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Journal of Environmental Biology
July, 2007
Inter-relationship between growth analysis and carbohydrate
contents of sweet sorghum cultivars and lines
A. Almodares*, R. Taheri and S. Adeli
*aalm odar es@yah oo.com
Department of Biology, College of Science, University of Isfahan-81846-73441, Isfahan, Iran
(Received: January 06, 2006; Revised received: October 18, 2006; Accepted: November 11, 2006)
Abstract: The carbohydrate contents of sweet sorghum (Sorghum bicolor L. Moench) is an important industrial factor for crystal sugar or bioethanol
production. In this study the relationship between growth analysis and carbohydrate contents were studied to recognize the best growth stages for sweet
sorghum harvesting. Five sweet sorghum cultivars and four sweet sorghum lines were evaluated for leaf area index (LAI), net assimilation rate (NAR), relative
growth rate (RGR) and stem crop growth rate (CGR) in relation to sucrose content, invert sugars and total sugar at booting, soft-dough, hard dough and post
grain maturity. Except at post grain maturity stage, the correlations among LAI, NAR and RGR for sucrose content and total sugar were positive and for invert
sugars were negative. The relationship between invert sugars including glucose, fructose, maltose and xylose at hard dough stage regarding LAI, NAR and
GRG were negative. As plant grows LAI, NAR and RGR increases which consequently increases sucroses content and decrease invert sugar.
Key words: Sweet sorghum, LAI, NAR, RGR, CGR, Sucrose, Invert sugar
PDF file of full length paper is available with author
Introduction
Sorghum (Sorghum bicolor L. Moench) is one of the 5
major cultivated species in the world. It can outproduce most other
cereals under marginal environmental conditions, especially dry
and hot climatic conditions which prevails the most parts of the
country (Almodares and Darany, 2006). Sorghum tolerates drought
relatively well, and it responds to adequate fertility and soil moisture
with faster growth (Wayne and Frederiksen, 2005). Many types of
sorghum are suitable for grain and forage production (Showemimo
et al., 2002) , as well as, alternative uses, such as energy, pulp for
paper, food products and sugar or ethanol products (Gnansounoua
et al., 2005 and Dolciotti et al., 1998). Three valuable growth
analysis functions are relative growth rate (RGR), crop growth rate
(CGR) and net assimilation rate (NAR) (Borras’s et al., 2007). Leaf
area index (LAI) is a dimensionless variable and was first defined
as the total one-sided area of photosynthetic tissue per unit ground
surface area (Inge et al., 2004). LAI is the component of crop
growth analysis that accounts for the ability of the crop to capture
light energy and is critical to understand the function of many crop
management practices. Leaf area index can have importance in
many areas of agronomy and crop production through its influence:
light interception, crop growth, weed control, crop weed competition,
crop water use, and soil erosion (Welles, 1990 and Sonnentag et
al., 2007). NAR measures the mean photosynthetic efficiency of
leaves in a crop community. The integration of weight and leaf area
measurement over time provides value that is highly useful for
studying the growth of crops (Shipley, 2006). According to Patterson
(1982), relative growth rate (RGR), net assimilation rate (NAR),
and leaf area ratio (LAR) are good measures of solar radiation
capture during growth with NAR and LAR for an individual plant
and LAI for population helping to explain differences in RGR.
Samba et al. (2003) found that interception of PAR is closely
followed by LAI. Reduced NAR interception causes reduction of
the RGR, NAR and LAR. Mansab et al. (2003) reported that for
maximum crop growth, enough leaves must be present in the
canopy to intercept most of the incident NAR. Therefore, growth is
often expressed on a leaf-area basis. According to Tsuni and
Fujise (1965), there is a linear relationship between leaf-area
and net assimilation. Growth analysis of sweet sorghum in relation
to sucrose has not been reported so the purpose of this experiment
was to study the relationship between growth analysis and
carbohydrate content of sweet sorghum cultivars and lines.
Materials and Methods
Field experiment was conducted at the University of Isfahan
Experimental Station. Five sweet sorghum cultivars (Soave, Rio,
Vespa, Turno and M81-E) and four sweet sorghum lines (IS-6962,
IS-18154, IS-4546 and IS-16054) were assessed in a complete
block design with four replications. Plots consisted of 8 rows, 5 m
long and 0.80 cm apart. Plots received 300 kg/ha of diammonium
phosphate and 100 kg/ha of urea disced into the soil before planting.
Plots were side-dressed with 100 kg/ha of urea subsurface banded
30 days after planting. Three meters from two centre rows were
harvested when the plants reached at booting, flowering, soft-dough,
hard-dough and two weeks after seed maturity. The fresh stalk,
after removing the panicles and leaves, were weighed and crushed
in a sugarcane crusher to extract the juice. After filtration through a
sieve to remove chaff, etc., reducing sugars were measured
according to Lane-Eynon (1970). The soluble solids (brix) and
sucrose (pol%) were measured according to Varma (1988). HPLC
Journal of Environmental Biology July 2007, 28(3) 527-531 (2007)
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Journal of Environmental Biology
July, 2007
528 Almodares et al.
was used for determination of glucose, fructose, maltose, xylose,
mannose, galactose and arabinose. The HPLC separation was
accomplished by using an aminopropyl column (4.6 x 250 mm) with
a mobile phase of 80% acetonitrile/ 20% water (Biermann and
McGinnis, 1989). Quantitation was accomplished by using three
point calibration curves. Growth analysis were performed by
determining mean LAI, NAR (g/m
2
day) and RGR(g/g day)
according to methods outlined by Hunt (1990) . Parameter results
are presented graphically with best-fit polynomial equations plotted
against growth degree days (GDD), calculated from emergence
using a base temperature of 10 degree centigrade. Leaf area was
determined using leaf area meter, then the samples were dried at
65 degree centigrade for 48 hr and then their dry weights were
determined.
Results and Discussion
To study th e correl ati o ns be tween growt h and
carbohydrates, growth indexes such as LAI, NAR, RGR and stem
CGR at different growth stages were used. The relationships among
LAI, for sucrose, invert sugar, and total sugar of sweet sorghum
cultivars and lines at different growth stages are shown in Table 1.
This correlation became significant after booting. As number of leaves
increase, the amount of photosyntate such as sucrose increase.
The leaves are the primary organ for light interception and
photosynthesis in crop plants. According to Gardner et al. (1994),
as the leaf area develops radiation interception by leaves increase.
Sweet sorghum leaves are staying green after flowering which
increase photosynthesis. Bulk of translocated substances, other
than water, is the result of photosynthesis and 90% of the total solid
in the phloem consists of carbohydrates, mostly. The predominant
sugar, translocated in the phloem of most crop species is sucrose.
The results show that at all growth stages, invert sugars are
negatively correlated to LAI except at hard-dought post grain maturity
stage (Table 4). As plants grow, there are more leaves which
through photosynthesis, produce more sucrose and less invert
sugar. The correlation among LAI, NAR and RGR with glucose,
fructose, maltose and xylose at hard dough stage is shown in Table
5. The correlation among LAI and all the above mentioned sugars
are negative which means that as a number of leaves increases,
the amount of all the above mentioned sugars decreases. Table 2
shows significant (p<0.01) relationship between RGR and sugar
content at soft dough. The relation between NAR and sugar content
is presented in Table 3. Table 4 shows the relationships between
stem CGR and the above sugars at booting and soft dough stage.
In this study, CGR is presented on a 10 degree-day basis. At the
end of the season, CGR will decrease. CGR in the beginning of the
season increases because of the increase in leaf size and stem
weight. According to Davidson and Campbell (1984) reduction of
CGR at the end of the growing season is due to the reduction of
photosynthesis rate and leaves abscission. In this study stem CGR
at the end of growing season was not reduced due to the stay
green conditioned leaves of sweet sorghum cultivars and lines.
The relationships among LAI, NAR, RGR and stem CGR for sucrose
and total sugar were positive and for invert sugar were negative
Table - 4: Coefficient of correlation (r) between stem CGR and sugar
contents of sweet sorghum cultivars and lines at booting and flowering- soft
dough stages
Carbohydrates Booting Soft dough
Sucrose 0.98** 0.97**
Invert sugar -0.85 -0.98**
Total sugar 0.93 0.97**
* significant at 0.05, ** significant at 0.01
Table - 5: Coefficient of correlation (r) between LAI, NAR, RGR and invert
sugars of sweet sorghum cultivars and lines at soft dough- hard dough
stage
Carbohydrates LAI NAR RGR
Glucose -0.99** -0.97** -0.85
Fructose -0.95** -0.96** -0.75
Maltose -0.87 -0.86 -0.65
Xylose -0.98** -0.93* -0.82
* significant at 0.05, ** significant at 0.01
Table - 3: Coefficient of correlation (r) between NAR and sugar contents of
sweet sorghum cultivars and lines at different growth stages
Carbohydrates NAR
Booting Soft Hard Post grain
dough dough maturity
Sucrose 0.95 0.94* 0.96** 0.95**
Invert sugar -0.64 -0.92* -0.89 0.83
Total sugar 0.90 0.98** 0.97** 0.89
* significant at 0.05 ** significant at 0.01
Table - 1: Coefficient of correlation (r) between LAI and sugar contents of
sweet sorghum cultivars and lines at different growth stages
Carbohydrates LAI
Booting Soft Hard Post grain
dough dough maturity
Sucrose 0.93 0.94* 0.98** 0.99**
Invert sugar -0.84 -0.91* -0.93* 0.95**
Total sugar 0.85 0.95** 0.98** 93.3*
* significant at 0.05, ** significant at 0.01
Table - 2: Coefficient of correlation (r) between RGR and sugar contents of
sweet sorghum cultivars and lines at different growth stages
Carbohydrates RGR
Booting Soft Hard Post grain
dough dough maturity
Sucrose 0.65 0.95** 0.83 0.41
Invert sugar -0.65 -0.88 -0.74 0.21
Total sugar 0.59 0.84 0.81 0.69
* significant at 0.05, ** significant at 0.01
Journal of Environmental Biology
July, 2007
529Growth analysis and carbohydrate contents of sweet sorghum
Fig 1: Correlation between LAI and sucrose content at physiological maturity
Fig 2: Correlation between RGR and sucrose content at physiological maturity
except during post grain maturity. During this stage, the amount of
invert sugars was small. The relationships indicates that as the plant
grows the amount of sucrose and total sugar increase while the
amount of invert sugar decreases. Almodares et al. (1994) found
that at all growth stages, total sugar and sucrose were higher than
other carbohydrates. The relationships among LAI, NAR, RGR
and invert sugars (glucose, fructose, maltose and xylose) are shown
in Table 5. The coefficient of correlation for all these carbohydrates
are negative which indicate that the plant grows LAI, NAR, and
RGR also increase and the amount of these carbohydrates
decreases. The correlations among LAI, RGR, NAR and CGR
with sucrose for sweet sorghum cultivars and lines at hard dough
stage are shown in Fig. 1-5 respectively. The relationships have
linear and positive responses which indicate that as the plant grows,
there are more leaves, more photosynthesis (NAR), and more
growth (RGR and CGR) which increase sucrose content in the
stem. The results showed as sweet sorghum cultivars and lines
grow, LAI, NAR and RGR will be increased. As a result, the amount
of sucrose increased while the amount of invert sugar (glucose,
fructose, maltose and xylose) decreased. The results indicate that
Sucrose (g/kg F.W.)
Sucrose (g/kg F.W.)
LAI
Sucrose = 15 (LAI) + 0.38
Sucrose = 2837.56 (RGR) + 15.52
RGR (g/g) 10 GDD
Journal of Environmental Biology
July, 2007
530
Fig 3: Correlation between NAR and sucrose content at physiological maturity
Fig 4: Correlation between stem CGR and sucrose content at physiological maturity
the time of harvesting sweet sorghum, cultivars and lines is dependent
on the purpose of planting. If the purpose of planting is to produce
crystal sugar, it is suggested to harvest sweet sorghum cultivars
and lines at hard dough- post maturity stage when plants had the
highest sucrose and lowest invert sugars, since invert sugars
interfere crystallization. Harvesting sweet sorghum cultivars and
lines at other growth stages are more suitable for liquid sugar
production. It should be mentioned that as plants grow, biomass
increases and plants at early growth stage had lower biomass than
other growth stages.
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Journal of Environmental Biology
July, 2007
531
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Growth analysis and carbohydrate contents of sweet sorghum