Journal of Agricultural Science; Vol. 7, No.

5; 2015
ISSN 1916-9752 E-ISSN 1916-9760
Published by Canadian Center of Science and Education

Effects of Drought Stress on Seed Germination and Seedling Growth of

Different Maize Varieties
Mingli Liu1, Meng Li2, Kaichang Liu2 & Na Sui1
1
Key Laboratory of Plant Stress Research, College of Life Science, Shandong Normal University, Jinan, P. R.
China
2
Shandong Academy of Agricultural Sciences, Jinan, P. R. China
Correspondence: Kaichang Liu, Shandong Academy of Agricultural Sciences, Jinan 250100, P. R. China. E-mail:
liukc@saas.ac.cn
Na Sui, Key Laboratory of Plant Stress Research, College of Life Science, Shandong Normal University, Jinan
250014, P. R. China. E-mail: suina800101@163.com

Received: March 4, 2015 Accepted: March 31, 2015 Online Published: April 15, 2015
doi:10.5539/jas.v7n5p231 URL: http://dx.doi.org/10.5539/jas.v7n5p231

Abstract
Maize is susceptible to drought stress, especially during germination. The experiment of seed germination and
seedling growth of two maize varieties under different concentration of mannitol solution was carried out. Four
mannitol concentrations were set: 0, 200, 400 and 600 mM, and the corresponding solution osmotic potential was:
0, -0.49, -0.99, and -1.49 Mpa. Results indicated that Zhengdan958 germinated more quickly and in greater
numbers than Liansheng15. The germination rate, germination potential, shoot and root fresh weight, shoot and
root length of two maize varieties both decreased under drought stress, but they decreased more in Liansheng15
than in Zhengdan958. However, the germination and vigor index of Zhengdan958 were obviously higher than
that of Liansheng15 under the stress of 200, 400 and 600 mM mannitol. These results showed that the drought
resistances of Zhengdan958 was higher than that of Liansheng15 by having more powerful root system and
higher osmotic adjustment ability.
Keywords: maize, mannitol stress, germination, drought resistance
1. Introduction
Abiotic stresses are governed by various factors, the most prominent is the elevated temperatures and scarcity of
water (Pena & Hughes, 2007; Narusaka et al., 2003). As drought is one of the most important environmental
factors affecting plant growth, development and crop yield, considerable progress has been made in
understanding the changes in physiological processes caused by drought stress. It has been shown that the
physiological responses of plants to drought stress are extremely complex and vary with plant species as well as
with the degree and time of the exposure to drought (Evans et al., 1990, 1991; King, 2011). Differences in
resistance to drought are known to exist within genotypes of plant species and were found in many studies, e.g.
in maize (Martinielio & Lorenzoni, 1985; Lorens et al., 1987; Grzesiak, 1990; Grzesiak et al., 2012), wheat
(Winter et al., 1988; Reynolds et al., 1998; Paknejad et al., 2007), rape seed (Richards & Thurling, 1978), oat
(Larsson & Go’rny, 1988), coconut (Gomez et al., 2008) and triticale (Royo et al., 2000; Grzesiak et al., 2012).
Unusual dry weather conditions adversely affect germination and seedling growth rates thus enhancing cell
elongation sensitivity to damages induced under stressed conditions (Taylor et al., 1982; Delachiave & Pinho,
2003; Hamayun et al., 2010a). Deleterious effects of water stress have been reported in different crops such as
tomato (Ragab et al., 2007), soybean (Sakthivelu et al., 2008; Hamayun et al., 2010b), corn (Khodarahmpour,
2011) and citrus (Ben-Hayyim, 1987).
Maize is one of the important crops, which serves as food and corn oil for human consumption, feed for
livestock and poultry and raw material for agro-based industries. Expanding populations with greater food and
energy needs are increasing demand for greater global maize production. Unfortunately, environmental
limitations such as temperature and drought continue to restrain maize production levels as they have in earlier
decades and in many areas, which is predicted to worsen with changing climates (Battisti & Naylor, 2009; Lobell
et al., 2011; Mishra & Cherkauer, 2010). The increasing frequency of dry periods in many regions of the world

231
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

result in the consecutive occurrence of drought on cultivated land (Hu & Schmidhalter, 2005). In China, maize
planted in arid and semi arid area. At present, China’s annual maize planting area is around 24 million hm2, of
which nearly 2/3 area is dry-land maize. Drought has become an important limiting factors of agricultural
production in the world, especially in arid agricultural production. As dry-land crops, corn not only need more
water, but also are more sensitive to water stress. Drought is the major cause of our country and around the world
maize production low and unstable, generally can be reduced by 20%-30%. Therefore, the breeding of new
maize varieties with strong drought resistance is one of effective ways for high and stable yield. However, the
accurate identification of drought resistance of maize varieties is the basis for breeding drought resistance maize
varieties.
In recent years, many scholars have studied the drought mechanism of maize, identification methods and
indicators of drought resistance. Wang et al. have studied the germination characters of maize varieties with a
hypertonic solution PEG-6000 simulated drought stress. Xu et al. have studied the germination potential,
germination rate and utilization efficiency of storage material, amylase activity. Hou et al. have studied the
drought resistance in the germination period of maize by hypertonic solution germination, TTC quantitative
measurement, seed water absorption detection method with mannitol solution simulated water stress. These
methods and indicators reflected the drought resistance of maize varieties from different angles and extent, also
see scattered reports on bud seedling stage of maize.
Mannitol is a kind of macromolecule osmotic regulation substances that can’t through the cell wall, small toxicity
to cells, an ideal material to simulate the soil drought. This experiment with mannitol hypertonic solution
simulated water stress and studied the difference of drought resistance on germination and seedling growth of
two maize varieties to filter out strong drought resistance varieties, in order to provide theoretical guidance for
drought resistance breeding and promotion of maize varieties.
2. Materials and Methods
2.1 Plant Culture and Treatments
Maize seeds were provided by the shandong academy of agricultural sciences. Dry seeds were stored in a
refrigerator at −4 °C before being used.
Selecting the grain full, same size, no worm holes maize seeds, 70% ethanol solution disinfected for 2 min and
then rinsed with running water. Each variety seeds germinated in different germination boxes which containing 20
seeds. Each variety sets four concentration: 0, 200, 400 and 600 mM mannitol, three times repeated and the
corresponding solution osmotic potential were: 0, -0.49, -0.99, and -1.49 Mpa. New mannitol solution of the same
amount and normal water were replaced every day. The seeds germinated in a glasshouse in the light condition of
14 h light/10 h dark. The temperature in the glasshouse was 25 °C during the day and 20 °C at night.
2.2 Germination Rate (%)
Observing germination situation of maize seeds every day and the germination rate of seeds was measured by the
national seed inspection standards. Computing the final germination rate after germination for seven days.
Germination rate was calculated as follows: Germination rate = (Germinated seed number/Test seed number) ×
100%
2.3 Germination Potential
Germination potential is the index to evaluate the seed germination rate and germination uniformity. Germination
potential was calculated as follows: Germination potential = (Germinated seed number at germination peak/Test
seed number) × 100%
2.4 Germination and Vigor Index of Drought Resistant
Germination and vigor index of drought resistant were calculated as follows: Seed germination index of drought
resistance = seed promptness index under water stress (PIS)/controlled seed promptness index (PIC); Promptness
index PI = nd2 (1.00) + nd4 (0.75) + nd6 (0.50) + nd8 (0.25); ndx = number of germinated seeds by the xth day of
measurement (Grzesiak et al., 2013).
Seed vigor index of drought resistance = Seed vigor index under water stress (VIS)/Controlled seed vigor index
(VIC); Vigor index VI = PI × Sx; Sx is the average length of bud for seventh days.
2.5 Shoot and Root Fresh Weight and Root-Shoot Ratio
After germination for seven days, seedlings grew consistently were chosen to be cut the root and shoot with a
shear and weighed respectively. Root-shoot ratio was calculated as follows: Root-shoot ratio = Root fresh

232
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

weight/Shoot fresh weight.

2.6 Shoot and Root Length
After germination of maize seeds for seven days, the shoot and root length were measured with a ruler.
2.7 Drought Resistance Coefficient
Drought resistance coefficient can reflect the sensitivity of plants to drought and it was calculated as follows:
Drought resistance coefficient = (Index value under drought stress/Index value under normal moisture) × 100%
3. Results
3.1 Germination Rate during Drought Stress
The germination of Zhengdan958 was better than Liansheng15 under different mannitol treatment (Figure 1).
The germination rate of two maize varieties declined obviously under drought stress. Under 200 mM mannitol
treatment, germination percentage of Zhengdan958 and Liansheng15 decreased 1.75% and 13.46%, respectively.
Under 400 mM mannitol treatment, germination percentage of Zhengdan958 and Liansheng15 decreased 28.07
% and 50.00%, respectively. Under 600 mM mannitol treatment, germination percentage of Zhengdan958 and
Liansheng15 decreased 50.88% and 75.00%, respectively (Figure 2).

Zhengdan958

Liansheng15

Figure 1. Photos of germination of Zhengdan958 and Liansheng15 under different concentrations of mannitol

Figure 2. Germination rate of different maize varieties in response to different concentrations of mannitol. Each
point represents the means ± SD of five measurements on each of five plants (n = 5)

3.2 Germination Potential during Drought Stress

The germination potential of two maize varieties declined significantly under mannitol stress and there was
significant difference between different varieties. Under 200 mM mannitol treatment, there was nearly no
decrease in germination potential of Zhengdan958, but it decreased 34.62% in Liansheng15. Under the stress of
600 mM mannitol, the germination potential of Zhengdan958 decreased 67.92% and the germination potential of
Liansheng15 was 0 (Figure 3).

233
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

Figure 3. Germination potential of different maize varieties in response to different concentrations of mannitol.
Values are means ±SD of five measurements for each of five plants (n = 5)

3.3 Germination and Vigor Index of Drought Resistance

The germination index (PIS/PIC) and vigor index (VIS/VIC) of drought resistance of the two maize varieties
both decreased with the increase of mannitol concentration. The PIS/PIC of Zhengdan958 was higher than
Liansheng15 under the stress of 200, 400 and 600 mM mannitol. The VIS/VIC of Zhengdan958 was also higher
than that of Liansheng15 under 200 and 400 mM mannitol and there was no obvious difference under the stress
of 600 mM mannitol (Figures 4 and 5).

Figure 4. Germination index of drought resistance of different maize varieties in response to different
concentrations of mannitol. Data are means of 5 replicates (n = 5) ± SD

234
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

Figure 5. Vigor index of drought resistance of different maize varieties in response to different concentrations of
mannitol. Each point represents the means ± SD of five measurements on each of five plants (n = 5)

3.4 Shoot and Root Fresh Weight and Root-Shoot Ratio during Drought Stress
Shoot and root fresh weight of two maize varieties under drought stress decreased obviously with the increase of
mannitol concentration. Under 200 mM mannitol treatment, shoot fresh weight of Zhengdan958 and
Liansheng15 decreased 71.13% and 79.51%, respectively (Figure 6). Under 400 mM mannitol treatment, shoot
fresh weight of Zhengdan958 and Liansheng15 decreased 90.84% and 95.55%, respectively. Under 600 mM
mannitol treatment, shoot fresh weight of Zhengdan958 and Liansheng15 decreased 95.49% and 98.36%,
respectively. Root fresh weight of Zhengdan958 and Liansheng15 under 200, 400 and 600 mM mannitol
treatment decreased 43.73% and 46.30%, 70.28% and 87.27%, 85.87% and 96.00%, respectively (Figure 7).
The root-shoot ratio of Zhengdan958 increased during mannitol treatment. It increased before 400 mM mannitol
treatment and then decreased in Liansheng15 (Figure 8). Root-shoot ratio of Zhengdan958 increased from 0.82
under the control condition to 3.11 under 600 mM mannitol treatment, and the ratio of Liansheng15 increased
from 1.04 under the control condition to 2.15 under 600 mM mannitol treatment.

Figure 6. Shoot fresh weight of different maize varieties in response to different concentrations of mannitol.
Values are means ±SD of five measurements for each of five plants (n = 5)

235
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

Figure 7. Root fresh weight of different maize varieties in response to different concentrations of mannitol. Data
are means of 5 replicates (n = 5) ± SD

Figure 8. Root-shoot ratio of different maize varieties in response to different concentrations of mannitol. Each
point represents the means ± SD of five measurements on each of five plants (n = 5)

3.5 Changes of Shoot and Root Length during Drought Stress

Shoot and root length of two maize varieties decreased obviously under the stress of 200, 400 and 600 mM
mannitol (Figures 9 and 10). Under the stress of 200, 400 and 600 mM mannitol, shoot length of Zhengdan958
decreased 68.17%, 88.66% and 94.33%, respectively. While shoot length of Liansheng15 under the stress of 200,
400 and 600 mM mannitol decreased 75.20%, 92.32% and 96.80%, respectively (Figure 9). Root length of
Zhengdan958 under the stress of 200, 400 and 600 mM mannitol decreased 37.73%, 64.53% and 84.83%,
respectively. And root length of Liansheng15 under the stress of 200, 400 and 600 mM mannitol decreased
49.65%, 82.39% and 92.74%, respectively (Figure 10).

Figure 9. Shoot length of different maize varieties in response to different concentrations of mannitol. Values are
means ±SD of five measurements for each of five plants (n = 5)

236
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

Figure 10. Root length of different maize varieties in response to different concentrations of mannitol. Data are
means of 5 replicates (n = 5) ± SD

3.6 Drought Resistance Coefficient

The drought resistance coefficients of shoot and root fresh weight and length of two maize varieties under
drought stress decreased obviously with the increase of mannitol concentration. But the drought resistance
coefficients of Zhengdan958 were higher than that of Liansheng15. The drought resistance coefficient of
root-shoot ratio of Zhengdan958 increased during mannitol treatment. While the ratio of Liansheng15 increased
first and then decreased (Table 1). This showed that the drought resistances of Zhengdan958 was higher than that
of Liansheng15 on seed germination condition.

Table 1. Drought resistance coefficient of two maize varieties under 200, 400 and 600 mM mannitol treatment.
Each point represents the means ± SD of five measurements on each of five plants (n = 5)
Mannitol concentration Shoot fresh Root fresh Root-shoot
Variety Shoot length Root length
(mM) weight weight ratio
Zhengdan958 200 0.29a 0.56a 2.34c 0.32a 0.62a
400 0.09b 0.30b 3.74ab 0.11b 0.34b
600 0.05c 0.14c 3.82a 0.06c 0.15c
Liansheng15 200 0.24a 0.54a 2.16b 0.25a 0.50a
400 0.06b 0.13b 2.33a 0.08b 0.18b
600 0.02c 0.04c 2.07c 0.03c 0.07c

4. Discussion
Soil drought is one of the most important environmental factors responsible for the limit of maize productivity in
many areas of the world. Maize appears very sensitive to drought stress during seedling emergence. Some
researches (Grzesiak, 1990; Lorens et al., 1987a, 1987b; Martiniello & Lorenzoni, 1985; Ristic & Cass, 1991)
confirmed the existence in maize of a wide range of genotypic variability of response to drought. Plants develop
different morphological, physiological and biochemical mechanisms which inhibit or remove the harmful effects
of drought stresses (Boyer, 1982; Larsson & Go’rny, 1988; Chaves et al., 2002; Reynolds et al., 1998; Asharaf,
2010). Relatively little research was carried out on the response to drought stress of some plant species.
Intragenotypic variability was studied in maize (Martiniello & Lorenzoni, 1985; Lorens et al., 1987a, 1987b;
Evans et al., 1990; Grzesiak, 1990; Ristic & Cass, 1991). Developing cultivars of maize that can perform well
under heat and drought stress is an important goal throughout the world. However, progress in breeding of
drought resistant maize genotypes requires future study into physiological mechanisms underlying the responses
of maize to water stress (Larsson & Go’rny, 1988; Richards, 1991). In maize, similarly to other species, genotypic
differences in response to soil water deficit were identified with respect to plant physiological characteristics. In
the present study, we investigated the effects of drought stress on seeds germination and seedlings growth of two
maize varieties.
The germination rate under drought stress can show the germination ability of drought resistance of various

237
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

varieties. In this experiment, we preliminary judged the strength of drought resistance according to the
germination rate of two maize varieties under the simulated drought stress with mannitol. The germination rate of
two maize varieties declined obviously under drought stress, and there was no significant difference between the
two varieties. Under the stress of 200, 400 and 600 mM mannitol, germination percentage of Zhengdan958 was
still higher than that of Liansheng15, suggesting that Zhengdan958 had a stronger drought resistance (Figure 2).
Germination potential is a index to evaluate the seed germination rate and germination uniformity. It has been
shown that the germination potential of two maize varieties both declined under drought stress. But Liansheng15
decreased more than that of Zhengdan958 (Figure 3). This suggested that Zhengdan958 germinated more quickly
than Liansheng15, showing a stronger drought resistance of Zhengdan958. In addition, the effect of drought
stress on the germination potential was greater than germination rate, explaining the greater influence of drought
stress on early seed germination. This may be due to the bigger seeds suction influence of Zhengdan958 under
drought stress on the early stage of germination.
Vigor index of drought resistance can better reflect the drought resistance and germination characteristics. The
germination and vigor index of drought resistance of two maize varieties decreased with the increase of mannitol
concentration, but Zhengdan958 decreased less than that of Liansheng15 under the stress of 200, 400 and 600
mM mannitol (Figures 4 and 5). The seedlings of Zhengdan958 grew better under osmotic stress, which could
also be reflected by the seedling fresh weight (Figures 6 and 7) and length (Figures 9 and 10).
Osmotic adjustment is one of main strategies by which plants ensured water uptake during drought stress
conditions. Drought resistance coefficient reflects the sensitive degree of plants to drought stress. From our results,
the drought resistance coefficients of Zhengdan958 were all higher than that of Liansheng15 (Table 1). Roots is
an important absorption organ of plants, developed level of roots is one of the main indexes of drought resistance.
From the physiological point of view, a great root-shoot ratio is beneficial to alleviate drought stress. From the
experimental results, Zhengdan958 had higher root-shoot ratio under drought stress (Figure 8), suggesting that
Zhengdan958 developed powerful root system to fully use of water in deep soil under drought condition, thus
showing stronger drought resistance.
5. Conclusion
In conclusion, under the same drought stress, the stronger drought resistance varieties which have higher
germination and vigor index of drought resistance can still maintain higher germination potential and germination
rate. In this experiment, Zhengdan958 germinated in greater numbers and more quickly than Liansheng15. And
Zhengdan958 also had more powerful root system and stronger ability of osmotic adjustment to ensure seedling
establishment in their drought environments, which might be the important reason for higher resistance to
drought stress of Zhengdan958.
Acknowledgements
We are grateful for financial support from the Supporting Plan of National Science and Technology of China
(2013BAD07B06), the NSFC (National Natural Science Research Foundation of China (31300205), Natural
Science Research Foundation of Shandong (ZR2013CQ009), the Science and Technology Development Projects
of Shandong Province (2014GNC113005) and the Program for Scientific research innovation team in colleges
and universities of Shandong Province.
References
Asharaf, M. (2010). Inducing drought tolerance in plants: Recent advances. Biotech Adv, 28, 199-238.
http://dx.doi.org/10.1016/j.biotechadv.2009.11.005
Battisti, D. S., & Naylor, R. L. (2009). Historical warnings of future food insecurity with unprecedented seasonal
heat. Science, 323, 240-244. http://dx.doi.org/10.1126/science.1164363
Ben-Hayyim, G. (1987). Relationship between salt tolerance and resistance to polyethylene glycol-induced water
stress in cultured citrus. Plant Physiol, 85, 430-433. http://dx.doi.org/10.1104/pp.85.2.430
Boyer, J. S. (1982). Plant productivity and environment. Science, 218, 443-448.
http://dx.doi.org/10.1126/science.218.4571.443
Chaves, M. M., Pereira, J. S., Maroco, J., Rodrigues, M. L., Ricardo, C. P. P., Osorio, L. M., ... Pinheiro, C.
(2002). How plants cope with water stress in the field? Photosynthesis and growth. Ann Bot, 89, 907-916.
http://dx.doi.org/10.1093/aob/mcf105
Delachiave, M. E. A., & Pinho, S. Z. D. (2003). Germination of Sennaoccidentalis link: seed at different osmotic
potential levels. Braz Arch Tech, 46, 163-166. http://dx.doi.org/10.1590/S1516-89132003000200004

238
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

Evans, R. O., Skagss, R. W., & Sneed, R. E. (1990). Normalized crop susceptibility factors for corn and soybean
to excess water stress. Trans ASAE, 33, 1153-1161. http://dx.doi.org/10.13031/2013.31452
Evans, R. O., Skagss, R. W., & Sneed, R. E. (1991). Stress day index models to predict corn and soybean relative
yield under high water table conditions. Trans ASAE, 5, 1997-2005. http://dx.doi.org/10.13031/2013.31829
Gomez, F., Oliva, M. A., Mielke, M. S., Almeida, A. A. F., Leita, H. G., & Aquino, L. A. (2008). Photosynthetic
limitations in leaves of young Brazilian Green Dwarf coconut (Cocos macifera L) “nana’’ palm under
well-watered conditions or recovering from drought stress. Environ Exp Bot, 62, 195-204.
http://dx.doi.org/10.1016/j.envexpbot.2007.08.006
Grzesiak, M. T., Marcin’ska, I., Janowiak, F., Rzepka, A., & Hura, T. (2012). The relationship between seedling
growth and grain yield under drought conditions in maize and triticale genotypes. Acta Physiol Plantarum,
34, 1757-1764. http://dx.doi.org/10.1007/s11738-012-0973-3
Grzesiak, M. T., Waligorski, P., Janowiak, F., Marcinska, I., Hura, K., Szczyrek, P., & Głab, T. (2013). The
relations between drought susceptibility index based on grain yield (DSIGY) and key physiological
seedling traits in maize and triticale genotypes. Acta Physiol Plant, 35, 549-565.
http://dx.doi.org/10.1007/s11738-012-1097-5
Grzesiak, S. (1990). Reaction to drought of inbreds and hybrids of maize (Zea mays L.) as evaluated in field and
greenhouse experiments. Maydica, 35, 303-311. Retrieved from
http://www.cabdirect.org/abstracts/19901617370.html
Hamayun, M., Khan, S. A., Shinwari, Z. K., Khan, A. L., Ahmad, N., & Lee, I. (2010b). Effect of polyethylene
glycol induced drought stress on physio-hormonal attributes of soybean. Pak J Bot, 42, 977-986. Retrieved
from https://scholar.google.com.hk/scholar?q=+Effect+of+polyethylene+glycol+induced+drought+stress+
on+physio-hormonal+attributes+of+soybean.&btnG=&hl=zh-CN&as_sdt=0%2C5
Hamayun, M., Sohn, E. Y., Khan, S. A., Shinwari, Z. K., Khan, A. L., & Lee, I. J. (2010a). Silicon alleviates the
adverse effects of salinity and drought stress on growth and endogenous plant growth hormones of soybean
(Glycine max L.). Pak J Bot, 42(3), 1713-1722. Retrieved from
https://scholar.google.com.hk/scholar?q=Silicon+alleviates+the+adverse+effects+of+salinity+and+drought
+stress+on+growth+and+endogenous+plant+growth+hormones+of+soybean+%28Glycine+max+L%29&bt
nG=&hl=zh-CN&as_sdt=0%2C5
Hu, Y., & Schmidhalter, U. (2005). Drought and salinity: A comparison of their effects on mineral nutrition of
plants. J Plant Nutr Soil Sci, 168, 541-549. http://dx.doi.org/10.1002/jpln.200420516
Khodarahmpour, Z. (2011). Effect of drought stress induced by polyethylene glycol (PEG) on germination
indices in corn (Zea mays L.) hybrids. Afr J Biotech, 10, 18222-18227. Retrieved from
http://www.ajol.info/index.php/ajb/article/view/98591
Larsson, S., & Go’rny, A. G. (1988). Grain yield and drought resistance indices of oat cultivars in field rain
shelter and laboratory experiments. J Agron Crop Sci, 161, 277-286.
http://dx.doi.org/10.1111/j.1439-037X.1988.tb00668.x
Lobell, D. B., Schlenker, W., & Costa-Roberts, J. (2011). Climate trends and global crop production since 1980.
Science, 333, 616-620. http://dx.doi.org/10.1126/science.1204531
Lorens, G. F., Bennett, J. M., & Loggale, L. B. (1987). Differences in drought resistance between two corn
hybrids. I. Water relations and root length density. Agron J, 79, 802-807.
http://dx.doi.org/10.2134/agronj1987.00021962007900050009x
Lorens, G. F., Bennett, J. M., & Loggale, L. B. (1987b). Differences in drought resistance between two corn
hybrids. II. Component analysis and growth rate. Agron J, 79(5), 808-813.
http://dx.doi.org/10.2134/agronj1987.00021962007900050009x
Martinielio, P., & Lorenzoni, C. (1985). Response of maize genotypes to drought tolerance tests. Maydica, 30,
361-370. Retrieved from http://agris.fao.org/agris-search/search.do?recordID=IT876013788
Mishra, V., & Cherkauer, K. A. (2010). Retrospective droughts in the crop growing season: Implications to corn
and soybean yield in the midwestern united states. Agric For Meteorol, 150, 1030-1045.
http://dx.doi.org/10.1016/j.agrformet.2010.04.002
Narusaka, Y., Nakashima, K., Shinwari, Z. K., Sakuma, Y., Furihata, T., Abe, H., ... Shinozaki, K. Y. (2003).
Interaction between two cis-acting elements, ABRE and DRE, in ABA-dependent expression of

239
www.ccsenet.org/jas Journal of Agricultural Science Vol. 7, No. 5; 2015

Arabidopsis rd29A gene in response to dehydration and high salinity stresses. The Plant Journal, 34(2),
137-149. http://dx.doi.org/10.1046/j.1365-313X.2003.01708.x
Paknejad, F., Nasri, M., Moghadam, H. R. T., Zahedi, H., & Alahmadi, M. F. (2007). Effects of drought stress on
chlorophyll fluorescence parameters, chlorophyll content and grain yield of wheat cultivars. J Biol Sci, 7,
841-847. http://dx.doi.org/10.3923/jbs.2007.841.847
Pena, R. D. L., & Hughes, J. (2007). Improving vegetable productivity in a variable and changing climate. SAT
eJournal, 4, 1-22. Retrieved from http://agropedia.iitk.ac.in/openaccess/sites/default/files/sp1.pdf
Ragab, A. R., Abdel-Raheem, A. T., Kasem, Z. A., Omar, F. D., & Samera, A. M. (2007). Evaluation of R1
tomato somaclone plants selected under poly ethylene glycol (PEG) treatments. Afr Crop Sci Soc, 8,
2017-2025. Retrieved from http://www.acss.ws/upload/xml/research/468.pdf
Reynolds, M. P., Singh, R. P., Ibrahim, A., Agech, O. A. A., Larque-Saavedra, A., & Quick, J. S. (1998).
Evaluation physiological traits to complement empirical selection for wheat in warm environments.
Euphytica, 100, 84-95. http://dx.doi.org/10.1023/A:1018355906553
Richards, R. A., & Thurling, N. (1978). Variation between and within species of rapeseed (Brasica campestris
and B napus) in response to drought stress. I. Sensitivity at different stages of development. Aust J Agric
Res, 29, 469-477. http://dx.doi.org/10.1071/AR9780491
Ristic, Z., & Cass, D. D. (1991). Leaf anatomy of Zea mays L in response to water shortage and high
temperature: a comparison of drought-resistant and drought-sensitive lines. Bot Gaz, 152(2), 173-185.
http://dx.doi.org/10.1086/337878
Royo, C., Abaza, M., Bianco, R., & Moral, L. F. G. (2000). Triticale grain growth and morphometry as affected
by drought stress, late sowing and simulated drought stress. Aust J Physiol, 27, 1051-1059.
http://dx.doi.org/10.1071/PP99113
Sakthivelu, G., Devi, M. K. A., Giridhar, P., Rajasekaran, T., Ravishankar, G. A., Nedev, T., & Kosturkova, G.
(2008). Drought induced alterations in growth, osmotic potential and in vitro regeneration of soybean
cultivars. Genet Appl Plant Physiol, 34, 103-112. Retrieved from
http://www.bio21.bas.bg/ippg/bg/wp-content/uploads/2011/06/08_pisa_1-2_103-112.pdf
Taylor, A. G., Moles, J. E., & Kirkham, N. B. (1982). Germination and seedling growth characteristics of three
tomato species affected by water deficits. J Am Soc Hort, 107, 282-285. Retrieved from
http://agris.fao.org/agris-search/search.do?recordID=US19830891444
Winter, S. R., Musick, J. T., & Porter, K. B. (1988). Evaluation of screening techniques for breeding drought
resistant winter wheat. Crop Sci, 28, 512-516.
http://dx.doi.org/10.2135/cropsci1988.0011183X002800030018x

Copyrights
Copyright for this article is retained by the author(s), with first publication rights granted to the journal.
This is an open-access article distributed under the terms and conditions of the Creative Commons Attribution
license (http://creativecommons.org/licenses/by/3.0/).

240