Published online June 12, 2015

Research

Early Vigor of Advanced Breeding Lines

and Modern Cotton Cultivars
S. Liu, M. Remley, F. M. Bourland, R. L. Nichols, W. E. Stevens,
A. Phillips Jones, and F. B. Fritschi*

S. Liu and F.B. Fritschi, Division of Plant Sciences, Univ. of Missouri,

ABSTRACT
Stand establishment and early vigor are critical to
the successful production of cotton (Gossypium
hirsutum L.). Rapid early growth could provide
significant advantages to young plants relative to
diseases, insects, and weeds. The objectives of
this research were to identify traits that contrib-
ute to differences in early growth, characterize
genotypic variation in early vigor among mod-
ern cultivars and advanced breeding lines, and
determine the effect of the seed production envi-
ronment on early vigor. Early growth of 10 geno-
types from private companies and 18 unreleased
breeding lines was compared through mea-
surements of leaf area and biomass under field
conditions in 2 yr. A positive correlation of seed
weight and rapid early growth was observed up
to 32 d after planting (DAP). Cotyledon area was
positively associated with shoot dry weight over
the course of the first 52 DAP. Significant geno-
typic differences were observed for cotyledon
and total leaf areas, and for cotyledon, first true
leaf, total leaf, stem, and total shoot dry weights
in both years within approximately the first 4
wk after planting. No correlation between lint
yield and rapid early growth was found. For the
first 17 (2009) and 32 (2008) DAP—but not after
that—plants grown from seeds produced in dis-
tinct environments differed in early vigor. Overall,
seeds from breeding lines produced more vig-
orous seedlings than did seeds of commercial
genotypes. Although heritability of the examined
traits has not been established, the broad range
in values and significant genotypic variation
suggest that breeding for enhanced early vigor
should be possible without compromising yield.
Columbia, MO; M. Remley, Darr School of Agriculture, Missouri
State Univ., Springfield, MO; F.M. Bourland, Northeast Research and
Extension Center, Univ. of Arkansas, Keiser, AR; R.L. Nichols, Cot-
ton Incorporated, Cary, NC; W.E. Stevens, Division of Plant Sciences,
Univ. of Missouri, Fisher Delta Research Center, Portageville, MO; A.
Phillips Jones, Univ. of Missouri, Fisher Delta Research Center, Por-
tageville, MO. Received 3 Oct. 2014. Accepted 3 Feb. 2015. *Corre-
sponding author (fritschif@missouri.edu).
Abbreviations: DAP, days after planting; RGR, relative growth rate.
E arly development of cotton tends to be slow, leaving plants
vulnerable to stress from various sources (El-Sharkawy et al.,
1965; Ortiz and Bourland, 1999). In production regions with a
short growing season or high disease, insect, and weed pressures,
slow early growth may be particularly detrimental to the crop.
Clearly, seedling growth and development are strongly influenced
by prevailing environmental conditions after planting. Since late
planting often delays crop maturity, which in turn can reduce lint
yields and quality, producers in the Mississippi Delta often plant
cotton early in the season (Wrather et al., 2008). However, early
planted seedlings can experience suboptimal growing conditions.
Cool conditions tend to favor pests and diseases that can cause
seedling damage and death and reduce plant populations, which
further delays cotton crop maturity (Jones and Wells, 1998; Smith
et al., 1979; Siebert et al., 2006; Wrather et al., 2008). Low soil
temperatures early in the season are associated with increased seed-
ling diseases and slow seedling emergence and stand development
(Christiansen and Thomas, 1969; Colyer et al., 1991). Early planted
Published in Crop Sci. 55:1729–1740 (2015).
doi: 10.2135/cropsci2014.10.0686
© Crop Science Society of America | 5585 Guilford Rd., Madison, WI 53711 USA
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crop science, vol. 55, july– august 2015  www.crops.org 1729

cotton appears to be very sensitive to weed competition,
particularly during the first 40 d, which are very critical for
cotton growth and development (Holt and Orcutt, 1991;
Murray et al., 1988; Papamichail et al., 2002).
While early planting associated with suboptimal grow-
ing conditions can negatively influence yields, late plant-
ing also results in reduced yields, lint percentages, and low
micronaire because of delayed boll maturity under cool fall
temperatures, even though conditions for seedling emer-
gence and stand establishment may have been ideal (Bilbro
and Ray, 1973; Guthrie, 1991; Cathey and Meredith, 1988;
Kittock et al., 1987; Wrather et al., 2008). On the other
hand, early planting without cold stress showed higher
yields than later planting due to the crop maturing during
warmer fall conditions (Bange and Milroy, 2004). Thus,
every year, cotton producers face the dilemma between
early planting with the risk of suboptimal conditions for
seedling growth and later planting that may be associated
with the crop maturing under cool fall temperatures.
Development of cultivars with more vigorous seed-
ling growth would help overcome some difficulties asso-
ciated with early planting. Healthy, vigorous growth can
allow plants to be more tolerant to pathogens and escape
disease (Patrick, 1989). When seedling growth is delayed,
cotton plants are vulnerable to pathogen attacks for a
longer period and are often impeded with regard to nutri-
ent uptake. Weak cotton seedlings are especially suscep-
tible to attack by pathogenic fungi because the extended
period with greater sensitivity generally means more
chances to be infected. Because vigorous seedlings harden
and develop woody tissue faster, the sensitive period is
shortened and fungal penetration and rotting is reduced
(Bourland, 1992; Collins, 2015). Similarly, more vigorous
seedlings can have an advantage when facing insect pres-
sure. For instance, Buntin (2013) reported that most corn
(Zea mays L.) fields with vigorous growth may completely
escape serious insect damage. Additionally, field experi-
ments by Elliott et al. (2008) showed that more vigorous
canola (Brassica napus L.) seedlings were more tolerant to
flea beetle (Phyllotreta spp.) damage than seedlings with
lower vigor. In cotton, early-season thrips (Frankliniella
spp.) infestation is very common and can decrease yields
(Cook et al., 2011), but cotton with more vigorous early
growth may be more tolerant to thrips damage.
Vigorous early growth resulting in more rapid canopy
development can inhibit weed growth by increased shad-
ing (Wilcut et al., 1995). For example, Kamboozia (1994)
reported that pea (Pisum sativum L.) genotypes with faster
emergence and greater percentage emergence were able
to better compete with weeds and produced greater yield
than did genotypes with slower emergence. For wheat
(Triticum aestivum L), Gill and Zerner (2006) found that
because of greater early leaf area development, plants
developing from larger seeds were better able to compete
1730 www.crops.org crop science, vol. 55, july– august 2015
with oat (Avena sativa L.) weeds than those developing
from smaller seeds. And, López-Castañeda et al. (1995),
indicated that more vigorous crop seedlings inhibit weed
growth, which in turn can contribute to a reduction in
herbicide use and may help counter the increasing prob-
lem of herbicide resistance in weeds.
As outlined above, there are many advantages of vig-
orous seedlings over weak seedlings, and it is clear that
vigorous growth following emergence is of interest in
many crops. Improving seedling vigor is an important
objective of breeding programs in several crops, including
rice (Oryza sativa L.) (Redona and Mackill, 1996; Lu et.al.,
2007), corn (Revilla et al, 1999; Engelen et al., 2003; Ruta
and Hund, 2010) and wheat (Richards and Lukacs, 2002;
Botwright et al., 2002; Condon et al., 2004). However,
for cotton, there is a lack of information on the genetics as
well as the physiological mechanisms determining vigor
early during crop development. Therefore, the objectives
of this study were to (i) characterize genotypic variation in
early vigor among modern cultivars and advanced breed-
ing lines, (ii) determine the effect of the seed production
environment on early vigor, and (iii) examine the plant
traits that contribute to differences in early vigor.
MATERIALS AND METHODS
Site Description and Crop Management
Field experiments were conducted at the Delta Research and
Extension Center in Southeast Missouri on a Tiptonville soil
(fine-silty, mixed, superactive, thermic Oxyaquic Argiudolls).
Seedling vigor comparisons were conducted on 28 genotypes,
including 18 cotton breeding lines from the University of
Arkansas Cotton Breeding Program (Bourland 2004, 2013) and
10 genotypes from private companies. For genotypes from pri-
vate companies, plant variety protection numbers are provided
for released cultivars but not for experimental lines. For simplic-
ity reasons, hereafter, all genotypes from private companies will
be referred to as cultivars. In addition, seeds of breeding lines
were produced in Arkansas and Arizona to compare the effect of
the seed production environment. Seed from both sources were
delinted at Keiser, AR, using standard wet sulfuric acid tech-
niques. Entries were planted in a randomized complete block
design with four replications. Acid-delinted, fungicide-treated
cotton seeds from each entry were planted in 11.3-m long and
3.86-m wide four-row plots at 2.5 cm depth and at a density
of 13.6 plants m−2. Plantings were made on 20 May 2008 and
1 June 2009 (rainfall in May delayed the 2009 planting, Fig.
1). Before planting, 67 kg N ha−1 in the form of urea was uni-
formly broadcast applied across all plots. Crop management,
including irrigation, herbicide, and insecticide applications, was
conducted based on weather, field conditions, and according to
University of Missouri Extension recommendations. Weather
information, including total precipitation and temperature, was
obtained from a weather station located within 1000 m of the
cotton field (historical agricultural weather database of Missouri
Weather Stations, Pemiscot County, Lee Farm).
Figure 1. Average temperature and precipitation during the 2008-A and 2009-B growing seasons at Portageville, MO. (In 2008, S1 = 22
DAP, S2 = 32 DAP, S3 = 52 DAP; In 2009, S1 = 17 DAP, S2 = 27 DAP, S3 = 35 DAP, S4 = 48 DAP)
Plant Sampling and Data Collection measurements. Leaf area measurements were conducted using
For each genotype, average seed weight was determined based an LI-3100 leaf area meter (Li-Cor, Inc.) on individual plants
on a random sample of 400 acid-delinted, cleaned seeds. To and included separate measurements for cotyledons and true
assess early season growth of the different genotypes, plant sam- leaves as described in Table 1. In addition, plant height and
ples were collected 22, 32, and 52 DAP in 2008 and 17, 27, 35, number of nodes were recorded for each plant. Individual plant
and 48 DAP in 2009. Measurements of leaf area, dry matter, fractions were oven-dried at 60C until samples reached stable
plant height, and number of nodes were obtained each time, weights. From these data, relative growth rate (RGR) was cal-
accounting for developmental differences as indicated in Table culated according to Fisher (1921):
1. At each sampling, five plants per plot were cut at ground
level and transported on ice to the laboratory for detailed RGR = (lnW2 − lnW1)/T

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Table 1. Overview of sampling times in days after planting Statistical Analyses
(DAP) and growing degree days (GDD) and variables mea- Statistical analyses were conducted using SAS software version
sured at the different sampling times. 9.2 (SAS, 2008). Analysis of variance was conducted using the
Samplings mixed model procedure (PROC MIXED) to examine geno-
Year DAP GDD† Variables sampled type effects, initially across both years to test for year effects
2008 22 209 Leaf area: Cotyledon, first leaf, total leaves and genotype-by-year interactions. Year, replication, and year-
Biomass: Cotyledon, first leaf, total leaves, by-replication effects were considered random. These analyses
stem and total shoot revealed highly significant year effects but no genotype-by-
Height and nodes year interactions. Nonetheless, because of the significant year
32 305 Leaf area: Cotyledon, total leaves effect, further analyses were conducted by year and sampling
Biomass: Cotyledon, total leaves, stem and time using block-by-genotype as the denominator of F for the
total shoot genotype effects. Breeding lines and commercial cultivars were
Height and nodes grouped and compared by PROC MIXED separately for each
52 525 Biomass: total leaves, stem, square and sampling time in each year using block-by-group as the denomi-
total shoot
nator of F for the group effects. For the seed source comparison,
Height and nodes
seed source, genotype, and replication-by-sampling time were
2009 17 151 Leaf area: Cotyledon, first leaf, total leaves
tested with PROC MIXED for each year separately using
Biomass: Cotyledon, first leaf, total leaves,
stem and total shoot block-by-genotype-by-location as the denominator of F. Mean
Height, hypocotyl height, and nodes comparisons were made using the least square means statement
27 297 Leaf area: Cotyledon, first, second, and third (LSMEANS) with the Tukey adjustment (P  0.05).
leaf, total leaves Relationships among variables were examined using the
Biomass: Cotyledon, first, second, and third PROC CORR procedure.
leaf, total leaves, stem, total shoot
Height, hypocotyl height, and nodes
35 382 Total leaf area RESULTS AND DISCUSSION
Biomass: total leaves, stem, and total shoot Environmental Conditions and Growing
Height and nodes
Degree Days
48 515 Biomass: total leaves, stem, square, and
total shoot; Precipitation and temperature during the first few weeks
Height after planting varied between the 2 yr (Fig. 1). In 2009,
†
Growing degree days calculated using 15.5°C as low-temperature threshold. precipitation delayed planting until June 1, but rainfall
accumulation of 60 mm by 27 DAP and 85 mm by 48
where W1 and W2 are the dry weights in grams from the beginning DAP was greater in both the first half and the second half
and the end of a specific time interval (∆T) in days, respectively. of the sampling period than the corresponding sampling
Cotton plants were defoliated mid-September and har- periods in 2008 (41 mm by 22 DAP and 68 mm by 52 DAP,
vested in October. Before harvest, plots were end-trimmed respectively). Growing degree days from planting in the
to 9.4-m length to minimize end-plant effects, and the two 2 yr differed considerably at the first samplings but were
center rows of each plot were mechanically harvested with a similar for the second sampling and last sampling in 2008
spindle picker (Case IH, model 2022) designed for small plots. and 2009 (Table 1). Because of large variations in temper-
Seed cotton was ginned with a 20-saw Continental micro-gin ature and precipitation, it was not possible to collect all the
(Continental Eagle Corp.) equipped with an incline and a sin- samplings strictly according to heat-unit accumulations.
gle-stage lint cleaner. After ginning, lint yields were calculated Thus, differences in growth and development between
and are reported in kilograms per hectare. Because there is no
the 2 yr were expected, and highly significant year effects
universal agreement on the definition of seedling vigor, shoot
dry weight and total leaf area during early plant development
(P < 0.001 for all examined traits) were found. Because
were used as the main parameters for vigor in this study. of the differences between years, results are presented by
To characterize the temperature conditions, growing year, even though genotype-by-year interactions were
degree days accumulated by each sampling date were calcu- largely absent, except for dry weight of the first true leaf at
lated as follows: the first sampling and plant height at the first, second, and
last samplings (data not shown).
DAP n é (Tmax + Tmin  ù
å ê
ê - 15.5°Cúú n Relationships among Seedling Traits
DAP 0 ëê 2 ûú To assess the early growth of cotton seedlings, traits
including area and dry weights of cotyledons, the first true
where Tmax is the daily maximum air temperature, Tmin is the leaf, and all remaining leaves were determined, and total
daily minimum air temperature, and n is the days from planting to shoot and stem dry weight, plant height, and number of
each sampling (Baskerville and Emin, 1969; Viator et al., 2005). nodes were measured for all 28 cotton genotypes. Total
shoot dry weights were positively correlated with plant

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Table 2. Correlation coefficients between shoot dry weights
at three samplings in 2008 and four samplings in 2009 and
seed weight, plant height, cotyledon area, total leaf area, and
relative growth rate for 28 field grown cotton genotypes.
Shoot first Total
dry leaf leaf Relative
weight Seed Plant Cotyledon area area growth
Year at weight height area at S1† at S1† rate
g cm ————— cm2 ————— g g−1 d−1
2008 22 DAP 0.28 0.68 0.77 0.02 0.91
** *** *** NS‡ ***
32 DAP 0.39 0.56 0.30 −0.03 0.37 0.62
*** *** ** NS *** ***
52 DAP 0.13 0.43 0.12 −0.03 0.22 0.63
NS *** NS NS * ***
2009 17 DAP 0.30 0.29 0.82 0.82 0.91
*** ** *** *** ***
27 DAP 0.28 0.61 0.49 0.46 0.52 0.44
*** *** *** *** *** ***
35 DAP 0.19 0.74 0.35 0.45 0.46 0.51
NS *** *** *** *** ***
48 DAP 0.12 0.53 0.20 0.21 0.23 0.60
NS *** * * * ***
*Significant at the 0.05 probability level.
** Significant at the 0.01 probability level.
*** Significant at the 0.001 probability level.
†
Correlation between the total leaf area measured at the first sampling (S1) and the
shoot weight at each of the indicated samplings.
‡
NS, not significant.

height and total leaf area (Table 2; Fig. 2). To examine

the influence of early leaf area development on shoot dry
matter accumulation, total leaf area at the first sampling
as well as cotyledon area were correlated with shoot dry
weights at later samplings (Table 2). Total leaf area at the
first sampling was positively correlated with the shoot dry
weight at later samplings. Similarly, cotyledon area (mea-
sured at Sampling 1) and total shoot dry weight of plants
up to 48 DAP were also positively correlated. Despite the
progressively smaller correlations at later harvests, these
correlations indicate the importance of early leaf area
development for shoot dry matter accumulation.
As indicated by the positive correlations between cot-
yledon area and total leaf area (2008, r = 0.74, P < 0.0001;
2009, r = 0.83, P < 0.0001), genotypes with large coty-
ledons also had large total leaf area at the first samplings.
In addition, significant correlations were also observed
between cotyledon area and total leaf area at the second
sampling (2008, r = 0.34, P < 0.001; 2009, r = 0.58,
P < 0.0001). While the high r values at the first samplings
were anticipated, since cotyledons represented 49% (2-yr
average) of the total leaf area, at the second sampling coty-
ledons accounted for only 10% (2-yr average) of the total
leaf area and r values for the relationship of cotyledon area
with remaining leaf area were 0.30 (P < 0.01) and 0.53 (P
< 0.0001) in 2008 and 2009, respectively.
Figure 2. Relationships between shoot dry weight and total leaf
area of 28 genotypes harvested 32 days after planting (DAP) in
2008-A and 26 genotypes harvested 35 DAP in 2009-B under
field conditions.
Correlations of seed weight and cotyledon area (mea-
sured at 22 DAP in 2008 and 17 DAP in 2009) revealed
r values of 0.34 (2008, P < 0.001) and 0.50 (2009, P <
0.001), while those of seed weight and cotyledon weight
were 0.25 (2008, P < 0.05) and 0.42 (2009, P < 0.0001).
Further, seed weight was positively correlated with seed-
ling dry weight up to 32 DAP (Table 2). The positive asso-
ciation of seed weight with cotyledon size and seedling
dry weight was not surprising as a positive relationship of
seed size and seedling vigor is commonly observed and has
been shown for cotton (Snider et al., 2014) and a number
of other crops, including barley (Hordeum vulgare L.), bread
wheat, durum wheat (T. turgidum L.), triticale (Triticosecale
spp.), oat, rice, chickpea (Cicer arietinum L.), lupin (Lupinus
albus L.), and pea (Chen et al., 1986; Kamboozia, 1994;
López-Castañeda et al., 1996).
El-Sharkawy et al. (1965), examined growth charac-
teristics of 26 Gossypium species, including G. hirsutum and
G. barbadense L., and, after 8 to 9 wk of growth, found

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Table 3. Analysis of variance for seedling traits of 18 (2008) or 16 (2009) breeding lines and 10 commercial cotton varieties
grown under field conditions.
Leaf area Biomass
Total
Year Sampling Source DF Cot† First leaf All leaves Cot Frst leaf All leaves Stem shoot Height
2008 22 DAP Genotype 27 *** NS‡ *** *** * *** *** *** ***
Replication 3 NS NS NS NS NS NS NS NS NS
32 DAP Genotype 27 *** *** ** *** * *** ***
Replication 3 * NS NS NS NS NS *
52 DAP Genotype 27 * * * ***
Replication 3 NS NS NS NS
2009 17 DAP Genotype 25 *** *** *** *** *** *** *** NS
Replication 3 NS NS * * * * * NS
27 DAP Genotype 25 *** ** ** *** *** *** *** *** ***
Replication 3 NS NS *** NS NS NS NS NS ***
35 DAP Genotype 25 ** *** *** *** ***
Replication 3 *** *** *** *** ***
48 DAP Genotype 25 NS NS NS ***
Replication 3 ** *** *** ***
* Significant at the 0.05 probability level.
** Significant at the 0.01 probability level.
*** Significant at the 0.001 probability level.
†
Cot, cotyledon.
‡
NS, not significant.

significant differences in dry matter accumulation among
the species, and that the differences were positively asso-
ciated with differences in leaf area. Further, the authors
found that cotyledon size was positively related to dry
matter accumulation across the 26 Gossypium species.
Early leaf area development has previously been shown
to be important for early vigor in other species as well.
López-Castañeda et al. (1996) suggested that the width
of the first seedling leaf is an indirect estimate of early
vigor in barley, wheat, triticale, and oat. For wheat, Liang
and Richards (1999) found that the area of the first two
leaves was the trait most closely associated with seedling
vigor. Thus, in addition to cotyledon size, the area of the
first true leaf was measured in this study as it may play
an important role in early development of cotton. In the
2009 sampling, the size of the first true leaf was positively
associated with shoot biomass at all sampling dates (Table
2). However, no significant correlations were observed in
2008. Nonetheless, these data suggest that cotyledon size
and, at least in 1 yr, the size of the first true leaf, were
important drivers of early growth. This is consistent with
results from Kerby and Keeley (1987) who showed that
the removal of cotyledons up to 15 d after emergence sig-
nificantly reduced shoot dry weight determined at 53 d
after emergence. Further, in a comparison of cotton with
a visible true leaf at emergence and normal phenotype
seedlings, Ortiz and Bourland (1999) found that total dry
weight and leaf area were greater in plants with a visible
true leaf than those without a true leaf at emergence.
Genotypic Variation in Early Vigor Traits
Differences in environmental conditions between the 2
yr strongly influenced growth and development resulting
in highly significant year effects (P < 0.001) for all exam-
ined traits. However, significant genotypic variation was
observed in both years for all seedling biomass and leaf
area traits, except the area of the first true leaf in 2008 and
biomass at the last sampling in 2009 (Table 3). In 2009,
no genotypic differences were observed for leaf, stem, and
total shoot biomass 48 DAP (last sampling) even though
such differences were observed in 2008 at 52 DAP. Geno-
typic differences in plant height were observed at every
sampling in both years, except at 17 DAP in 2009 when
the average plant height was only 8.6 cm. As shown in
Tables 4 and 5, a considerable range in values among the
genotypes was observed for many traits at the first sam-
pling of each year. Genotypic differences for total shoot
dry matter were highly significant (through 35 DAP;
Table 3), with the most vigorous genotype accumulating
2.5-, 2.4-, and 2.3-fold more biomass than the least vig-
orous genotype, at the first, second, and third samplings,
respectively (averaged across years). Leaf area was mea-
sured at the first two samplings in 2008 and the first three
samplings in 2009. The genotypes differed significantly
in total leaf area at each of these samplings. At the first
and second samplings, a 2.6-fold larger total leaf area was
observed for the genotype with the maximum compared
with the genotype with the minimum leaf area (aver-
aged across both years; data not shown). While genotypes
differed significantly in both total leaf area and total leaf
biomass, specific leaf area (on a total leaf area basis) was

1734 www.crops.org crop science, vol. 55, july– august 2015

Table 4. Mean, standard deviation, minimum, and maximum values for shoot characteristics of 28 cotton genotypes grown
under field conditions in 2008.

Year Sampling Trait Mean  SD Min. Max.

2008 22 DAP Cotyledon area (cm2) 25.99  3.85 19.24 31.10
First true leaf area (cm2) 9.66  2.24 7.10 13.08
Total leaf area (cm2) 64.86  11.83 46.58 82.24
Cotyledon dry weight (g) 0.18  0.033 0.13 0.23
First true leaf dry weight (g) 0.059  0.015 0.044 0.081
Total leaf dry weight (g) 0.39  0.067 0.28 0.48
Stem dry weight (g) 0.17  0.029 0.13 0.22
Total shoot dry weight (g) 0.56  0.090 0.41 0.70
Plant height (cm) 10.27  1.26 8.65 12.68
Leaf area ratio (cm2 g−1) 116.66  8.79 106.68 125.39
Specific leaf area (cm2 g−1) 168.45  10.40 153.44 178.71
2008 32 DAP Cotyledon area (cm2) 27.66  4.79 20.14 34.58
Total leaf area (cm2) 315.70  56.78 232.7 377.01
Cotyledon dry weight (g) 0.25  0.051 0.18 0.33
Total leaf dry weight (g) 1.90  0.33 1.46 2.23
Stem dry weight (g) 1.15  0.24 0.87 1.51
Total shoot dry weight (g) 3.05  0.53 2.33 3.68
Plant height (cm) 24.79  2.58 21.35 28.63
Leaf area ratio (cm2 g−1) 103.90  9.56 94.66 122.31
Specific leaf area (cm2 g−1) 166.55  16.44 152.43 196.46
2008 52 DAP Total leaf dry weight (g) 6.91  1.18 5.40 8.54
Stem dry weight (g) 6.36  1.12 4.79 7.98
Total shoot dry weight (g) 14.00  2.37 10.60 17.27
Plant height (cm) 45.44  4.45 37.45 51.50
2008 Growth RGR† (~22–32 DAP) (g g−1 d−1) 0.17  0.020 0.15 0.19
RGR (~22–52 DAP) (g g−1 d−1) 0.11  0.0071 0.095 0.12
RGR (~32–52 DAP) (g g−1 d−1) 0.076  0.012 0.062 0.087
†
RGR, relative growth rate.

the same for all genotypes (data not shown). Cotyledon
area was strongly influenced by the genotype and, when
averaged across all measurements, a difference of 18.6 cm 2
was observed between the genotype with the greatest and
the genotype with the smallest cotyledons. Thus the dif-
ference between the extreme genotypes is more than the
average size of the cotyledons (15.8 cm 2 average across all
samplings) observed for the genotypes with the smallest
cotyledons. In contrast to cotyledon area, genotypes dif-
fered in area of the first true leaf only in 2009 and not in
2008. However, a genotype effect was observed for the
weight of the first true leaf in both years (Table 3).
Results presented in Table 2 show a significant posi-
tive association of shoot dry weight with RGR for the
time periods between the first and subsequent samplings.
While genotypes did not differ in RGR in 2008, a signifi-
cant genotype effect (P = 0.02) was observed for RGR in
2009 when calculated for the time interval between the
first and the last sampling. Analyses of the RGR for the
time intervals between each of the four samplings revealed
a highly significant genotype effect (P < 0.001) during
the period between 35 and 48 DAP only. During that
period, maximum RGR was 0.151 g g−1 d−1 and mini-
mum RGR was 0.113 g g−1 d−1. Mean RGRs calculated
across all genotypes for the time period between the first
and the last sampling averaged 0.107 and 0.149 g g−1 d−1 in
2008 and 2009, respectively.
Variation in early growth among the cotton geno-
types was expected and has previously been shown, albeit
based on fewer traits and genotypes. For instance, Snider
et al. (2014) observed differences in seedling vigor (seed-
ling fresh weight) among 11 cotton cultivars grown at
five locations. Ortiz and Bourland (1999) compared early
growth of four cotton genotypes under greenhouse con-
ditions and showed genotypic differences in leaf area and
dry weight at 10, 20, and 30 d after emergence. Simi-
larly, Cook and El-Zik (1992) compared seedlings from
six cotton genotypes under normal and drought stressed
field conditions and found that root dry weight, shoot dry
weight and root-to-shoot ratio, varied among genotypes
when sampled 28 DAP. Basal et al. (2005), also showed
significant genetic variation in root and shoot dry weight
in six genotypes harvested 44 d after emergence in non-
stressed and stressed conditions.
Findings for RGR are similar to those reported by
Fakorede and Ojo (1981) for maize. While they found some
variation in RGR among genotypes, differences in RGR
were only significant for three of the five sampling intervals

crop science, vol. 55, july– august 2015  www.crops.org 1735

Table 5. Mean, standard deviation, minimum, and maximum values for shoot characteristics of 26 cotton genotypes grown
under field conditions in 2009.

Year Sampling Trait Mean  SD Min Max

2009 17 DAP Cotyledon area (cm2) 22.10  2.80 16.41 27.47
First true leaf area (cm2) 16.66  4.73 7.79 24.09
Total leaf area (cm2) 38.75  6.78 24.19 48.31
Cotyledon dry weight (g) 0.11  0.016 0.086 0.14
First true leaf dry weight (g) 0.062  0.018 0.03 0.09
Total leaf dry weight (g) 0.18  0.031 0.12 0.21
Stem dry weight (g) 0.082  0.017 0.06 0.11
Total shoot dry weight (g) 0.26  0.045 0.18 0.32
Plant height (cm) 8.63  2.10 6.36 11.53
Leaf area ratio (cm2 g−1) 150.75  10.91 137.19 160.96
Specific leaf area (cm2 g−1) 221.49  16.30 204.61 233.49
2009 27 DAP Cotyledon area (cm2) 23.43  3.05 18.65 27.76
First true leaf area (cm2) 19.39  4.47 13.75 25.32
Total leaf area (cm2) 203.11  38.04 143.32 230.31
Cotyledon dry weight (g) 0.16  0.022 0.13 0.21
First true leaf dry weight (g) 0.12  0.032 0.077 0.18
Total leaf dry weight (g) 1.08  0.16 0.75 1.25
Stem dry weight (g) 0.65  0.12 0.43 0.80
Total shoot dry weight (g) 1.73  0.28 1.19 2.04
Plant height (cm) 20.64  2.50 17.06 24.27
Leaf area ratio (cm2 g−1) 117.61  14.05 107.06 127.84
Specific leaf area (cm2 g−1) 187.47  22.21 172.34 203.69
2009 35DAP Total leaf area (cm2) 515.78  105.84 364.54 648.21
Total leaf dry weight (g) 2.94  0.62 2.10 3.83
Stem dry weight (g) 2.09  0.46 1.30 2.63
Total shoot dry weight (g) 5.03  1.03 3.40 6.47
Plant height (cm) 30.03  3.92 23.8 36.45
Leaf area ratio (cm2 g−1) 102.88  9.51 92.27 119.46
Specific leaf area (cm2 g−1) 176.70  19.52 163.81 196.92
2009 48 DAP Total leaf dry weight (g) 13.59  2.34 11.23 16.57
Stem dry weight (g) 12.52  2.62 10.06 15.37
Total shoot dry weight (g) 26.84  4.90 22.44 32.32
Plant height (cm) 77.41  7.35 64.85 86.05
2009 Growth RGR† (~17–27 DAP) (g g−1 d−1) 0.19  0.017 0.17 0.21
RGR (~17–35 DAP) (g g−1 d−1) 0.13  0.012 0.12 0.14
RGR (~17–48 DAP) (g g−1 d−1) 0.15  0.0068 0.14 0.16
RGR (~27–35 DAP) (g g−1 d−1) 0.13  0.026 0.10 0.16
RGR (~27–48 DAP) (g g−1 d−1) 0.13  0.011 0.12 0.14
RGR (~35–45 DAP) (g g−1 d−1) 0.13  0.014 0.11 0.15
†
RGR, relative growth rate.

between 9 and 25 DAP. Interestingly, comparative analy-
ses of seedling characteristics contributing to variation in
the early vigor between barley, bread wheat, durum wheat,
triticale, and oat revealed similar RGRs for whole plant,
shoots, and roots for all species and cultivars even though
barley and triticale had greater total dry weight and leaf
area (López-Castañeda et al., 1995, 1996). For cotton, Ortiz
and Bourland (1999) did not find any differences in RGR
among four genotypes even though seedling leaf areas and
dry weights of the genotypes differed significantly when
sampled at 10, 20, and 30 d after emergence.
Even though, as indicated above, environmental con-
ditions and, consequently, planting dates and crop growth
differed significantly between the two growing seasons,
the absence of genotype-by-year interactions (except for
dry weight of Leaf 1 at the first sampling and plant height
at all three samplings) indicated that the performance of
the genotypes was consistent across years. This also can be
illustrated by the highly correlated total shoot dry matter
(r = 0.91, P < 0.001) and total leaf area (r = 0.52, P <
0.001) between 22 DAP in 2008 and 27 DAP in 2009. At
these sampling times, the two genotypes with the greatest
biomass accumulation in 2008 (Ark 0001-01-03 and Ark
0016-05-10) were also among the top three in 2009 (gen-
otype Ark 0016–05–10 was later released as Arkot 0016
[Bourland and Jones, 2011]). Similarly, the four genotypes

1736 www.crops.org crop science, vol. 55, july– august 2015

with the lowest biomass accumulation were the same in
2008 and 2009 (BCSX 0721 B2F, FM 1740 B2F [PVP
200800163], BCSX 0888LLB2, and ST 5288 B2F [PVP
201000066]). While the correlation coefficient for dry
matter accumulation at the first and the last sampling was
only 0.39 (P = 0.05) in 2008 and 0.42 (P = 0.05) in 2009,
genotype Ark 0001-01-03 remained the entry with the
greatest or second greatest accumulation of dry matter at
the last samplings, and FM1740 B2F and ST 5288 B2F
were among the entries with the lowest amount of dry
matter at the last samplings in 2008 and the two lowest
in 2009. As indicated above, shoot dry weight and total
leaf area were highly correlated at all samplings. Thus,
it was not surprising to find genotypes Ark 0001-01-03
and Arkot 0016 among those with the greatest leaf areas.
In addition to these two genotypes, Ark 0002-03-02 also
consistently ranked among the top four in leaf area. The
genotype BCSX 0721 B2F exhibited the smallest leaf area
among all examined genotypes at every sampling across
both years, and ST 5288 B2F also consistently ranked
among the three genotypes with the lowest leaf areas.
Significant variations in seedling traits among 28
cotton genotypes were observed in this study (Table 4,
5). As indicated by the lack of genotype-by-year interac-
tion and significant correlations between 2008 and 2009
data, the responses of the genotypes were consistent across
the 2 yr, suggesting that the examined traits are largely
influenced by the genotype and that genotype-by-envi-
ronment interactions may not play a large role. The large
ranges observed for the various traits (Table 4, 5) provide
diversity that may be useful for genetic improvement.
Importantly, further exploration of the cotton germ-
plasm beyond the relatively few genotypes examined in
this study will likely reveal more extreme phenotypes that
could be used for breeding purposes and analyses of the
genetics underlying seedling vigor traits of interest.
Early Vigor and Yield
High seed vigor should provide a high percentage of uni-
formly emerged, rapidly developing seedlings and result in
early season cotton stands with potential for high yields.
Lint yields in this study ranged from 960 to 1639 kg ha−1
(average, 1333 kg ha−1) in 2008 and 904 to 1364 kg ha−1
(average, 1214 kg ha−1) in 2009. Despite differences in
early vigor, no significant correlations between seedling
dry weight or seedling leaf area and lint yield were found
in either year. These results are in agreement with the con-
clusions from a review by TeKrony and Egli (1991) that
examined literature encompassing crops harvested at dif-
ferent developmental stages. They found that the effect of
seed vigor on yield depends on the harvested organ of a
plant and on harvest timing. Positive relationships between
seed vigor and yield are commonly observed for species
for which leaves or roots are the harvested portions (i.e.,
crop science, vol. 55, july– august 2015  www.crops.org 1737
Lactuca sativa L. and Daucus carota L.) or species harvested
during early reproductive growth (i.e., pea and Lycopersicon
esculentum L). In contrast, positive relationships between
seed vigor and yield of grain crops harvested at maturity
are not consistently found in situations where plant den-
sity did not influence yield. Nonetheless, seedling vigor is
important for lowering risks of stand establishment and for
providing a competitive advantage in situations such as low
stand density (Fuck et al., 1962), late planting (Khah et al.,
1989), insect pressure (Elliott et al., 2008), or weed infes-
tation (Kamboozia, 1994; Gill and Zerner, 2006) where
greater seedling vigor has been shown to increase yields.
Cultivar and Breeding Line Comparisons
To examine seedling vigor in commercial cultivars, 10
entries developed by private companies were selected for
evaluation and comparison with advanced breeding lines
from the University of Arkansas cotton breeding pro-
gram. Comparison of the commercial cultivars to the
breeding lines revealed that, as a group, early vigor of the
commercial cultivars was lower than that of the breeding
lines at all sampling dates in both years (Fig. 3). Signifi-
cantly lower areas of cotyledons and all leaves and lower
dry weights of cotyledons and all leaves and total shoot (all
P < 0.001) at the first sampling in both years as well as the
second and third sampling in 2009 were found (data not
shown). As indicated by the significant differences in total
shoot dry weight, the differences between the commer-
cial cultivars and the breeding lines persisted in 2008 and
2009. On average and across all sampling dates and both
years, the commercial cultivars accumulated 88% of the
shoot dry matter and 89% of the leaf area that was amassed
by the advanced breeding lines. At the early samplings,
four (2008) or five (2009) commercial cultivars exhibited
the lowest biomass accumulation while the commercial
cultivar with the greatest biomass accumulation (PHY
315 RF) ranked eighth (2008) or seventh in total biomass
accumulation among all the entries.
Overall, the commercial cultivars examined in
this study tended to have lower seedling vigor than the
advanced breeding lines from the University of Arkansas
breeding program. This provides possibilities for future
breeding by introducing early vigor traits into the high
yield commercial cultivars.
Seed Source Influences Early
Seedling Growth
Environmental conditions strongly influence plant growth
and yield, and conditions during seed development may
subsequently influence the growth of the seedlings emerg-
ing from these seeds. To evaluate the impact of different
production environments on early cotton growth, seeds
of 18 breeding lines grown in either Arkansas or Arizona
(seed source) were examined. Assessment of leaf area, leaf
Figure 3. Mean shoot dry weights of 18 (2008-A) or 16 (2009-B)
cotton breeding lines and 10 commercial cultivars grown under
field conditions. (Values are means ±SE. Within day after plant-
ing, bars with a different letter indicate a significant difference at
P <0.05.)
dry matter, stem dry matter, total shoot dry matter, and
plant height revealed that seed source influenced these
traits but that the effect could only be observed at the first
sampling (17 DAP) in 2009 and up to ~4 wk after plant-
ing in 2008 (Fig. 4). While no genotype-by-seed source
interactions were observed in 2008, interactions were
found at the first sampling in 2009 but were largely absent
at all other samplings, indicating that by about 3 wk after
planting, the response of the 18 genotypes was consis-
tent across both seed source environments. Thus, using
shoot dry matter and total leaf area as integrated measures
of seedling vigor, seedlings from Arizona-grown seeds
exhibited greater vigor than those from Arkansas-grown
seeds at early stages (when sampled at 22 DAP and 32
DAP in 2008 and 17 DAP in 2009), but the vigor advan-
tage was lost in later stages of development (Fig. 4). At the
first sampling in 2009—but not 2008—the cotyledon area
in developing seedlings from Arizona-grown seed was
1738 www.crops.org crop science, vol. 55, july– august 2015
Figure 4. Mean shoot dry weights of 18 (2008-A) or 16 (2009-B)
cotton breeding lines grown under field conditions over the course
of early development. Seeds for these plants were harvested from
cotton grown in Arizona or Arkansas. (Values are means ±SE.
Within day after planting, bars with a different letter indicate a sig-
nificant difference at P <0.05.)
significantly greater than that in seedlings from Arkan-
sas-grown seed (data not shown). Since the average seed
mass across genotypes was 97.3 and 91.9 mg seed−1 for
seed grown in Arizona and Arkansas, respectively, larger
cotyledons in Arizona-grown seed should be expected as
larger seeds generally accumulate more organic metabolic
reserves (lipids and proteins), which benefit initial growth.
For instance, Krieg and Carroll (1978) demonstrated that
cotton seed maturity and quantity of available substrate
(weight) determined radicle growth rates irrespective of
cultivar. Further, Perry (1978, 1980) showed that heavier
seeds had better emergence and growth potential than
lighter seeds. More recently, Snider et al. (2014) reported
positive relationships between cotton seedling vigor (seed-
ling fresh weight) and both total seed oil content and seed
size. In addition, a significant effect of seed source envi-
ronments on seedling weight has previously been shown
for barley (Pieta and Ellis, 1991).
CONCLUSIONS
Positive associations of seed weight with cotyledon size
and early seedling dry weight and no significant correla-
tions between seedling dry weight or seedling leaf area
and lint yield were found. The examined genotypes dif-
fered significantly for all seedling biomass traits, cotyledon
area, and total leaf area in both years. The consistent per-
formance of the genotypes across years indicates that main
seedling vigor traits are largely influenced by the genotype
and that genotype-by-environment interactions may not
play a major role for early vigor. The presence of signifi-
cant genotypic variation coupled with the broad ranges in
observed values within the various traits and greater early
vigor of breeding lines than commercial cultivars suggests
that seedling vigor of high-yielding commercial culti-
vars could be improved through breeding. Future efforts
should include screening of a large and more diverse
group of cotton genotypes, identify the mechanisms asso-
ciated with differences in seedling vigor, characterizing
the underlying genetics, and investigating the agronomic
implications of increased seedling vigor.
Acknowledgments
Funding for this project was provided by Cotton Incorporated
and the Cotton State Support Committee of Missouri.
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