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International journal of Rural Development, Environment and Health Research(IJREH) [Vol-2, Issue-2, Mar-Apr, 2018]
https://dx.doi.org/10.22161/ijreh.2.2.1 ISSN: 2456-8678
www.aipublications.com/ijreh Page | 1
Determination of Sugar Content in Lactuca
sativa L. Grown at Different Temperatures
Najia Shwerif 1*
, Kirsten Brandt2
, Stephen Wilcockson3
1*
Department of Plant Science, Gharian University, Libya.
2
Human Nutrition Research Centre, School of Agriculture, Food & Rural Development Newcastle University, Newcastle
Upon Tyne, NE1 7RU, UK.
3
Nafferton Ecological Farming Group, Newcastle University, Nafferton Farm, Stocksfield, NE43 7XD, UK
Abstract— Temperature has a large impact on the
growth and development of plants. The temperature
conditions that the plant is grown in may affect the
content of phytochemicals, which in turn affects the
quality of crops. This study investigates the effects of
different growth temperature regimes on sugar content of
two variety of lettuce grown in a controlled environment.
The results showed that, highly significant effects of the
temperature growth regimes on the lettuce varieties were
observed. There were higher levels of sugar in plants
grown at low temperature compared with high
temperature.
Keywords— Abiotic stress, Leafy vegetables, Sugar
content, Temperature.
I. INTRODUCTION
Temperature is an environmental factor with large effects
on growth, development, yield and quality of food crops
(Berry and Bjorkman, 1980; Kopsell, 2010). Global
climate and temperature changes will affect vegetable
productivity, nutritional value with consequences for diet
and health. Lettuce is an important worldwide, dietary,
leafy vegetable that is primarily cultivated and consumed
as a fresh product or in salad mixes. It is an important
source of phytonutrients, (Liu et al., 2007; Khoo et al.,
2011; Cruz et al., 2012).
Sugars are organic compounds in plants resulting from
photosynthesis, which play an important role in
respiration by providing the energy. Sugars are condensed
to store the energy in the form of starch, whilst the energy
is transported in sucrose form and sugars play a role in
cell wall structure, (Harborne, 1998). In addition to these
functions, sugars perform a regulatory role in numerous
mechanisms of plant growth and development (Rosa et
al., 2009). They aid control of plant metabolism and
different stress responses during the whole growth stages
from embryogenesis to senescence, involving a number of
sugar signals that are generated depending on the
surrounding environmental conditions, (Rolland et al.,
2006). The production and consumption of sugars are
performed by plants, whereas the environmental abiotic
stresses are one of the main factors that have an influence
on soluble sugars due to their sensitivity. However, the
soluble sugar levels increase in a plant with low
temperature conditions (Rosa et al., 2009). According to
molecular size the sugars are divided into three groups:
monosaccharaides such as glucose and fructose,
oligosaccharide and polysaccharides (Harborne, 1998).
The leafy vegetables including lettuce, fructose
contributed the highest concentration (about 55 %) of
total soluble sugars. In a lettuce plant sugar content
changes during the late stages of growth and development
as plants age and continues to increase. Furthermore,
sugar levels increase from winter to summer when grown
in greenhouses (Gent, 2012).
II. METHODS AND MATERIALS
2.1 Plant and growth condition
Different varieties of lettuce (Dixter and Exbury) were
grown in growth chambers, at different day temperatures:
12, 17, 22, 27 and 32℃ with a 4℃ day/night temperature
difference, an 11 hour photoperiod and with controlled
light and water supply. The light was constant 150 µmol
/m2/s. Each temperature treatment was applied at a
separate time and the plants were harvested after having
produced 10-12 leaves.
2.2 Sugar measurement
For sugars measurement of freeze dried lettuce samples,
1ml ultra –pure water was added to 100mg of samples,
vortex mixed for a few minutes then the concentrated
extracts were measured by hand with a refractometer
(Bellingham and Stanley Eclipse refractometer, 0-30
BRIX⁰). According to (Gaweda, 2007), the concentration
of sugars were proportionally distributed as glucose 2.9g ,
fructose 5.4g and sucrose 1.7g per 100ml. The standard
curve provide the conversion of BRIXº unit to sugar gram
per 100ml, therefore one BRIX corresponded to
1.1125g/100ml, Figure 1.
International journal of Rural Development, Environment and Health Research(IJREH) [Vol-2, Issue-2, Mar-Apr, 2018]
https://dx.doi.org/10.22161/ijreh.2.2.1 ISSN: 2456-8678
www.aipublications.com/ijreh Page | 2
Fig. 1: Standard curves for the sugar determination of
lettuce.
III. RESULTS
Temperature regimes had a highly significant effect on
shoot sugar contents of both lettuce varieties. There
highest content of sugar was at low temperature (12/8°C)
and it decreased as temperature increased (Figure 2).
Lettuce Dixter had higher sugar concentrations than
Lettuce Exbury, but the interaction between the lettuce
varieties and temperature was not significant. The non-
sugar components of varied between the varieties and
plant parts with a highly significant effect of temperature
Figure 2.
IV. DISCUSSION
Accumulation of sugar in shoot of Lettuce was highest at
a low temperature and decreases with increasing
temperatures. According to (Pietrini et al., 2002), at a low
temperature the respiration rates slow down and there is
less enzyme activity, which tends to reduce energy and
leads to a high accumulation of sugar. By contrast, at a
high temperature, increasing respiration (more energy
requirement) leads to a decrease in the sugar content with
a different response between the genotypes, as a result of
the associated active nitrogen metabolism process that
utilizes sugars, (Champigny, 1995). This offers an
explanation for current results whereby, the mechanism of
sugar production results in high levels in response to the
effect of low temperatures, leading to the inhibition of
nitrate accumulation associated with an increase in the
rate of nitrogen metabolism. Similar observations were
made in lettuce leaves by (Zhou et al., 2013) with the
same light intensity but a different light quality.
Fig. 2: Sugar and non-sugar content of Lettuce grown in different temperature.
International journal of Rural Development, Environment and Health Research(IJREH) [Vol-2, Issue-2, Mar-Apr, 2018]
https://dx.doi.org/10.22161/ijreh.2.2.1 ISSN: 2456-8678
www.aipublications.com/ijreh Page | 3
Additionally, the greater concentration of sugar is
associated with slow growth and development of plants,
due to the main effect of low temperatures on sugar
accumulation. This contrasts with higher temperature
regimes that lead to more rapid growth and less sugar.
Therefore the physiological process (the photosynthesis
and respiration alterations caused by temperature) is a
possible reason, since dark respiration slows down and
causes inhibition of carbohydrate consumption under low
temperature conditions (Khayat et al., 1988). Thus
carbohydrate levels are influenced by the maturation
extent of leaves and shoots (Jiao et al., 1989) in rose
plants. Although effects on sugar content were consistent
in the recent study, this was not the case for the content of
non-sugar materials (e.g. cellulose and protein) of lettuce
plant.
V. CONCLUSION
Clearly growth temperature regimes have an effect on the
physiological processes of plants and involve
components, which either respond to or defend against
extreme temperatures by means of large alterations in
phytochemical compounds.
REFERENCES
[1] Berry, J. and Bjorkman, O. (1980) 'Photosynthetin
Response and Adaption to Temperature in Higher-
Plants.', Annual Review of Plant Physiology and
Plant Molecular Biology, 31, pp. 491-543.
[2] Champigny, M. L. (1995) 'Integration of
photosynthetic carbon and nitrogen metabolism in
higher plants', Photosynthesis Research, 46(1-2), pp.
117-127.
[3] Cruz, R., Baptista, P., Cunha, S., Pereira, J. A. and
Casal, S. (2012) 'Carotenoids of Lettuce (Lactuca
sativa L.) Grown on Soil Enriched with Spent Coffee
Grounds', Molecules, 17(2), pp. 1535-1547.
[4] Gaweda, M. (2007) 'Changes in the contents of some
carbohydrates in vegetables cumulating lead', Polish
Journal of Environmental Studies, 16(1), pp. 57-62.
[5] Gent, M. P. N. (2012) 'Composition of hydroponic
lettuce: effect of time of day, plant size, and season',
Journal of the Science of Food and Agriculture,
92(3), pp. 542-550.
[6] Harborne, J. B. (ed.) (1998) Phytochemical Methods
A Guide to modern techiques of plant analysis. 3 edn.
London UK: Chapman & Hall.
[7] Jiao, J., Gilmour, M., Tsujita, M. J. and Grodzinski,
B. (1989) 'Photosynthesis and Carbon Partitioning in
Samantha roses', Canadian Journal of Plant Science,
69(2), pp. 577-584.
[8] Khayat, E., Zieslin, N., Mortensen, L. and Moe, R.
(1988) 'Effect of Alternating Temperature on Dark
Respiration and 14C Export in Rose Plants', Journal
of Plant Physiology, 133(2), pp. 199-202.
[9] Khoo, H.-E., Prasad, K. N., Kong, K.-W., Jiang, Y.
and Ismail, A. (2011) 'Carotenoids and Their
Isomers: Color Pigments in Fruits and Vegetables',
Molecules, 16, pp. 1710-1738.
[10]Kopsell, D. A. (2010) 'Improving Carotenoid
Phytochemical Concentration in Vegetable Crops.
[11]Liu, X., Ardo, S., Bunning, M., Parry, J., Zhou, K.,
Stushnoff, C., Stoniker, F., Yu, L. and Kendall, P.
(2007) 'Total phenolic content and DPPHd radical
scavenging activity of lettuce (Lactuca sativa L.)
grown in Colorado', LWT, 40, pp. 552-557.
[12]Pietrini, F., Iannelli, M. A. and Massacci, A. (2002)
'Anthocyanin accumulation in the illuminated surface
of maize leaves enhances protection from photo-
inhibitory risks at low temperature, without further
limitation to photosynthesis', Plant Cell and
Environment, 25(10), pp. 1251-1259.
[13]Rolland, F., Baena-Gonzalez, E. and Sheen, J. (2006)
'Sugar Sensing and Signaling in Plants: Conserved
and Novel Mechanisms', Annual Review of Plant
Biology 57, pp. 675- 709.
[14]Rosa, M., Prado, C., Podazza, G., Interdonato, R.,
Gonzalez, J. A., Hilal, M. and Prado, F. E. (2009)
'Soluble sugars--metabolism, sensing and abiotic
stress: a complex network in the life of plants', Plant
signaling & behavior, 4(5), pp. 388-93.
[15]Zhou, W., Liu, W. and Yang, Q. (2013) 'Reducing
Nitrate Content in Lettuce by Pre-Harvest Continuos
Light Delivered by Red and Blue Light-Emitting
Diodes.', Journal of Plant Nutrition, 36(3), pp. 481-
490.

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  • 1. International journal of Rural Development, Environment and Health Research(IJREH) [Vol-2, Issue-2, Mar-Apr, 2018] https://dx.doi.org/10.22161/ijreh.2.2.1 ISSN: 2456-8678 www.aipublications.com/ijreh Page | 1 Determination of Sugar Content in Lactuca sativa L. Grown at Different Temperatures Najia Shwerif 1* , Kirsten Brandt2 , Stephen Wilcockson3 1* Department of Plant Science, Gharian University, Libya. 2 Human Nutrition Research Centre, School of Agriculture, Food & Rural Development Newcastle University, Newcastle Upon Tyne, NE1 7RU, UK. 3 Nafferton Ecological Farming Group, Newcastle University, Nafferton Farm, Stocksfield, NE43 7XD, UK Abstract— Temperature has a large impact on the growth and development of plants. The temperature conditions that the plant is grown in may affect the content of phytochemicals, which in turn affects the quality of crops. This study investigates the effects of different growth temperature regimes on sugar content of two variety of lettuce grown in a controlled environment. The results showed that, highly significant effects of the temperature growth regimes on the lettuce varieties were observed. There were higher levels of sugar in plants grown at low temperature compared with high temperature. Keywords— Abiotic stress, Leafy vegetables, Sugar content, Temperature. I. INTRODUCTION Temperature is an environmental factor with large effects on growth, development, yield and quality of food crops (Berry and Bjorkman, 1980; Kopsell, 2010). Global climate and temperature changes will affect vegetable productivity, nutritional value with consequences for diet and health. Lettuce is an important worldwide, dietary, leafy vegetable that is primarily cultivated and consumed as a fresh product or in salad mixes. It is an important source of phytonutrients, (Liu et al., 2007; Khoo et al., 2011; Cruz et al., 2012). Sugars are organic compounds in plants resulting from photosynthesis, which play an important role in respiration by providing the energy. Sugars are condensed to store the energy in the form of starch, whilst the energy is transported in sucrose form and sugars play a role in cell wall structure, (Harborne, 1998). In addition to these functions, sugars perform a regulatory role in numerous mechanisms of plant growth and development (Rosa et al., 2009). They aid control of plant metabolism and different stress responses during the whole growth stages from embryogenesis to senescence, involving a number of sugar signals that are generated depending on the surrounding environmental conditions, (Rolland et al., 2006). The production and consumption of sugars are performed by plants, whereas the environmental abiotic stresses are one of the main factors that have an influence on soluble sugars due to their sensitivity. However, the soluble sugar levels increase in a plant with low temperature conditions (Rosa et al., 2009). According to molecular size the sugars are divided into three groups: monosaccharaides such as glucose and fructose, oligosaccharide and polysaccharides (Harborne, 1998). The leafy vegetables including lettuce, fructose contributed the highest concentration (about 55 %) of total soluble sugars. In a lettuce plant sugar content changes during the late stages of growth and development as plants age and continues to increase. Furthermore, sugar levels increase from winter to summer when grown in greenhouses (Gent, 2012). II. METHODS AND MATERIALS 2.1 Plant and growth condition Different varieties of lettuce (Dixter and Exbury) were grown in growth chambers, at different day temperatures: 12, 17, 22, 27 and 32℃ with a 4℃ day/night temperature difference, an 11 hour photoperiod and with controlled light and water supply. The light was constant 150 µmol /m2/s. Each temperature treatment was applied at a separate time and the plants were harvested after having produced 10-12 leaves. 2.2 Sugar measurement For sugars measurement of freeze dried lettuce samples, 1ml ultra –pure water was added to 100mg of samples, vortex mixed for a few minutes then the concentrated extracts were measured by hand with a refractometer (Bellingham and Stanley Eclipse refractometer, 0-30 BRIX⁰). According to (Gaweda, 2007), the concentration of sugars were proportionally distributed as glucose 2.9g , fructose 5.4g and sucrose 1.7g per 100ml. The standard curve provide the conversion of BRIXº unit to sugar gram per 100ml, therefore one BRIX corresponded to 1.1125g/100ml, Figure 1.
  • 2. International journal of Rural Development, Environment and Health Research(IJREH) [Vol-2, Issue-2, Mar-Apr, 2018] https://dx.doi.org/10.22161/ijreh.2.2.1 ISSN: 2456-8678 www.aipublications.com/ijreh Page | 2 Fig. 1: Standard curves for the sugar determination of lettuce. III. RESULTS Temperature regimes had a highly significant effect on shoot sugar contents of both lettuce varieties. There highest content of sugar was at low temperature (12/8°C) and it decreased as temperature increased (Figure 2). Lettuce Dixter had higher sugar concentrations than Lettuce Exbury, but the interaction between the lettuce varieties and temperature was not significant. The non- sugar components of varied between the varieties and plant parts with a highly significant effect of temperature Figure 2. IV. DISCUSSION Accumulation of sugar in shoot of Lettuce was highest at a low temperature and decreases with increasing temperatures. According to (Pietrini et al., 2002), at a low temperature the respiration rates slow down and there is less enzyme activity, which tends to reduce energy and leads to a high accumulation of sugar. By contrast, at a high temperature, increasing respiration (more energy requirement) leads to a decrease in the sugar content with a different response between the genotypes, as a result of the associated active nitrogen metabolism process that utilizes sugars, (Champigny, 1995). This offers an explanation for current results whereby, the mechanism of sugar production results in high levels in response to the effect of low temperatures, leading to the inhibition of nitrate accumulation associated with an increase in the rate of nitrogen metabolism. Similar observations were made in lettuce leaves by (Zhou et al., 2013) with the same light intensity but a different light quality. Fig. 2: Sugar and non-sugar content of Lettuce grown in different temperature.
  • 3. International journal of Rural Development, Environment and Health Research(IJREH) [Vol-2, Issue-2, Mar-Apr, 2018] https://dx.doi.org/10.22161/ijreh.2.2.1 ISSN: 2456-8678 www.aipublications.com/ijreh Page | 3 Additionally, the greater concentration of sugar is associated with slow growth and development of plants, due to the main effect of low temperatures on sugar accumulation. This contrasts with higher temperature regimes that lead to more rapid growth and less sugar. Therefore the physiological process (the photosynthesis and respiration alterations caused by temperature) is a possible reason, since dark respiration slows down and causes inhibition of carbohydrate consumption under low temperature conditions (Khayat et al., 1988). Thus carbohydrate levels are influenced by the maturation extent of leaves and shoots (Jiao et al., 1989) in rose plants. Although effects on sugar content were consistent in the recent study, this was not the case for the content of non-sugar materials (e.g. cellulose and protein) of lettuce plant. V. CONCLUSION Clearly growth temperature regimes have an effect on the physiological processes of plants and involve components, which either respond to or defend against extreme temperatures by means of large alterations in phytochemical compounds. REFERENCES [1] Berry, J. and Bjorkman, O. (1980) 'Photosynthetin Response and Adaption to Temperature in Higher- Plants.', Annual Review of Plant Physiology and Plant Molecular Biology, 31, pp. 491-543. [2] Champigny, M. L. (1995) 'Integration of photosynthetic carbon and nitrogen metabolism in higher plants', Photosynthesis Research, 46(1-2), pp. 117-127. [3] Cruz, R., Baptista, P., Cunha, S., Pereira, J. A. and Casal, S. (2012) 'Carotenoids of Lettuce (Lactuca sativa L.) Grown on Soil Enriched with Spent Coffee Grounds', Molecules, 17(2), pp. 1535-1547. [4] Gaweda, M. (2007) 'Changes in the contents of some carbohydrates in vegetables cumulating lead', Polish Journal of Environmental Studies, 16(1), pp. 57-62. [5] Gent, M. P. N. (2012) 'Composition of hydroponic lettuce: effect of time of day, plant size, and season', Journal of the Science of Food and Agriculture, 92(3), pp. 542-550. [6] Harborne, J. B. (ed.) (1998) Phytochemical Methods A Guide to modern techiques of plant analysis. 3 edn. London UK: Chapman & Hall. [7] Jiao, J., Gilmour, M., Tsujita, M. J. and Grodzinski, B. (1989) 'Photosynthesis and Carbon Partitioning in Samantha roses', Canadian Journal of Plant Science, 69(2), pp. 577-584. [8] Khayat, E., Zieslin, N., Mortensen, L. and Moe, R. (1988) 'Effect of Alternating Temperature on Dark Respiration and 14C Export in Rose Plants', Journal of Plant Physiology, 133(2), pp. 199-202. [9] Khoo, H.-E., Prasad, K. N., Kong, K.-W., Jiang, Y. and Ismail, A. (2011) 'Carotenoids and Their Isomers: Color Pigments in Fruits and Vegetables', Molecules, 16, pp. 1710-1738. [10]Kopsell, D. A. (2010) 'Improving Carotenoid Phytochemical Concentration in Vegetable Crops. [11]Liu, X., Ardo, S., Bunning, M., Parry, J., Zhou, K., Stushnoff, C., Stoniker, F., Yu, L. and Kendall, P. (2007) 'Total phenolic content and DPPHd radical scavenging activity of lettuce (Lactuca sativa L.) grown in Colorado', LWT, 40, pp. 552-557. [12]Pietrini, F., Iannelli, M. A. and Massacci, A. (2002) 'Anthocyanin accumulation in the illuminated surface of maize leaves enhances protection from photo- inhibitory risks at low temperature, without further limitation to photosynthesis', Plant Cell and Environment, 25(10), pp. 1251-1259. [13]Rolland, F., Baena-Gonzalez, E. and Sheen, J. (2006) 'Sugar Sensing and Signaling in Plants: Conserved and Novel Mechanisms', Annual Review of Plant Biology 57, pp. 675- 709. [14]Rosa, M., Prado, C., Podazza, G., Interdonato, R., Gonzalez, J. A., Hilal, M. and Prado, F. E. (2009) 'Soluble sugars--metabolism, sensing and abiotic stress: a complex network in the life of plants', Plant signaling & behavior, 4(5), pp. 388-93. [15]Zhou, W., Liu, W. and Yang, Q. (2013) 'Reducing Nitrate Content in Lettuce by Pre-Harvest Continuos Light Delivered by Red and Blue Light-Emitting Diodes.', Journal of Plant Nutrition, 36(3), pp. 481- 490.