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Published in Crop Sci. 44:1553-1559 (2004).
© 2004 Crop Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA

CROP BREEDING, GENETICS & CYTOLOGY

Fiber Length Development in Near-Long Staple Upland Cotton

Chris A. Braden* and C. W. Smith

Dep. of Soil and Crop Sci., Texas A&M Univ., College Station, TX 77843-2474

* Corresponding author (cbraden{at}ag.tamu.edu).


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Advanced Fiber Information System (AFIS) mean fiber length by weight (Lw), fiber length development period (FLDP), and average daily fiber length growth rates (ADGR) were determined for four experimental strains and three cultivars of upland cotton, Gossypium hirsutum L., in 1998 and 1999 in College Station, TX. TAM 94L-25 and TAM 94M-14, near-long staple sib lines developed by the Texas Agricultural Experiment Station (TAES), were compared with five upland cotton genotypes that varied in high-volume instrument (HVI) upper half mean (UHM) fiber length. The FLDP was determined as the number of days postanthesis (DPA) until fibers reached a Lw greater than or equal to their Lw at boll opening. Final average daily growth rate (FADGR) was calculated as the average Lw d–1 from anthesis to open boll. Length by weight was determined at 3-d intervals from 16 DPA until boll opening. With a higher FADGR and longer FLDP, TAM 94L-25 and TAM 94M-14 exhibited the longest final Lw. TAM 94L-25 had a higher FADGR than the commercial cultivars Suregrow 125 and Tamcot CAMD-E, while TAES strain 94WD-17 exhibited the lowest FADGR. The broad-sense heritability estimate for final Lw across years was 0.88.

Abbreviations: ADGR, average daily growth rate • AFIS, advanced fiber information system • DPA, day(s) postanthesis • FADGR, final average daily growth rate • FLDP, fiber length development period • HVI, high-volume instrument • Lw, length by weight • SADGR, segmented average daily growth rate • TAES, Texas Agricultural Experiment Station • UHM, upper half mean


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
COTTON BREEDERS MUST BE concerned with fiber quality as well as yield if U.S. cotton is to maintain a strong, competitive edge in today's world market. Fiber quality is important to the textile industry because it directly relates to processing performance, productivity, and yarn quality. Fiber length is one of the most important properties of cotton fibers relative to marketing and processing (Perkins et al., 1984). Production efficiency, amount of waste, fly generation, and cleaning are a few parameters affected by fiber length. A premium is paid for longer fibers because length is correlated to yarn quality parameters such as strength, elongation, hairiness, and evenness, and because such fibers allow the mill flexibility in the type of yarn produced. Therefore, selecting genotypes that produce longer fibers is a fiber quality objective of many U.S. cotton breeding programs.

A cotton fiber is a seed hair, a single hyperelongated cell arising from the protodermal cells of the outer integument layer of the seed coat. The elongation phase of the fiber development process encompasses the major expansion growth phase, and Xie et al. (1993) reported that there are three stages of fiber elongation: initiation, early elongation, and late elongation. They, along with Gipson and Joham (1969), stated that early-stage fiber elongation was temperature dependent and late fiber elongation was temperature independent. A number of researchers have documented the growth and development of cotton fibers through the elongation phase (Anderson and Kerr, 1938; Balls, 1915; Barre et al., 1931; Jasdanwala et al., 1977; Kohel et al., 1974; Schubert et al., 1973). The majority of these papers have reported the growth and development of one cultivar in a given year. These studies also were limited in scope because they reported the nonginned fiber length from individual seeds from 1 to 5 locules of 1 to 6 different bolls, or 10 seeds from each boll. The fiber lengths were measured on the convex side of a watchglass after a jet of water had flared and straightened the fibers away from the seed surface.

A large amount of information describing the chronology of cotton fiber development, both field studies and cotton ovule cultures, and the effects of the growth environment on fiber quality, especially suboptimal temperatures, has been accumulated (Anderson and Kerr, 1938; Gipson and Joham, 1968, 1969; Gipson and Ray, 1969; Haigler et al., 1991; Hawkins and Serviss, 1930; Hesketh and Low, 1968; Hessler et al., 1955; Morris, 1962; Quisenberry and Kohel, 1975; O'Kelly and Carr, 1953; Ramsey and Berlin, 1976; Sturkie, 1934; Thaker et al., 1989; Xie et al., 1993). Basra and Malik (1984), and Gipson and Joham (1969) documented the variability in the rate and the duration of elongation among different cotton cultivars. Gipson and Joham (1969) stated that the rate of fiber growth appeared to be dependent upon cultivar, fiber age, and night temperature. Quisenberry and Kohel (1975) reported that variations in fiber length and the fiber elongation period were associated with heat-unit accumulations. Regression analyses showed that genotypes that produced longer fibers were more responsive to heat unit accumulation levels than were genotypes that produced shorter fibers. These studies, together with the genetic studies by Kohel et al. (1974), suggest that rate and duration of fiber elongation can be altered through traditional breeding techniques.

The natural variability and complex relationship among fiber quality characteristics (Behery, 1993; Bradow et al., 1997a, 1997b; Davidonis and Hinojosa, 1994; DeLanghe, 1986) also has been documented. Significant genetic and environmental variations of fiber properties occur among individual plants, among bolls on the same plant, across seeds of an individual boll, and on a single seed. Variations in fiber length of a cultivar from one season to another can be attributed to the environmental conditions during the fiber elongation period.

Average staple length in the USA was 27.8 mm in 1995, whereas the average staple length in Texas was 26.9 (Smith, 1999). Cotton Incorporated designates upland cotton with an UHM fiber length of 28.2 to 32.0 mm as long and anything > 32.0 mm as extra long (Cotton Incorporated and Textile World, 2003). When tested in 19 cultivar performance trials across nine locations in Texas (12 irrigated and 7 dryland) in 2000 and 2001, TAM 94L-25 averaged 30.2 mm UHM length (Smith et al., 2001, 2002). Its sister line, TAM 94M-14, averaged 31.2 mm UHM across six locations x years from 1996–1998 (Smith, 1998, unpublished data). The objectives of this study were to (i) determine AFIS Lw of TAM 94L-25, TAM 94M-14, and a number of medium staple genotypes varying in fiber length when grown under irrigated conditions, (ii) determine the FLDP using Lw, and (iii) determine phenologically how TAM 94L-25 and TAM 94M-14 attain their longer average fiber length.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS AND DISCUSSION
 REFERENCES
 
Four upland cotton strains [TAM 94L-25 (Smith, 2003), TAM 94M-14, TAM 94WD-17, and TAM 91C-95Ls (Smith, 2001)] and three commercial cultivars [Suregrow 125 (PVP 9400063), Tamcot CAMD-E (Bird, 1979a), and Acala Maxxa (PVP 9000168)] were planted in a randomized complete block design with four replications in 1998 and eight replications in 1999 at the Texas A&M Research Farm near College Station, TX. Soil type for both years was a Weswood silt loam, a fine-silty, mixed, superactive, thermic Udifluventic Haplustept, intergraded with Ships clay, a very fine, mixed, active, thermic Chromic Hapludert. Plots were two rows, 12 x 1 m, with a blank row on each side. In 1998, approximately 63 ha-cm of preplant irrigation was applied. Genotypes were planted on 20 Apr. 1998 with skips replanted on 30 April. Early in the growing season, plots were thinned to single plant culture (approximately 40 cm between plants). Cultural practices including furrow irrigation were designed to maximize boll retention. Because of high demand and limited supplemental water resources, plots were furrow irrigated only twice, on 3 June and 1 July, receiving around 31 ha-cm of water at each application. Tagging of white flowers started on 22 June and continued until 17 July 1998. Harvesting of green bolls of different ages began 20 July and ended 14 August. Because of timely and beneficial rainfall in 1999, no supplemental irrigation was needed before planting nor during the growing season. Plots were planted on 12 Apr. 1999. Tagging of white flowers was initiated on 20 June and continued through 16 July. Harvesting of green bolls started 28 July and concluded on 4 Aug. 1999.

Strains and cultivars were chosen on the basis of their HVI fiber length. The general genetic background and description of each genotype follows.

The AFIS is the preferred method to measure fiber length because of its ability to analyze immature fibers from a collection of bolls. The UHM length does not always agree with AFIS length data from HVI because of effects of sample pooling, fiber crimp, specimen crimp characteristics, and other factors (Behery, 1993). Also, AFIS is not dependent upon maintaining a ribbon as in HVI machinery. Manual classer's staple length, HVI UHM length, and the USDA staple standards (Sutter Web Array) are Lw referenced measurements. Fiber Lw is weight biased in that individual length groups are weighted and individual fibers are not counted. Weight bias influences the length calculations and puts more weight percentage to the longer fibers. Therefore, Lw is always greater than mean length by number for a given sample.

Analysis of variance for AFIS Lw was conducted by means of the PROC GLM procedure of SAS (SAS Institute, 1999). Where GLM procedure showed significant main effects, Waller–Duncan K ratio LSD was used to separate genotypic means.

Tagging of flowers was not initiated until all genotypes exhibited blooms. Flowers were then tagged on the day of anthesis with dated tags. The chronology of fiber length development was determined by harvesting approximately 30 tagged green bolls, regardless of sympodial position, from within each plot for each predetermined age. This procedure ensured better representation of fibers within a DPA sample. These predetermined ages were 16, 19, 22, 25, 28, 31, 35, 38, and 42 DPA. All harvested bolls for each genotype and DPA were collected from the same anthesis date, except when the allotment of 30 bolls was not fulfilled and then the next anthesis date was used. Untagged bolls among plants within a plot were not detached to maintain normal boll load and consistency from plant to plant. Tagged bolls were harvested in a manner in which the lower fruiting zone constituted 35, 38, and 42 DPA; middle fruiting zone represented 25, 28, and 31 DPA; and the upper fruiting zone included 16, 19, and 22 DPA bolls. Thus, a single harvest date would include multiple DPA and as narrow of a time span as possible to lessen the environmental variance component. Bolls of all genotypes harvested at 38 DPA in 1998 and 42 DPA in 1999 were considered fully mature when sutures started to dehisce naturally. Harvested bolls were allowed to dry in a greenhouse, after which the seedcotton was hand removed from the bur and ginned on a roller gin. The Lw measurement from AFIS instrumentation was determined by Cotton Incorporated and used to determine fiber length developmental time and time needed for each genotype to reach its maximum fiber length.

Average daily growth rates and segmented average daily growth rate (SADGR) were calculated by dividing the Lw at a given DPA by the DPA or by determining the Lw for a given segment of DPA [e.g., (Lw at 19 DPA) – (Lw at 16 DPA)], and dividing by the appropriate segmented DPA. FADGR was determined by dividing the final Lw by the number of days from anthesis to open boll. Boll maturation period (open boll) was determined by 10 plants in each of two reps in 1998 and four reps in 1999 of each genotype that were selected randomly and marked early in the growing season (data not shown). To determine the date the tagged bolls matured (defined as when the sutures dehisce naturally or under pressure between the thumb and forefinger), tagged bolls from the 10 plants in each plot were monitored daily beginning about 35 d after the first flowers were tagged. Date of maturity was recorded on the same tag, which also identified the date of anthesis. Estimates of variance components and broad-sense heritabilities of Lw were calculated from mean squares across years (Fehr, 1987). Confidence intervals for heritability estimates were calculated according to Knapp et al. (1985).


   RESULTS AND DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
REFERENCES
 
Thirty-five days with temperatures at or above 37.8°C, with 25 consecutive days at or above 37.8°C, and only 1.29 cm of total rain characterized the 1998 growing season at College Station, TX. The high heat probably led to excessive abscission of fruit, which resulted in an extended green boll harvesting period. In an attempt to obtain 30 bolls for each age in each plot, bolls of the same age were harvested from different anthesis dates.

On the basis of the amount of rainfall received during the growing season, 1999 was the inverse of 1998. Timely and beneficial rainfall enabled the cotton plants to retain a large quantity of fruit at most positions and to develop cotton fibers under essentially no moisture stress. This more ideal growing season resulted in a shorter harvest interval and almost all bolls within a given age in each plot came from a single anthesis date.

Final Fiber Length
Final Lw for mature bolls (38 DPA in 1998 and 42 DPA in 1999) differed (P = 0.05) across years, but were pooled since the ANOVA indicated no significant genotype x year interaction (Table 1). The lack of a significant effect of genotype x year for fiber length supports the premise that there is a strong genetic basis for this trait (May, 2000) and that an appropriate irrigated test was accomplished in 1998 despite the high evaporative demands created by high temperatures and near-full sun (Hearn, 1994). Genotypes differed (P = 0.05) in Lw as expected.


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  		Table 1. Mean squares for fiber length by weight (Lw) for specific days postanthesis (DPA) and final Lw of seven cotton genotypes grown at College Station, TX, in 1998 and 1999.

 
TAM 94L-25 and TAM 94M-14 developed longer Lw fibers than the commercial cultivars and other genotypes in this study (Table 2). Genotypes TAM 94L-25 and TAM 94M-14 averaged 29.2 and 29.1 mm, respectively, and based on Cotton Incorporated's fiber classification chart (Cotton Incorporated and Textile World, 2003), they are long-staple upland cotton genotypes. These two genotypes, developed at the TAES, have consistently averaged UHM fiber length about 30.0 mm across multiple locations and multiple years (Smith, 2003; Smith, 1998, unpublished data). As expected, Tamcot CAMD-E, along with TAM 94WD-17 exhibited the shortest Lw fibers at only 25.4 and 25.8 mm, respectively. TAM 91C-95Ls had Lw fibers just above the average for the study, while Acala Maxxa and Suregrow 125 had Lw fibers below the average of the seven genotypes.


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  		Table 2. Average fiber length by weight (Lw) for seven cotton genotypes when harvested at eight different days postanthesis (DPA) and at open boll in 1998 and 1999.

 
Fiber Length Development
The objective of this research was to gain a better understanding of the phenology of long-staple TAM genotypes. Since TAM 94L-25 and TAM 94M-14 produce long or near-extra long HVI fiber lengths, then the logical question is, do they derive this extra length through a greater average daily fiber growth rate, or longer FLDP? Analysis of variance of Lw for each DPA revealed significant year effects for the first six DPA, 16, 19, 22, 25, 28, and 31, and genotype x year interaction for 35 and 38 DPA (Table 1). Genotypes differed (P = 0.05) in Lw early in fiber development, 16 and 19 DPA, but not again until 31, 35, and 38 DPA. The ANOVA of Lw across DPA for each genotype showed a year effect for TAM 91C-95Ls, Acala Maxxa, TAM WD-17, and Tamcot CAMD-E (Table 3). The significant D x Y term for each genotype, except Acala Maxxa, resulted from a slightly faster developmental period in 1998 than 1999. Lw at 25 DPA of TAM 94L-25 and TAM 94M-14 were not different (P = 0.05) from their mature boll Lw in 1998, while this relationship was not reached until 28 DPA in 1999 (data not shown). The other genotypes reached Lw equivalent to their final Lw at 22 DPA in 1998, but not until 25 DPA in 1999. The significant D x Y for CAMD-E exhibited longer Lw at 31 DPA than at boll maturation in 1999, but not in 1998. These data do not change the relationship of Lw across genotypes, so years were pooled to look at Lw development within genotypes across DPA.


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  		Table 3. Mean squares for fiber length by weight for seven cotton genotypes grown at College Station, TX, in 1998 and 1999.

 
At 16 DPA, Suregrow 125, Acala Maxxa, Tamcot CAMD-E, and TAM 91C-95Ls had longer fibers (P = 0.05) than TAM 94L-25 and TAM 94M-14, which averaged 16.8 and 16.9 mm, respectively (Table 2). Three days later, Suregrow 125 exhibited Lw longer than all other genotypes except Acala Maxxa. At 22 DPA, all genotypes had equal Lw and at 25 DPA, Suregrow 125 and TAM 91C-95Ls were the only genotypes with Lw longer than Tamcot CAMD-E. At 28 DPA all genotypes again were equal in Lw, however by 31 DPA, TAM 94L-25 exhibited longer Lw than all other genotypes. TAM 94L-25 was longer than all genotypes except TAM 94M-14 by 35 DPA and these two genotypes were not different at 38 DPA and longer than all other genotypes.

Kohel et al. (1974) noted a decline in fiber length when nearing the conclusion of the boll maturation period. This reduction in fiber length is present in these data and the data of other fiber development studies such as Hawkins and Serviss, (1930) and Schubert et al. (1973). The only genotype not exhibiting this phenomenon was TAM 94M-14. When averaged across years, all genotypes in this study reached their maximum Lw by 31 DPA. Although Suregrow 125 Lw at 25 DPA, which appears to be an anomaly in the data, and Tamcot CAMD-E Lw at 28 DPA were similar to their Lw at 31 DPA (Table 2), their maximum numerical average Lw was not attained until 31 DPA and thus 31 DPA was considered to be the FLDP for all genotypes in this study.

The ADGR at selected DPA and FADGR were pooled across years since the ANOVA indicated no significant genotype x year interaction (Table 4). The year effect was significant for 16, 19, 22, 25, 28, 31, and FADGR. Genotypes differed (P = 0.05) in ADGR at 16 and 19 DPA, but not again until 31 DPA (Table 4). The ADGRs, which are Lw at a given date divided by the number of days from anthesis, revealed that the five genotypes with shorter final Lw actually grew at a faster rate per day during the early part of the elongation phase but their daily rate of growth slowed during the latter part of the elongation phase (Table 5). This is reflected in the Lw data in Table 2 which shows TAM 94L-25 and TAM 94M-14 having shorter Lw at 16 DPA than all other genotypes. At 16 DPA, Suregrow 125, Acala Maxxa, Tamcot CAMD-E, and TAM 91C-95Ls had attained between 68 to 74% of their final length and had greater ADGR (P = 0.05) than TAM 94L-25 and TAM 94M-14, which only averaged 1.04 and 1.05 mm d–1, respectively (Table 5). Three days later, Suregrow 125 exhibited ADGR higher than all other genotypes except Acala Maxxa. At 22 and 25 DPA all genotypes had equal ADGR, and at 25 DPA all genotypes except TAM 94L-25 and TAM 94M-14 had attained near 100% of their final length. By 28 DPA there were again no differences in ADGR among the genotypes, but the five genotypes with the shorter final Lw had ceased accumulating percentages toward their final length.


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  		Table 4. Mean squares for average daily growth rate (ADGR) at specific days postanthesis (DPA) and final ADGR of seven cotton genotypes grown at College Station, TX, in 1998 and 1999.

 

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  		Table 5. Average daily length by weight growth rate (ADGR) at selected days postanthesis (DPA), percentage of final length (FL) at each DPA, and final ADGR (FADGR) for seven cotton genotypes grown at College Station, TX, in 1998 and 1999.

 
TAM 94L-25, TAM 94M-14, and Acala Maxxa had the highest FADGR, 0.67, 0.67, and 0.66 mm d–1 respectively, with TAM 94L-25 being different (P = 0.05) from the other two commercial cultivars (Table 5). These data agree with Gipson and Joham (1969) and Quisenberry and Kohel (1975), who found that longer staple genotypes produced longer fibers through faster rates of elongation, coupled with longer elongation period. TAM 94L-25 and TAM 94M-14, with a higher FADGR and longer FLDP, produced longer fibers than Acala Maxxa, which had a shorter FLDP. Suregrow 125 and Tamcot CAMD-E had similar FADGR of 0.65 and 0.63 mm d–1, respectively. Although TAM 91C-95Ls had a low FADGR of 0.62 mm, this genotype attained a fiber length similar to TAM 94L-25 and TAM 94M-14 through its FLDP. The FADGR of TAM 94WD-17 was the lowest at 0.57 mm d–1. These data suggest that longer Lw in these seven genotypes was attained by higher FADGR and longer FLDP, yet these genotypes varied in ADGR across the season. Thus, ADGR were determined for each 3-d segment from 16 to 31 DPA.

Breaking the daily growth rates into 3-d segments, SADGR within the elongation phase revealed significant year effects between 16 to 19, 22 to 25, and 25 to 28 DPA (Table 6). Genotype and genotype x year terms were nonsignificant. Year and SADGR were significant for all genotypes when ANOVA was analyzed for each genotype and only TAM 94WD-17 had an increment x year interaction (Table 7). All genotypes had their highest numerical SADGR between 16 to 19 DPA (Table 8). TAM 91C-95Ls and Acala Maxxa maintained a SADGR from 19 to 22 DPA similar to the increment between 16 to 19 DPA, and TAM 91C-95Ls maintained this SADGR for the 22 and 25 DPA segment. TAM 94L-25, TAM94M-14, and Tamcot CAMD-E sustained a similar SADGR for the last four increments. Suregrow 125 fiber length growth pattern showed fluctuations across the SADGR.


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  		Table 6. Mean squares for segmented average daily growth rate of seven cotton genotypes grown at College Station, TX, in 1998 and 1999.

 

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  		Table 7. Mean squares for segmented average daily growth rate for each of seven cotton genotypes grown at College Station, TX, in 1998 and 1999.

 

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  		Table 8. Segmented average daily growth rate (SADGR) for seven cotton genotypes grown at College Station, TX, in 1998 and 1999.

 
Variance components for the different effects are presented in Table 9. For all DPA except 35, 38, and final, the error variance exceeded the genotypic variance, while the genotypic x year variance constituted only a small percentage of the total variance. Broad sense heritability estimates for Lw ranged from 0.27 to 0.88 for each individual DPA. High heritability estimates for Lw were obtained at 16, 31, 35, 38, and final DPA. For 22, 25, and 28 DPA, genetic variances are not different from 0 and therefore heritability estimates are not different from 0.


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  		Table 9. Components of variance, broad-sense heritability, and percentage of the total variance (in parentheses) of length by weight (Lw) for each day past anthesis (DPA) and final Lw of seven cotton genotypes grown at College Station, TX, in 1998 and 1999.

 
These data support previous reports that cotton fiber growth is complex in nature and has a strong genetic basis early and late in fiber length development. Across 2 yr of diverse climatic conditions, no genotype x year interaction was detected for the final Lw, and all genotypes reached their maximum length by 31 DPA. TAM 94L-25 and TAM 94M-14 exhibited Lw fibers longer than the three cultivars and two other genotypes in this study. These breeding lines attained longer fibers through a greater FADGR coupled with a longer FLDP. TAM 94L-25 and TAM94M-14 also exhibited a more sustained ADGR through 31 DPA than all other genotypes which tended to fluctuate in SADGR. Investigation into the decrease of fiber length after 31 DPA needs to be addressed to determine if this phenomenon is biological or due to mechanical deficiency in both the AFIS and HVI systems.

Received for publication October 29, 2003.


   REFERENCES
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 




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