Indian Journal of Fibre & Texti le Research Vo l. 28, Septembe r 2003, pp. 348-362
Review Article
Structure and structure-property relationship in native cotton Some challenges to breeders A V Moharir" Di vis io n o f Agricultural Ph ys ics. Indi a n Agri c ultural Resea rc h Institute, Ne w Delhi 110012, Indi a
Received 27 May 2002 ; accepled 27 August 2002 Thi s paper presents a brief review o f the literature on the current statu s of the supra molecular s tructure o f native cotto n fibre and of th e ex pe rime ntal data on th e phys ica l properties and cell ulose crystallite X-ray orientation parameters in respec t o f a large number of the same cotton vari e ti es. belong in g to diffe rent species of cotton g rown at widely differe nt ag roc lim atic local io ns, name ly Coimbatore. Nagpur. New De lhi and Si rsa, betwee n 11.0 0 and 29.0 0 North latitudes in India. Al so prese nted a nd di sc ussed are the data on cellulose synthesis in sixtee n cotton va rie ti es g rown under ide nti cal ag roc lima tic co nditi o ns at one loc ati o n. Possible reasons for the variations in the physical and technolog ical prope rti es o f fibres have been obse rved and the implicati o ns of these as c hall e nges to cotton breeders in India are foresee n in the li ght o f the de mand of the mode rn cotto n processin g tec hno logies. Keywords: Cotto n, Struc ture-prope rty re lation ship, Cellulose sy nthes is
1 Introduction The association of cotton fibre and textiles with man dates back to the Indus Valley civilization that flourished in India some 3000 years B.C. when cotton was not only cultivated as a commercial crop but was also being spun into yarns and woven into textiles 1.3 . Ever since, cotton has held a pl ace of pride amongst the cash crops all over the world and more particularly in the Indi an sub-continent and in othe r cotton growing countries. The tradition all y unmatched beauty and comfort of cotton and cotton fabrics has mainly been responsible for the spread of its cultivation in areas beyond the boundaries of its original natural habitat 1.3. The history of cotton cultivation, development and its spread to variou s locations all over the world has almost become synonymous with the progress of mankind. No surprise, the James Watt ' s steam engine, James Hargreave's spinning je nny, Richard Arkwright' s water frame spinning machine (mule), Samuel Cartwright's power operated loom, John Kay's flying shuttle, Reverend William Lee's knitting frame and EI Whitney 's cotton gin have become the symbols of new industrial revolution that ushered In an unprecedented prosperity to several nations, and reasonable purchasing power at the hands of poor "Pho ne: 25 841178 125848853: Fax: 0091-0 I 1-25842321: E-mail : amo harir @iari.res.in
workers across the world . 3. This commercial sought after aspect in the spread of cotton cultivation has al so been mainly responsible for vigorous genetic, agronomic, scientific and technological researches for improved fibre quality besides economic yields for cotton growers l.). No other agricultural commodity has invited and attracted the attention of th e most talented people from all di sciplines o f science, technology and humanities and integrated thei r ideas, as the cotton cropl .3. l
A cotton fibre is a tubular outgrowth of a single tiny cell on the outer epidermi s of seed / fertili zed ovule. This fibre progresses through four developmental stages: initiation, elongation , secondary wall synthesis and deposition, and dehydration 1.5. Whereas the first three stages occur inside the cotton boll while the fibre is alive and actively growing, the last stage occurs when the developin g cotton bolls are cut off from the tran spiration stream of the cotton plant and the fibres are physiologically dead. Excellent reviews on the developmental biology of cotton plant and fib re have been published 6. 13. From the present knowledge of the structure of cotton fibre, it constitutes about 95-99% pure cellulose, which is wholly crystalline in nature6.9. 14.17. This cellulose is synthesized within the cells and long unending threads called deposi ted as
MOHARIR: STRUCTURE AND STRUCTURE-PROPERTY RELATIONSHIP IN NATIVE COTTON
t
microfibrils ; the maximum amount is deposited into the diurnal secondary layers of individual fibres during boll maturity period 6. 13-19. Cotton is primarily cultivated all over the world for its fibres as industrial raw material 1-3. 6. The mature cotton fibre used in textile processing is the end product of this boll maturation period and upon which depend almost all the physical and chemical properties of cotton and its technological processing into textiles. Cellulose biosynthesis and its deposition into secondary layers of developing cotton fibres is a very complex process and excellent reviews have been published on this aspect6.10-17.19-25. No wonder, increments in fibre length, strength, fineness and maturity have always been sought as improvements in fibre quality to accommodate newer developments in spinning and processing techniques . In view of its purity, cotton cellulose has been the subject of extensive investigation, employing diverse physical and chemical techniques for biosynthetic pathways of cellulose synthesis, extent of cellulose polymer chain length, degree of polymerization, optimum environmental conditions for synthesis, rate and amount of cellulose synthesis, density, crystallite size, crystal lattice type, unit cell size and its group, structure and disposition of cellulose, polymorphs, orientation of cellulose crystallites within microfibrillar polymer and within the matrix of diurnal secondary growth layer of developing cotton ' . 6 10-17. 19-71 -. fI'b res, an d extent 0 f crysta II Inlly' Cellulose synthesis and its deposition is known to be influenced by several factors such as so il condition, location of the growth, humidity and soil moisture, maximum and minimum diurnal temperatures, and variety and species of cotton 1.2.4-9.2836. The unit cell describes the dimensions of the cellulose crystallite and the one that is most well known is that of Meyer and Misch 6.26 and its dimensions are: a = 0.835 nm, b = 1.03 nm, C = 0.79 nm and monoclinic angle ~ = 84°; this crystal is generally but not unanimously accepted to be monoclinic and of space group P2. The degree of polymerization of native cellulose is estimated to be 3500-12000 and in purified cotton linters, it is 10003000. Cellulose from various sources, particularly from the cotton linters, has been widely researched and yet the basic processes of its biosynthesis and disposition in cotton fibre matrix are still not known for certain. All these parameters have been used and found to have practical applications in determining structure-property correlations and assessment of
349
technological performance of cotton fibres with considerable success 1.2.9.26.37, and yet several aspects of the structure of native cotton fibre are still wide open to debate. Some such areas are: ( i ) rate and amount of cellulose synthesis, and (ii ) orientation and disposition of crystalline cellulose to the fibre axis within the individual diurnal secondary layers of developing cotton fibres, complicated by the . l" generatIOn 0f reversa extinctIOn b an d s 38-4?- an d convolution twists2.6.9.26.42.43, discussion over reversals and convolutions (being genetic or environmental in origin) and variation of their frequency within and between varieties of different species, their influence over physical and technological performance of fibres6.9.26.38.40.43 and debate over the constancy / variation of the true spiral angles in orientation of crystalline cellulose between varieties and species of cotton6-26. 44-46 in never-dried state of fibres . Genetic improvement in the physical properties of cotton fibre have been remarkable in several parts of the world and yet has lagged behind the corresponding improvements in the machines that spin threads from the fibres . Genetic improvement is hampered by the strong association of poor quality and high yields and the lack of knowledge about the genes that contribute or control the individual traits of fibre quality. Another serious di fficulty with cotton research being that the same set of developing fibres cannot be progressively observed every day during the entire developmental period. This is partic ularly serious in view of a very large variability in fibre properties observed from one single plant, single boll and from over single seed of cotton 1-9. In this paper, an attempt has been made to look at the various structural aspects of cotton fibres in the light of some recent developments and experimental observations, and explore the possible challenges to cotton breeders in the near future . And while doing so, some of the well known aspects of the development and growth of cotton plant in general a nd of cotton fibres inside the cotton boll have been ignored and the description has been confined only to the principal aspect of the organization and disposition of cellulose in cotton fibres and structureproperty linkages with it in the light of new experimental data.
2 Reversal Extinction Bands Cotton fibres, examined under crossed polarized light in a polarizing optical microscope, are known to show transverse, dark bands along the length of
INDI AN J. FIBR E TEXT. RES., SEPTEMB ER 2003
350 47
48
fibres. Balls and Ball s and Hancock proposed for the first time that the longitudinal spirals of micromacro fibrillar cellulose wi thin the secondary diurnal growth layers of developing cotton reverse their sense of orientation several times along the length of the fibres and in the process come into configuration where the crystallites lay parallel to the fibre ax is in a very narrow region and thereby exhibit an extinction band under crossed polarized light. Ball s called them as "Reversa ls" and ever since then, everyone has call ed them so. Debate over the origin and growth of reversals along the length of cotton fibres, their being geneti c or environmental in origin, and their contribution to strength or weakness of fibres is very extensive and lon g drawn over interva l of time 38 -4o. In 41 a recent communication , a significant increase in reversa l frequency has been shown with the exogenou s application of g ibberellic acid hormone during the growth of cotton plants. However, it is not clear whether the gibbere llic ac id contributes to increased crystalli zation of cellulose within the developing seco ndary walls of cotton fibres or it promotes the deposition of more number of secondary layers, enhancing thereby the boll maturation period. Further, increased reversal frequency means inc reased ori entation of cellulose to fibre ax is and crystalli zation and these changes must consequently be reflected in the increased tenacity of fibres. 40 Moh arir et al.38 - explained the origin and growth of ex tincti on / reversal bands without resortin g to the Ball s' co ncept of abrupt reversals on the bas is of relat ive phase shifts between success ive uni-modal spiral di sposition of crystalline cellulose microfibril s 111 consecutive, diurn al and coaxial-cylindrical secondary layers of developing cotton fibres_ However, the recent data40 on the frequency of reversa l ex tinction bands in field- matured cotton fibres of a large number of the same varieti es, grown at four different agro-c1 imatic locations in Indi a (Tab le 1), indicate that the frequency of reversal extinction bands per centimeter length of fibre remains practically invari ant within individual variety with location of growth, although its values vary between individual varieties. Apparently , the reversals appear to be genetically determined and th ey have been demonstrated to be defi nitely formed durin g th e seco ndary thickenin g phase of cellulose deposition in developing cotton fibres 38 .39 . Minor variatio n in the reversal frequency can be attributed to several compl ex local env ironmen tal conditions of growth and the mann er in which the crystalline ce llulose
aggregates are actually formed, since thi s process is known to be very sensitive to diurnal ni ght temperatures 28 -37 .
3 Convolution Twists, Convolution Number and Convolution Angle Field-matured naturally deh ydrated cottons, irrespective of species, are know n to generate characteristic twi sts along the length of fibres, known as convolutions. Convolutions have been believed to provide inter-fibre grip in yarns. Convolution frequency and convolution ang le have been studied ex tensively and di scussed in de tail keepi ng in view the variation between varieties a nd species, being genetic or environmental in origin, their influe nce in mod ify ing the sp iral-geo metry of cellulose di spos ition in cotton fibres and consequently the strength of fibres and yarns 2 .6 .42.43.49-54. Rece nt data on average convolutions and convolution angles in respect of a large number of the same diploid and tetraploid cotton varieti es grown at di fferent agro-climatic location s in Indi a during 1992, 1994 and 1995 crop years (Table 2) suggest that both convolutions and convolution ang les are genetically determ ined . Their values between differe nt varieties of cotton differ but within a variety the valu es remain practically invariant with th e location of growth of cotton_ Further, the spiral structure laid down during the secondary wall thickening period a nd not the temperature during dehydration pl ays a determini stic role in the formation of convolutions. The convolutions also appear to depend upon th e ori e ntati on and the cell wall thickness and conseq ue ntl y on the rate of cell ulose biosy nthes is and cellulose depositi on within the walls of develop ing cotton fibre, as observed from the very si g nificant correlati ons w ith sp iral a ng les (Tabl e 3); lower sp iral ang les correspond to lower convo lution angles, hi gher amount of ce llulose in fibres and consequently increased orientation of cellulose to fibre axis_ Thus, the convolutions and reversa l extitnction bands remain practically invariant withi n individual varieties with the change in locati on of growth , and minor variations in the ir frequency may be seen to ari se as a result of the complex genotype- metabo licenvironmental interacti o n at the pl ace of growth of cotton in view of the fact that there are no basic differences between the mass density of cellul ose in never-dri ed f ibres 55 , ori entation of cellul ose cry stall ites to the fib re ax is 56 a nd the size of the crystallographic units 56 .57 of cellu lose within the
MOHARIR: STRUCTURE AND STRUCTURE-PROPERTY RELATI ONS HIP IN NATIV E COTTON
35 1
Table I- Location-wise data on number o f ex tinctio n bands per unit length of cotton fibres in dip lo id and tetrap loid varieti es Cotton vari ety
Year of growth
Locati on o f growth
Ex tin cti on ba nds / cm
Av . ex tincti on band di stance ( cm )
Tetraploid BN
1992
Nagpur
22.80
0.043
BN
1992
Coimbatore
20.36
0.049
BN
1994
Sirsa
22.97
0.043
BN
1994
New Delhi
19.65
0.050
New Delhi
18.45 (L)
0.054 (H)
BN
1995
BN
1995
Nagpu r
2 1.83
0.045
BN
1995
Coimbatore
23.5 1 (H)
0.042 (L)
Range within variety (H-L)
5.06
0.0 12
Average va lue
21.36
0.046
Standard dev iati on
1.9 1
0.004
Suvin LRA-5166
1995 1994
New Delhi
22.70
0 .044
Sirsa
20.78
0 .048
LRA-51 66
1994
New De lhi
22.27
0.045
LRA-5166
1994
Nagpur
21.35
0.046
LRA-5 166
1994
Coilll batorc
22. 19
0.045
LRA-5 166
1995
New Delhi
2 1. 84
0.045
LRA-5166
1995
Nagpur
18.89 (L)
0.053 (H)
LRA-5 166
1995
Coimbatore
22.74 (H)
0.044 (L)
Range within vari ety (H-L)
8.85
0.009
Average va lue
2 1.43
0 .046
Standard devi ati on
1.29
0.003
AC-738
1994
Sirsa
20.40
0 .049
AC-738
1994
New Delh i
19.26 (L)
0.052 (H)
AC-738
1994
Nagp ur
20.88 (H)
0.047
AC-73 8
1995
Coilll batorc
19.35
0.05 1
Range within varie ty (H-L)
1.62
0.005
Average va lue
19.97
0.050
Standard devi ati on
0 .79
0.002
Diploid Y- I
1994
Sirsa
16.33
0.06 1
Y-I
1994
New De lhi
18.21 (H)
0.054(L)
Y- I
1995
Coimbatore
15.85(L)
0.063 (H)
Range wi thin vari ety (H-L)
2 .36
0.009
Average va lue
16.79
0 .059
Standard deviation
3. 12
0 .008
Sirsa
15.21 (L)
0 .065 (H) 0.059 (L)
AKA-5
1994
AKA-5
1995
New Delhi
16.79( H)
AKA -5
1995
Coimbatorc
16.39
0.06 1
1.58
0.006
Range withi n variety (H-L)
Contd
INDIAN J. rIBRE TEXT. RES. , SEPTEMBER 2003
352
Table I-Location-wise data on number of extinction bands per unit length of cotton fibres in diploid and tetraploid varieties -Conld Cotton variety
Year of growth
Location of growth
Extinction bands / cm
Av. extinction ba nd distance ( cm )
Average value
16.13
0.062
Standard deviation
0.82
0.003 0.039 (L)
SRT-IGCot.IO
1992
Nagpur
25 .38 (H)
SRT-IG Cot.JO
1992
Coimbatore
23.70
0.042
SRT-IG Cot.lO
1994
New Delhi
20.13 (L)
0.049 (H)
Range within variety (H-L)
5.25
0.010
A verage value
23.07
0.043
Standard deviation
2.68
0.005
AKH-4
1995
New Delhi
21.80 (H)
0.045 (L)
AKH-4
1995
Nagpur
19.96
0.050 (H )
Range within variety (H-L)
1.84
0.005
A verage va lue
20.88
0.047
Standard deviation
1.30
0.003
Maljari
1992
Nagpur
14.19
0.070
Maljari
1992
Coimbatore
13 . 18 (L)
0.075 (H)
Maljari
1994
Sirsa
17.68 (H)
0.056 (L)
Maljari
1995
New De lhi
16.8 1
0.059
Ran ge within variety (H-L)
4.50
0.019
Average value
15.46
0.064
Standard deviation
2. 12
0.009
H- Highest value; and L-
Lowest value
different varieties and species of cotton, and particularly in view of the strong argument that the reasons for apparent differences in cotton varieties must be soug ht in some higher order of structural organization 27 .55 -58. The knowledge of relative variations in degree of polymerization, crystallite size, crystallinity and orientation of crystallites to the fibre axis is helpful in understanding the inter- cotton differences, fibre .. 2'" 6 26 44-46 "59 60 . . an d c h emlca . I reactivity properties
4 Crystallinity, Crystallite Size and Crystallite Orientation The detection of crystalline cellulose in the early stages of developing cotton fibre by X-rays have led to the extensive researches on the fine structure of cotton cellulose and its disposi tion in fibre. The hi story of the use of X-rays in cotton researches is very extensive and has been rev iewed by Moharir and others2.6.26.61 -66. The data on crystallite sizes in raw cotton fibres of seven
cultivars belonging to d iploid Gossypium arboreum al1id tetraploid Gossypium hirsutum species are given in Table 4. Cellu lose crystallite sizes in directions perpendicul ar to (WI), (lOT) and (002) planes were estimated from X-ray powder diffraction patterns. The diffraction peaks were resolved using FITX-RA Y diffraction data analysis program (written by SOCABIM, Siemens DlFFRAC AT Software System, Germany). The comp lete data for all the three equatorial planes were analyzed for 2d values, full width at half maximum (FWHM), and the normalized area under the three eq uatori al diffraction peaks. The mean crystallite sizes were determined using the Scherrer's equation. This comprehensive study on the same cotton eultivars, grown at different locations and in different crop years, show variations in crystallite sizes corresponding to crystallographi c planes (101), (lOT) and (002) within individual varieties of both species, but the combined average crystalli te size corresponding to (101) and (lOT) planes taken together is
MO HA RIR: STR UCT URE AND STRUCT URE- PROP ERTY RELATI O NSHIP IN NAT IVE COTTON
Tab le 2- Loca ti o n-wi se average va lu es of co nvo luti o ns and co nvo luti o n angles fo r all years of crop growth COllo n variety
Averagc valuc
Locati o n o f grow th
Convo luti o ns jJcr cm
Co nvo lutio n angle (8)
dcg
A KH -4
Si rsa
43 .86
7 .6 1
A KH -4
New De lh i
43.36
6.87
A KH -4
Nag pur
43.83
704
A KH -4
Coimbatore
4 1. 28
7.53
1.39
0.36
Standa rd deviatio n AC-738
S irsa
62.03
11.62
AC-738
New De lhi
52 .57
8.76
AC-738
Nag pu r
5 1.83
8.8
Coi mbaLOre
58 .77
9.6
4.92
1.34
AC -73 8 Stand ard deviati o n BN
Sirsa
6 1.47
10.88
BN
New Delhi
52 .62
8. 84
BN
Nagpur
49. 8 1
8 09
BN
Coi mbato re
5 1.99
8.8
5. 14
1.2
StaIl dard devi ati o n Y- I
S irsa
40 .61
7. 13
Y- I
New De lhi
40.56
6.7
Y- I
Nag pur
47.88
7.2
Y-I
Coilll bato re
44.52
7.3
3.52
0.26
Standard dev iati o n Malj ari
Sirsa
48.67
8.45
Malj ari
New De lh i
41.00
6.00
Malj ari
Nag pur
43.85
7.63
Malj ari
Coilllbatore
43.29
7.26
3.22
1.02 7.46
Standard deviati o n A KA-5
Sirsa
43.67
AKA-5
New Delhi
42. 20
7.4 1
AKA -5
Nagpur
38.43
6. 11
AKA-5
Coilllbatore
48.85
8.36
4 .31
0.92 10.49
Standard deviation LH-900
Sirsa
62.55
LH-900
New Delhi
47.11
7.20
LH-900
Nag pur
56.84
10.62
LH-900
CoimbaLOre
52.61
8.23
6.53
1.69
Standard deviati o n
Conrd
353
INDIAN J . FIBR E TEXT. RES .. SEPTEMBER 2003
354
Tab le 2-Location-wise average values or convolutions and co nvoluti o n ang les ror al l yea rs or crop growth--Contd Co tton va ri ety
Location or growth
Average va lu e Co nvolut io ns per CIll
Convo luti o n ang le (8) c1eg
LRA-5 166
S irsa
64 .04
9.98
L RA -5166
New Delhi
56.56
9.03
LRA-5 166
Nag pur
46.39
7.20
LRA-5 166
Coilllbatore
55 .78
9.97
7.23
1.3 1
5 1.11
~).64
Standard de viation SRT- I G.Cot- IO
Sirsa
SRT- I G .Cot-IO
New Delhi
S RT-I G .Co t- IO
Nagpur
49. 10
7.37
SRT- I G.Co l.- IO
Coilllbatore
52.50
8.36
1.71
0.66
Standard dev iati on Suvin
Si rsa
Suv in
New Delh i
48.96
7.50
Suvin
Nagpur
44.26
6.40
Suvin
Coilllba tore
52.60
8. 35
4. 18
0.98
46.93
7.50
Sta ndard deviati o n Jyoti
S irsa
Jyoti
New Delhi
Jyo ti
Nagpur
Jyoti
Coilllbatore
Standard deviation
45.65
7.2 1
0 .90
0. 20
G.Cot- 13
Dhanclhuka
43 .6 1
6.77
G.Cot - 13
Chharocli
44.99
6.90
0.97
0.09
6 1.36
11 .40
44.44
6.40
11.96
3.53
Standa rd devia ti on G .Cot- IOO
S irsa
G.Cot-IOO
New Delhi
G.Cot-IOO
Nagpur
G.Cot- IOO
Coilllbatore
Standard deviation
always equal to the average crystallite size of (002) plane. It is visualized th at the variations in crystallite s izes arise as a result of the differences in the amount of cellulose sy nthesized within fibres of varieties and their deposition into crystalline matrix within developing cotton fibres, since cellulose molecules from various sources at the molecular level are same but they differ in the c rysta lline structures and binding by other biochemicals67 . Th. Malutan et.a!' (http://omicron.ch.tuiasi.ro/- thmalu Isymp99_I .pdf )
estimated the indi vidua l contribution of (IOI and lOT) crystallog raphic planes to about 24.36 % towards crystallinity as co mpared to only 5.28 % by the (002 and 02 1) planes. A cotton breeder, therefore, has been suggested to look for cottons where both the length-to68 width ratio of the pol ymer .69 and the orientation of cellulose crystallites to the fibre axis 26 are hi gh ancl to use these parameters in screening parent genotypes for breeding and evolving newer strains with increased strength of fibres.
MOHARIR: STRUCTURE AND STRUCTURE-PROPERTY RELATIONS HIP IN NATIVE COTTON
Table 3- Correlations of convoluti on angle with other orientat ion parametcrs Convolution angle (8), deg Convolutions I cm
p > 0.c)01
=- 0.132,
NS
Av. angle of orientation (a 111)
r = 0. 198,
NS
40 % X-ray angle
r= 0.222 ,
NS
50 % X-ray angle
=0.206, r =0.3 12,
NS p > 0.0 1
(40 % - 8)
r = - 0.436,
p > 0.001
(50 % - 8 )
r
=- 0.481, =- 0.355, r =- 0. 126,
p > 0.001
(a ",- 8)
r
p > 0.01
Hermans factor
r
r
75 % X-ray angle
...
r = 0.875,
True - spiral angle"
Relati ve orientation index W.r.t. ramie
NS
NS: Not signi fican t. "For definiti on, see reference Moharir el a/.. J App/ Po/ym Sci, 44 ( 1992) 19 13.
The o ri e ntatio n of cellulose crysta llites with respect to th e fibre ax is is known to dete rmin e the inter-cotton differences and most of the techn o logically important properties of the fibre 2.6.26 ,44-46.54.59-7o. Both the optical and X-ray me thods are used to determine orie ntation2.6.26.62,63, Since the optical methods are generally very tedi o us and require a large amount of data for a representative value of the fibre, X-ray methods are genera ll y favoured in o ri entation studi es. The most widely accepted parameters for characteri zing the o ri entation of the crystallites with respect to th e fibre axis are the Herman s crysta llite orie ntat io n factor and 40% and 50% X-ray orientation ang les fro m the most intense eq uatoria l (002) X-ray diffraction peak intensity profile, and these have been show n to corre late w ith importa nt fibre pro perti es, particularly the strength2.6.26.61.64 . Moharir, el al.60 , from their data on Hermans crystallite orie ntation factor, the average a ng le of o ri entatio n (a Ill) and 40%, 50% and 75 % X-ray angles in respect of the same 13 cotton varieties grown at different agro-c limatic locations and in di ffe re nt crop years , have observed that whereas the average val ues of the X-ray o rientation para meters are different for different vari eties, these remain practi ca ll y invariant w ith in individual variety with the cha nge in the location o f g rowth o f cotton and the X-ray orientation parameters appear to be geneti c in thei r ori g in and independe nt of agro-c1imati c co nditi o ns and locati o n of growth , The reaso ns for minor vari ati o ns in the valu es of these
355
orientation para meters for the same varl ettes with location of growth can possibl y be soug ht in the rate and amount of cellulose sy nthes ized , which considerab ly varies in individual variety with locatio n of 'growth of cotton and conseq ue ntl y with the environmental condition s of growth , as ev ide nced by the variation in the maturity of cotton with latitude of the place of growth2,6, 19,20,26.67,7 1.83. The above conclusions are in contradiction to that of Hebert et al. 55 who concluded from e lectron diffracti o n studi es that the degree of o ri e ntati o n of crysta llites within and between cotton varieties did not de vi ate apprec iab ly from one another. The conviction and recommendati o n26,54.70 in using the Hermans crystallite orientation factor as the parameter for sc reening cotto n genoty pes for increased stre ngth of fibres and its use in cotto n breeding programs for evolving newer strain s with improved stre ngth of fibres through hybridizati o n is the reby streng the ned . It may a lso be pertinent to mention here that the genetic inheritance of the Hermans o ri e ntation factor and X- ray orientation 84 ang les had been seen earli er in the inte r-spec ifi c FI hybrids of Gossypium barbadense and Gossypium hirsutum pare nts, and particul arl y in view of the fact that the streng th of cotton fibres has not o nl y been found to be genetica lly inherited but e nvironmenta ll y the most stabl e of a ll cotton fibre properties 85 .
5 Rate of Cellulose Synthesis in Cotton with Change in Location of Growth Table 5 shows the data on average rates of cellulose sy nthes is at 10, 20, 30 and 40 days pos tanthesis boll maturity stages in fo ur of the twe lve c ultivars of cotton, grown at two different agroclimatic locations in 1992,1994 and 1995 crop years, in re lati o n to o ther physical and crysta ilite o ri e ntation parameters 73 ,86. Cellulose_content of cotton bolls was estimated fo llowi ng the method described by Updegraff7 and the average values of three repli cates for each maturity stages of bo lls are reported. It may be obse rved that whereas the rates of cellul ose synthesis fo r the tetraplo id vari eti es are hi gher between \0 days and 20 days post-a nthesis peri od at Nagpur, the same are co nsiderably hi g her between 30 day s and 40 days post-a nthes is peri od at Coimbatore . The ove rall average va lu es of cell uiose sy nthes is are lowe r fo r Nagpur as co mpared to those for C oimbatore. Further, with not very si g nifi cant variati o ns in the va lues of Hermans crystallite orientation factor fo r a ll the indi vid ual cotto n vari eties
INDIAN J. FIBRE TEXT. RES .. SEPTEMBER 2003
356
Tab le 4-Cry ·tallite sizes fo r the COllon cultivars grown at different locations and in different crop yea rs COllon variety and specics
Year of
locati on of
growth
growth
Crystallite size (A) 101
10 I
Av. of 101+10 T
002
Av . of 10 1+1 0 i +002
Gossypium arboreul1I Y-I
1994
Sirsa
27.1
Y-I
1994
New Delhi
25.8 (l )
Y-I
1994
Nagpur
34.7
Y-I
1995
Nagpur
44.2
73. 1
50.1
49.9
80.4
53. 1
54. 6 (H)
53.!i
86.0 (1-1 )
60.3 (H)
5 1.0
57.'2 (H)
43.4 (L)
43.8 (l )
49.0 (l )
45.6 (l )
50.0
Average within the va ri ety
32.9
70.7
51.8
51.2
51 .6
Ran ge of va ri at io n (II - l)
18.4
42 .6
8.5
3.6
11.6
Maljari
1992
Nagpur
27.6 (l)
49.9
38.7
50.8
42.S
Maljari
1992
Coilllbatore
39.0
66.2
52.6 (H)
53.8 (1-1 )
53.0 (H)
Malja ri
1994
Sirsa
35.5
70.7
53. 1
5 1.5
52.6
Maljari
!994
Nagp ur
33.6
69. 1
51.3
52.2
51.6
Malja ri
1995
New Delhi
27.4
47.7 (l)
37.5 (l)
50.5
41. 8 (l )
Maljari
1995
Nag pur
32.6
72. 1 (1-1)
52.3
5 1.6
52.1
Maljari
1995
Coilllbatore
43.9 (H)
52.0
47.9
49.4 (l)
48.4
Average within the va riety
34.2
6 1.1
47.6
5 1.4
48 .9
Range of vari ation (1-1 - L)
16.3
24.4
15 . 1
4.4
11. 2
AKH-4
1992
Nagpur
27. 1
72.9
50.0
54.2
51.4
AKH-4
1992
Coilll batore
23.0 (L)
140.7 (1-1)
S I. 8 (H)
56.1
73.3 (H)
AKH-4
1994
Sirsa
36.S
75.7
56.2
56.2 (H)
56.2 (l )
AKH -4
1994
Nagpu r
36.2
6 1.2
48.7
50. 1
49.2
AKH-4
1995
New Delhi
26.4
58.3
42.3 (l)
5 1. 5
45.4
AKH-4
1995
Nagpur
45.6 (1-1)
44.9 (L)
45.2
49.0 (L)
46.5
Average within the va riet y
32.5
75 .6
54.0
52.9
53 .7
Range o f variation (H - l)
22.6
95.8
39.5
7.2
17. 1
1992
Nagpur
26.7 (l)
75 .0
50.8
53.5 (H)
51.7
AKA-5
1992
Coilllbatore
27.4
112 .9 (1-1)
70.1 (1-1 )
5 1. 5
63.9 (H)
AKA-5
1994
Sirsa
33 .7
69 .9
5 1.6
53.4
52.2
33.8
85.6
59.7
5 1.0
56.S
AKA-5
AKA-5
1994
Nagpur
AKA-5
1995
New Delhi
29 .7
43 .3 (l)
36.5 (L)
49.2 (l)
40. 7 (l )
AK A-5
1995
Nagpu r
36.4 (1-1)
63 .0
49.7
50.7
49 .8
Average within the variety
31.3
74.9
50.3
51.5
52. 5
Range o f variati on (1-1 - l)
9.7
69.6
34.6
4.3
23. 2
Gossypiulll hirsutu11I
BN
1992
Nagpur
27.7
89.2
58.4
53.2
56.7
63.2
50.6
52.5
51.9
BN
1992
Coi Illbatore
40.0
BN
1994
Sirsa
24.2 (L)
93.1
58 .6
56.9 (1-1)
58. 1
BN
1994
New Delhi
25.6
103.6 (1-1 )
64.6 (1-1)
52.0
60.4 (1-1 )
BN
1994
Nagpur
34. 1
65 .3
49.7
51.8
50.4
BN
1995
New Delhi
48.1 (1-1)
62.4
55.2
55.7
55.4 COl1ld
357
MOHARIR: STRUCTURE AND STRUCTURE-PROPERTY RELAT IONS HIP IN NATIVE COTTON
Ta ble 4-Crystallit e sizes fo r the collon culti va rs grown at different loca ti ons and in different crop years--Collid Colton vari ety and spec ies
Year o f
l ocation of
grow th
grow th
Crystallite size (A) 101
10 I
Av. of 101+ 10 I
002
Av. of 101+1 0 I +002
39.4
54. 1 (l)
46.7 (l )
48.7 (l )
47.4 (l)
Average wit hin the va ri ety
34.2
75.8
54.8
53.0
54.3
Ra nge of variat ion (H - l )
13.9
49.5
18.9
12.2
13.0
BN
1995
Nag pur
lH -900
1992
Nagpur
43.8
62.5
53. 1
54 .6 (1-1 )
53 .6
lH -900
1992
Coimbatore
28 . 1 (l )
83 . 1
55 .6
51.8 (l )
54. 3
ll-l -900
1994
Sirsa
28 .9
94.9 (1-1 )
6 1. 9(1-1)
54.4
59.4 (1-1)
ll-l-900
1994
Nagpur
29.5
72.7
5 1.1 (l )
53.6
51.9 (l l
l l-l -900
1995
New Delhi
44.5 (1-1 )
60.8 (l )
52.6
52.4
52.6
Ave rage within th e vari ety
35.3
74.8
54. 8
53.3
54.3
Range of variati on (1-1 - l )
i 6.4
34 .1
10.8
2.8
7.5
lR A-5 166
1992
Nagpur
41. 8
54.0 (l)
47 .9
49.7
48 .8
lR A-5 166
1992
Co imbato re
44.9 ( H)
57.4
5 1.1
52. 1
51. 6
lR A-5 166
1994
Sirsa
27 .8
67.0
47.4
54.4 (1-1 )
50.9
lR A-5166
1994
New Delhi
28.3
58 .7
43.5 (l )
44.6 (l )
44.0 (l)
lR A-5166
1994
Nag pur
33.3
80 .7
57.0
52 .8
54 .9
lR A-5 166
1994
Co imbatore
25 .6 (l)
105.9( 1-1 )
65.7 (1-1)
53.3
6 1.6 (1-1 )
lR A-5 166
1995
New Delhi
37.6
58. 1
47.8
48.3
48.0
lR A-5 166
1995
Nagpur
43.4
58.0
50.7
47.5
49.1
Ave rage wi thin the va ri ety
34.0
67.4
51.3
50.3
51.1
Ran ge of variation (1-1 - l )
19.3
51.9
22 .2
9.8
17.6
33 .5
71.6
52 .0
5 1.9
52. 3
CO!llbi ned ave rage of all varieties trrken together 1-1-
hi ghest va lue within the variety ; l -
lowest value within th e variety; and (1-1 - l ) -Ran ge of va ri ation within the ind ividual variety .
between the two locations. the increased tenacity of cotton fibres for Nagpur can be attributed to the higher initi al sy nthesis of cellulose between 10 days and 20 days post-anthes is peri od and perhaps to the manner in which this cellulose is deposited 73 .86 . For the diploid cotton variety Maljari, the rates of cellulose synthesis are consistently higher for Nagpur at a ll the fo ur post-anthesis stages of boll maturity . Extensive data 86 on twelve cotton genotypes, however, indicate significa nt reduction in cellulose sy nthesis at Nagpur as compared to that at Coimbatore. Thi s observation suggests that whereas Coimbatore, located at lower latitude, brings out the inherent capac ity differences between the genotypes, the higher latitude location Nagpur induces a ll the same genotypes to sy nthesize more amount of cellulose. On ly the variety LRA-5166 shows almost equiva lent value for cellulose sy nthesis at both the locations, indicating perhaps greater climatic
' 7386 . a dapta bt'l 'Ity for t h'IS cu Itlvar ' . Th ese 0 bservatlOns are in agreement with those of Lewis a nd Bened ict 88 , who showed that cellulose deposited between 15days and 20 day s post-anthesi s period determine 50-80% of the bundle tenacity of fibres. Secondary cell wall deposition in developing cotton fibres is reported to be much greater at ni ght than in the day. Since sucrose is the major precursor for cellulose biosynthesis in deve loping cotton fibres, diurnal variation in sucrose content may be anticipated to affect the rate of cellulose formation 77.89. Thi s observation is in full agreement with the findings 9094 reported eariier in respect of the variability between cotton genotypes for cellulose synthesi s. 75 Table 6 presents the recent data on cellulose content in fibres of sixteen cotton varieties of Gossypium arboreum and Gossypium hirsutum species at various maturity stages, grown in 200 I crop seaso n under the identical agro-climatic conditions at
358
INDIAN J. FIBR E T EXT. RES., SEPTEMB ER 2003
Table 5-
Data on cellulose co ntent and structural parameters of four cull. ivars grow n at two locati ons in [nd ia (The va lues are the averages of 1992, 1994 and 1995 crops) C ulti va rs BIK ANERI NARM A
Rate of cellulose sy nthesis (% ) g / boll / day at boll maturation stage 10 Days PA 20 Days PA 30 Days PA 40 Days PA A verage va lue
LRA-5 166
AC-738
MA LJ ARI
Nagpm Coimbatore
Nag pur Coim batore
Nag pur Coim batore
Nagpur
Coimbatore
2.09 1 3.98 1 3.806 3.042 3.230
0.683 3.397 5.320 4.887 3.571
1.5 11 3.529 4.335 3.222 3. 149
1.1 66 3.555 4.5 87 3.41 6 3. 18 1
1.209 3.299 4.769 4.547
22 .3 1 49 .8 1 8.09
2 1.93 5 1. 99 8.80
20. 12 46.39 7.20
47 .73 0.4765
43.82 0.4864
35.84
35.82
3.456
0.667 3.097 4.867 6.021 3.663
0.739 3. 15 7 4.5 12 3.538 2.986
0. 82 1 2.707 3. 187 2. 852 2. 39 1
22.46 55.7 8 9.97
20.88 5 1.83 8.80
19.35 58.77 9.60
14. 19 43.85 7.63
13. 18 43 .29 7.26
47.2 1 0.5356
40.3 1 0.4845
40.99 0.4607
43.06 0.46 11
43.76 0.50 14
42.40 0.5224
34.65
35.88
36.82
36.80
35.20
35.69
28.75 24.25 18.95
3 1.00
26. 12
27 .00 19.40
22.25 19.46
27.45 22.83 20.64
Fibre property parameters Av . reversals / cm Av. convoluti ons / cm Av . convoluti on angle (8), deg Av. bund le fi bre tenac ity, g / tex Av . Hermans ori entation factor Av. angle of ori entatio n (8), deg 40 % X- ray angle, deg 50 % X- ray angle, deg ( 40 % - 8 ), deg
28 .00 24.40 20.70
28.83 24.66 20.05
28.00 24.50 18.00
28.75 25.29 18.35
( 50 % - 8 ), dcg
17.03
15.80
14.52
14.97
15.45
16.40
15.58
15.2 1
( a m- 8 ), deg
28.80
27 .04
24.67
25.96
27.02
27.20
3 1.03
28.45
ag pur - Latitu de: 2 10 - 10' North , Longitude: 79() - 12' East; Coimbatore- - La ti tude: 11 0 - 00' No rth. Lo ngitude: 76') - 58' Eas t; and PA - Post-anth esis.
New Del hi . Fi g. 1 stri kin g ly shows the re lat ive variati o n of average cellul ose conte nt at any five-day interva l of post-anth es is boll maturity stages whe n these 16 vari eti es were alTan ged in in c reas in g o rder o f their average value of cellul ose co nte nt. It is a lso ev ident from Table 6 that th e vari eti es o f Gossypium hirsulum sy nthes ized mo re a mount of cellul ose as compared to th e vari eti es o f Gossypium arboreum and th at the variation between the lowest and hi ghest values wi thin the vari eties of indi vidua l spec ies was 216 % and 286% respecti vely. A lmost s imil ar g3 obse rvation was repo rted earli er by Jihu a and Hs ieh el 01. 93 . Days post-anthes is (DPA ) shows very sign ifica nt positive corre lati on of (> 0.80) with average value of cellul ose content at any five-day in terval w ithin al l the individual variet ies, irrespective of spec ies, whe n grown unde r unifo rm agro-c limatic condit ions at one location 75 . Thi s indicates their mutual in ter-dependence a nd since the slopes of
curves betwee n ce llul ose co nte nt days post- anth es is are diffe rent fo r indi vidual vari eties, it impli es that the rates of ce llul ose sy nthesis at a partic ular locati o n of g rowth of cotto n are a lso di fferent in di ffe re nt varieti es 75 .
6 Breeding Priority and Challenges to Breeders Reasonabl e success in bringi ng fo rth improve ments in o ve rall properti es, y ield a nd te nsile stre ngth of fibres throu g h conve nti o na l breedin g has been ach ieved in almost a ll the countri es g row ing cotto n o n com merc ial scale. Yet, th ere is a bi g gap between the leve ls of te nsi Ie strength of fibres of the ex istin g cul ti vated ge notypes a nd strength require men ts o n raw cotton fibre by the modern effic ient cotton . . 6 26 .6 1'g5. T h ese d eve Iopme nts processll1g tec I111 0 Iog les' have already reversed the pri o rity from breed in g cotton fo r increa::;ed stap le length of fibres (keepin g in view the demand of the old ring-frame spllln lllg
MOHARIR: STRUCTURE AND STRUCTURE-PROPERTY RELATIONSH IP IN NATIVE COTTON
00
""'
00
,
CO
Iii
0'
>
q:
o L-----------------------------------~ o , 2 3 4 5 6 7 8 9 10 " '2 '3 '4 '5 '6 Co tto n va ri e ti es
Fig. i -Collon va rieti es arranged in increasing order of ave rage ce llul ose content at an y 5-day interval rC. a rbo r elllll: I- HD- I07 , 2- CA-5, 3- NCA-7, 4- NCA- I, S- LD-327, 6- AAH- 1, 7HD- 123; and C. hirSlIllllll : 8- LH- 1556, 9- PBKH-4, IO- F84(). II - Bray brown , 12- RS -87S. 13- HS -6. 14- H- 1098 . 15- RS-20 13, and 16- Pw,a-3431
Heritability of a fibre property measured by sing lefibre property meas uring instruments (fibrograph , micronai re and stelometer) and meas ured by integ rated Hi gh Volume Instrument (HVl) shows th at it is grea ter with "telometer than with HVI instrument94 . Since automated fibre property meas uring instruments are very likely to co mpl etely replace the manu al methods in the years to come, it is imperative to develop new and reliabl e re lati onships between th e technologica ll y important fibre parameters and HVI instrumental calibration s. And for doing so, cotton breeders have to stop working in isolation in laboratori es and fi elds but see k close cooperati on with fibre science speci ali sts and tech nologists. As disc ussed above, the im provements in cotton fibre properties revo lve around th e amount and mann er of cellul ose sy nth esis and deposition within the matrix of second ary laye rs of developing cotton fibre , either in a natural way or through any other mea ns such as ferti li za tion and nutrition, irri gation, genotype selection, agronomic and cultural practices, enzy me and hormone applications or throu gh introg ression of effic ient cellulose syn thesizing genes into fibre cell s for modification of cellulose in amount and pattern s of its di sposition. And the methods range from conventional breedin g to molecular genetics, biotechnology and recombinant DN A technology, and introgression through marker-aided selection . R M Brown 79 of Molecular Geneti cs and Microbiology at UT Austin 's College of Natural Sciences hopes that "some day it will be poss ible to produce a fibre product that has better quality and more diverse trait for ya rn and texti le production . If scienti sts can
experimentall y alter dimensions of the co tton fibre as wel l as cell Ulose biosy nthesis, it should be possible to alter its strength and thu s change the way cotton can be handl ed". Multi-di sc iplinary and mu lti-p ronged research have already begun to ex hibit eco nomi c and tech nical benefits in th e form of th e emergence of the Bt-cotton, genetically modi fied naturall y co loured cotton in vari ous shades and the terminator seeds. These develop ments, howsoever spectac ul ar in scien ti fic merit , have also raised new co mplex issues of global patents and sa feguardin g th e commercial interest.s of th e patent holders throug h new laws and legislat ions, intellectual property rights and piracy; fanner' s ri ghts, pri vileges and apprehensions ; moral and ethi cal iss ues In geneti c mod ification / manipul ation of naturall y evolved germpl asm; hum an and anim al tox icity from geneticall y modifi ed seeds, fibres and oth er products; and environmental, co mmunity and personal safety aspects arisi ng out of these actio ns. Cotton breeder in the twenty-first cen tury wi ll not on ly have to increasing ly fac e th ese problems, but will be required to address th em fo r effec ti ve solurions. And these are further go ing to be serious in view of the impending globa l climati c changes wi th land surface temperature likely to go up by 3-4°C. How a cotton breeder will face these challenges, onl y time will tell . However, the on ly fact th at will prevail is that th e cotton fibre will continue to rul e as th e kin g of all tex til e fibres for the unm atched beauty and co mfort it provides. 7 Conclusion Staple length of cotton fibres, finen ess, frequency of reversa l ex tin ction bands and co nvo luti on twi sts, density of cellulose, degree of po lymeri zation , cellulose crystallite sizes and cry stallite ori entation to fibre ax is fo r a cotton vari ety remai ni ng constant / practically invariant with change in location of growth 86 , the rate and amo unt of cellul ose sy nth esis and degree of polymerizati095 sho uld be directl y incorporated in cotton breeding program for sc reeni ng cotto n genotypes for hi gher cellulose sy nthesis, location specific genotypes, and conseq uentl y for increased maturity and strength of fibres.
Acknowledgement The author wishes to thank the Director, Indian Agricultural Research Insti tute, New Delhi , for providing necessary facilities to carry out thi s work.
MOHARIR: STRUCTURE AND STRUCTUR E- PROPERTY RELATIONSHIP IN NATIV E COTIO N
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30 31 32 33 34 35 36 37 38 39 40
41 42 43
44 45 46
47 48 49
50 51 52 53 54
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