degrees Celsius plus or minus 1/2 degree Celsius), and the relative humidity is kept at 65 percent plus or minus 2 percent.

5.2 Sample conditioning

The moisture content of the samples is conditioned to match the permitted atmospheric conditions. Moisture level in conditioned samples will range from 6.75 to 8.25 percent (on a dry-weight basis). The moisture content of the conditioned samples is examined at random to ensure that the correct moisture content has been achieved. Samples can be passively or actively conditioned. The samples are put in single layers in trays for passive conditioning.

Advertisement

6. Breeding methods for the development of Colored cotton

Natural colored cotton cultivation dates back to 2300 BC [49]. Anciently, colored cottons were domesticated from G. hirsutum and G. barbadense. Subsequently, Due to the varied dyeing advantages in white cotton, the colored cotton was underrated and lost its preference among people and industrialist during the latter half of 19th century. But with the increasing environmental concern, demand for natural colored cotton gained its momentum during the past decade [16]. Generally the fiber color was negatively correlated with fiber yield, fiber quality [50] and limited color choice [51]. Correspondingly, the studies on genetics, inheritance and correlation of cotton fiber color and fiber yield, quality, pigmentation were stenuously carried out [29, 52]. These studies have led to the development of next generation colored cotton to overcome the short-comings encountered previously.

Primarily, colored cottons were identified as mutants of white cotton predominant from G. hirsutum and G. barbadense [53]. So far varieties of colored cotton were developed mostly by selection and recurrent crossing approaches from the germplasm [54]. Hybrids were also developed from crossing suitable germplasm with white cotton varieties to enhance the yield and fiber quality. Cocanada 1 & 2 and Red northers were the brown linted tree cotton varieties selected from G. arboretum. Vaidhehi 95 (MSH 53) is a introgressed cultivar developed from G. hirsutum which is also a brown linted variety produced by Central Intsitute for Cotton Research, Nagpur, India (CICR). CICR also produced some brown linted cotton varieties viz., CNA 405, CNA 406 and CNA 407 but no green linted cotton varities were developed so far. In order to develop more vibrant, diverse fiber color, high yielding, quality colored cotton varieties needs the combined usage of conventional and biotechnological method of plant breeding. Also integrating the results of omics studies on pigmentation of colored cotton would help in a big leap in developing colored cotton varieties [16].

Advertisement

7. Conclusion

To recapitulate, environmentalists are urging scientists and farmers to improve and grow NCC due to their concern about the effects of dyes and the advantages of naturally colored cotton over white cotton. Color development methods in green and brown cotton have been disclosed by the studies on the biochemical and molecular mechanisms of fiber color development in cotton and flavonoid biosynthesis genes. GhC4H, GhCHS, GhCHI, GhF3’H, GhDFR, and GhANR are the genes responsible for the structural flavonoid biosynthesis pathway that play a major role in color formation in brown fiber cotton. Cloning and genome-wide association studies of the fiber color genes help us comprehend the complicated biological mechanism of color development in cotton fiber which would have been hampered if the cotton genome sequences were not developed. Changing the cellulose and flavonoid biosynthesis pathways to improve fiber quality will be a step toward manufacturing cotton fiber with a wide range of hues and high quality for the textile industry. The recent improvements in several NCC research areas offer opportunities to overcome barriers to commercial NCC breeding, while further studies including multi-omics techniques are needed. The ability to reduce the fiber-quality and yield gap between NCC and white-fiber cotton will determine its growth in the textile market.

References

  1. 1. Barros MA, Silva CR, Lima LM, Farias FJ, Ramos GA, Santos RC. A review on evolution of cotton in Brazil: GM, white, and Colored cultivars. Journal of Natural Fibers. 2020;18:1-3
  2. 2. Statista. Cotton production by country worldwide in 2017/2018 (in 1, 000 metric tons). The Statistics Portal. 2018. Accessed September 12. https://www.statista.com/statistics/263055/cotton-production-worldwide-by-top-countries/
  3. 3. Meshram, JH, Mahajan SS, Nagrale D, Gokte-Narkhedkar N, Kumbhalkar H. Understanding Root Biology for Enhancing Cotton Production. In Plant Roots; IntechOpen: London, UK, 2021; p. 13
  4. 4. Fryxell PA. A revised taxonomic interpretation of Gossypium L. (Malvaceae). Rheedea. 1992;2(2):108-165
  5. 5. Lee JA, Fang DD. Cotton as a world crop: Origin, history, and current status. Cotton. 2015;57:1-23
  6. 6. Buainain AM, Alves E, Silveira JM, Navarro Z. O mundo rural no Brasil do século 21: a formação de um novo padrão agrário e agrícola. Brasília, DF: Embrapa, 2014;4:1125-1156
  7. 7. Weisburger JH. Comments on the history and importance of aromatic and heterocyclic amines in public health. Mutation Research, Fundamental and Molecular Mechanisms of Mutagenesis. 2002;506:9-20
  8. 8. Kimmel LB, Day MP. New life for an old Fiber: Attributes and advantages of naturally Colored cotton. Aatcc Review. 2001;1(10):32-35
  9. 9. Rathinamoorthy R, Parthiban M. Colored cotton: Novel eco-friendly textile material for the future. In: Handbook of Ecomaterials. Springer, New York; 2017. pp. 1-21
  10. 10. Murthy MS. Never say dye: The story of coloured cotton. Resonance. 2001;6(12):29-35
  11. 11. Crews PC, Hustvedt G. The ultraviolet protection factor of naturally-pigmented cotton. The Journal of Cotton Science. 2005;9:47-55
  12. 12. Yatsu LY, Espelie KE, Kolattukudy PE. Ultrastructural and chemical evidence that the cell wall of green cotton fiber is suberized. Plant Physiology. 1983;73(2):521-524
  13. 13. Dutt Y, Wang XD, Zhu YG, Li YY. Breeding for high yield and fibre quality in coloured cotton. Plant Breeding. 2004;123(2):145-151
  14. 14. Von Tunzelmann GN. Technology and Industrial Progress: The Foundations of Economic Growth. Edward Elgar Publishing limited, UK; 1995
  15. 15. Gong W, Du X, Jia Y, Pan Z. Color cotton and its utilization in China. In: Cotton Fiber: Physics, Chemistry and Biology. Springer, USA; 2018. pp. 117-132
  16. 16. Sun J, Sun Y, Zhu QH. Breeding next-generation naturally colored cotton. Trends in Plant Science. 2021;26(6):539-542
  17. 17. Naoumkina M, Thyssen GN, Fang DD, Jenkins JN, McCarty JC, Florane CB. Genetic and transcriptomic dissection of the fiber length trait from a cotton (Gossypium hirsutum L.) MAGIC population. BMC Genomics. 2019;20(1):1-4
  18. 18. Zhang D, Chen L, Zang C, Chen Y, Lin H. Antibacterial cotton fabric grafted with silver nanoparticles and its excellent laundering durability. Carbohydrate Polymers. 2013;92(2):2088-2094
  19. 19. Gong W, He S, Tian J, Sun J, Pan Z, Jia Y, et al. Comparison of the transcriptome between two cotton lines of different fiber color and quality. PLoS One. 2014;9(11):e112966
  20. 20. Rauf S, Shehzad M, Al-Khayri JM, Imran HM, Noorka IR. Cotton (Gossypium hirsutum L.) breeding strategies. In: Advances in Plant Breeding Strategies: Industrial and Food Crops. Springer, USA; 2019. pp. 29-59
  21. 21. Carvalho LP, Farias FJ, Lima MM, Rodrigues JI. Inheritance of different fiber colors in cotton (Gossypium barbadense L.). Crop Breeding and Applied Biotechnology. 2014;14(4):256-260
  22. 22. Xiao YH, Zhang ZS, Yin MH, Luo M, Li XB, Hou L, et al. Cotton flavonoid structural genes related to the pigmentation in brown fibers. Biochemical and Biophysical Research Communications. 2007;358(1):73-78
  23. 23. Feng H, Yang Y, Sun S, Li Y, Zhang L, Tian J, et al. Molecular analysis of caffeoyl residues related to pigmentation in green cotton fibers. Journal of Experimental Botany. 2017;68(16):4559-4569
  24. 24. Sun Y, Zhang D, Zheng H, Wu Y, Mei J, Ke L, et al. Biochemical and expression analyses revealed the involvement of Proanthocyanidins and/or their derivatives in fiber pigmentation of Gossypium stocksii. International Journal of Molecular Sciences. 2022;23(2):1008
  25. 25. Richmond TR. Inheritance of green and brown lint in upland cotton. Journal of the American Society of Agronomy. 1943;35(11):967-975
  26. 26. Fletcher E. Mendelian heredity in cotton. The Journal of Agricultural Science. 1907;2:281-282
  27. 27. Li Z, Su Q , Xu M, You J, Khan AQ , Li J, et al. Phenylpropanoid metabolism and pigmentation show divergent patterns between brown color and green color cottons as revealed by metabolic and gene expression analyses. Journal of Cotton Research. 2020;3(1):1-1
  28. 28. Shi YZ, Du XM, Liu GQ , Qiang AD, Zhou ZL, Pan ZE, et al. Genetic analysis of naturally colored lint and fuzz of cotton. Cotton Science. 2002;14(4):242-248
  29. 29. Sun DL, Sun JL, Du XM. Genetic study on the color of fiber and linter in brown cotton. Journal of Anhui Agricultural Sciences. 2008;36:6254-6255
  30. 30. Yan Q , Wang Y, Li Q , Zhang Z, Ding H, Zhang Y, et al. Up-regulation of Gh TT 2-3A in cotton fibres during secondary wall thickening results in brown fibres with improved quality. Plant Biotechnology Journal. 2018;16(10):1735-1747
  31. 31. Liu Q , Luo L, Zheng L. Lignins: Biosynthesis and biological functions in plants. International Journal of Molecular Sciences. 2018;19(2):335
  32. 32. Li T, Fan H, Li Z, Wei J, Lin Y, Cai Y. The accumulation of pigment in fiber related to proanthocyanidins synthesis for brown cotton. Acta Physiologiae Plantarum. 2012;34(2):813-818
  33. 33. Lu N, Roldan M, Dixon RA. Characterization of two TT2-type MYB transcription factors regulating proanthocyanidin biosynthesis in tetraploid cotton, Gossypium hirsutum. Planta. 2017;246(2):323-335
  34. 34. Humphries JA, Walker AR, Timmis JN, Orford SJ. Two WD-repeat genes from cotton are functional homologues of the Arabidopsis thaliana TRANSPARENT TESTA GLABRA1 (TTG1) gene. Plant Molecular Biology. 2005;57(1):67-81
  35. 35. Aleksandra M, Ksenia S, Elena K. The genes determining synthesis of pigments in cotton. Biological Communications. 2019;64(2):133-145
  36. 36. Fan L, Shi WJ, Hu WR, Hao XY, Wang DM, Yuan H, et al. Molecular and biochemical evidence for phenylpropanoid synthesis and presence of wall-linked phenolics in cotton fibers. Journal of Integrative Plant Biology. 2009;51(7):626-637
  37. 37. Feng H, Li Y, Wang S, Zhang L, Liu Y, Xue F, et al. Molecular analysis of proanthocyanidins related to pigmentation in brown cotton fibre (Gossypium hirsutum L.). Journal of Experimental Botany. 2014;65(20):5759-5769
  38. 38. Li YJ, Zhang XY, Wang FX, Yang CL, Liu F, Xia GX, et al. A comparative proteomic analysis provides insights into pigment biosynthesis in brown color fiber. Journal of Proteomics. 2013;78:374-388
  39. 39. Xiao YH, Yan Q , Ding H, Luo M, Hou L, Zhang M, et al. Transcriptome and biochemical analyses revealed a detailed proanthocyanidin biosynthesis pathway in brown cotton fiber. PLoS One. 2014;9(1):e86344
  40. 40. Hua S, Yuan S, Shamsi IH, Zhao X, Zhang X, Liu Y, et al. A comparison of three isolines of cotton differing in fiber color for yield, quality, and photosynthesis. Crop Science. 2009;49(3):983-989
  41. 41. Campbell BT, Dever JK, Hugie KL, Kelly CM. Cotton fiber improvement through breeding and biotechnology. In: Cotton Fiber: Physics, Chemistry and Biology. Springer, USA; 2018. pp. 193-215
  42. 42. Schmutz A, Jenny T, Amrhein N, Ryser U. Caffeic acid and glycerol are constituents of the suberin layers in green cotton fibres. Planta. 1993;189(3):453-460
  43. 43. Hua S, Wang X, Yuan S, Shao M, Zhao X, Zhu S, et al. Characterization of pigmentation and cellulose synthesis in colored cotton fibers. Crop Science. 2007;47(4):1540-1546
  44. 44. Ryser U, Meier H, Holloway PJ. Identification and localization of suberin in the cell walls of green cotton fibres (Gossypium hirsutum L., var. green lint). Protoplasma. 1983;117(3):196-205
  45. 45. Basavaradder AB, Maralappanavar MS. Evaluation of eco-friendly naturally coloured Gossypium hirsutum L. cotton genotypes. International Journal of Plant Sciences. 2014;9(2):414-419
  46. 46. Frydrych I. Cotton quality evaluation: New possibilities. In: 62nd ICAC Plenary Meeting, Gdańsk. ICAC. Poland. 2003
  47. 47. Nath J, Patil PG, Shukla SK, Arude VG. Design of pocket colorimeter for cotton. Agricultural Engineering Today. 2002;26(3 and 4):79-88
  48. 48. Naeem-Ullah U, Ramzan M, Bokhari SH, Saleem A, Qayyum MA, Iqbal N, et al. Insect pests of cotton crop and management under climate change scenarios. In: Environment, Climate, Plant and Vegetation Growth. Springer, Nature Switzerland; 2020. pp. 367-396
  49. 49. Vreeland J. The revival of colored cotton. Scientific American. 1999;280(4):112-118
  50. 50. Feng H, Guo L, Wang G, Sun J, Pan Z, He S, et al. The negative correlation between fiber color and quality traits revealed by QTL analysis. PLoS One. 2015;10(6):e0129490
  51. 51. Blas-Sevillano RH, Veramendi T, La Torre B, Velezmoro-Sánchez CE, Oliva AI, Mena-Martínez ME, et al. Physicochemical characterization of several types of naturally colored cotton fibers from Peru. Carbohydrate Polymers. 2018;197:246-252
  52. 52. Ruhsora R, Muhabbat R. The nature of inheritance of the colored cotton fiber trait. Nature. 2021;7(5):229-232
  53. 53. Kohel RJ. Genetic analysis of fiber color variants in cotton 1. Crop Science. 1985;25(5):793-797
  54. 54. Khan NU, Hassan G, Kumbhar MB, Marwat KB, Khan MA, Parveen A, et al. Combining ability analysis to identify suitable parents for heterosis in seed cotton yield, its components and lint% in upland cotton. Industrial Crops and Products. 2009;29(1):108-115

Written By

Sathees Nagarajan, Yazhni Purushothaman, Monika Selvavinayagam, Pandidurai Govindharaj and Aasif Musthafa

Submitted: 04 April 2022 Reviewed: 12 April 2022 Published: 05 October 2022