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Traits Affecting Low Temperature Tolerance in Tomato and Its Application to Breeding Program
Plant Breed. Biotech. 2019;7:350-359
Published online December 1, 2019
© 2019 Korean Society of Breeding Science.

Rajametov Sherzod1†, Eun Young Yang1†, Myeong Cheoul Cho1, Soo Young Chae1, Jeong Ho Kim1, Chun Woo Nam1, Won Byoung Chae2*

1National Institute of Horticultural & Herbal Science, Rural Development Administration, Wanju 55365, Korea
2Department of Environmental Horticulture, Dankook University, Cheonan 31116, Korea
Corresponding author: *Won Byoung Chae, wbchae75@dankook.ac.kr, Tel: +82-41-550-3642, Fax: +82-41-559-7881
These authors contributed equally.
Received September 4, 2019; Revised October 21, 2019; Accepted October 25, 2019.
This is an Open-Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.
Abstract
It is essential to develop tomato (Solanum lycopersicum L.) cultivars with tolerance to low temperature (LT) for reducing production cost and increasing fruit quality in winter. This study was conducted to investigate the effects of LT on 40 tomato accessions for establishing selection criteria to select accessions with LT tolerance. Tomato plants were grown in two polyethylene film greenhouses with night temperature set-points of 10°C and 15°C. LT significantly affected the photosynthetic parameters as well as both vegetative and reproductive growth in tomato. There was a significant difference in plant height between plants grown in 10°C and 15°C. Leaf length and width were also significantly lower in 10°C. The stem diameter was generally lower in 10°C but 12 accessions in 10°C had thicker stem diameter than those in 15°C. The retarded growth was due to lower photosynthetic rate in 10°C than 15°C. Significant delay in flowering was observed among tomato accessions in 10°C. For the number of flowers, 62.5% showed the increase but 37.5% did the decrease in 10°C among 40 tomato accessions, showing genotype specific interaction with LT. Accessions with small fruits showed reduced fruit set in 10°C; however, it was not obvious among accessions with large fruits. Interestingly, 75% of accessions showed increased chlorophyll contents in 10°C compared to 15°C. An accession ‘VI037163’ was selected for LT tolerance based on good performance in six out of seven traits considered such as plant height, leaf length, stem diameter, chlorophyll contents, days to flowering, the number of flowers and fruit set.
Keywords : Tomato, Breeding, Low temperature during the night, Flowering, Chlorophyll content, Photosynthesis
INTRODUCTION

Tomato (Solanum lycopersicum L.) is one of important greenhouse vegetables grown all-year-round and its cultivation area and production in 2017 were 5,782 ha and 355,107 metric tones, respectively, in Korea (Korean Statistical Information Service, http://kosis.kr). Greenhouses with heating system are mandatory in the winter for tomato cultivation in Korea and the heating cost is the most important factor affecting the production cost of tomato fruits. Approximately 16% of energy cost can be reduced if temperature set-point is reduced by 2℃ in greenhouses (Elings et al. 2005). Therefore, developing tomato cultivars with LT tolerance is crucial for reducing production cost and increasing fruit quality in winter.

Temperature is one of the main environmental factors affecting vegetative and reproductive growth of tomato. In LT, which is defined as sub-optimal temperature above the chilling temperature, tomato plants exhibit reduction in both growth rate (Paul 1984; Hoek et al. 1993; Venema et al. 1999) and photosynthesis (Venema et al. 1999). LT also delays the formation of leaf, truss appearance (Hurd and Graves 1985; Khayat et al. 1985; Nieuwhof et al. 1997; Adams et al. 2001) and fruit growth (Hurd and Graves 1985; De Koning 2000; Adams et al. 2001).

Night temperature in greenhouses is more important to reduce energy cost since ambient temperature is generally lower in the night than day. Besides, high temperature during the daytime can compensate for low temperature during the night (LTN), in terms of the growth of tomato plants (De Koning 1988). The reduction in energy cost can be achieved by using covering materials such as thermal screens but the best way is simply to decrease the night temperature in greenhouses. Therefore, breeding program for tomato cultivars with LT tolerance should focus on the night temperature.

In order to establish the optimal selection criteria in tomato breeding program for LT tolerance, it is important to investigate the response of tomato accessions to LT in large scale. Several studies investigated the effect of the LT on the growth and yield of tomato; however, most of studies were conducted with less than five cultivars and only few studies with more than ten cultivars (Der Van Ploeg and Heuvelink 2005). This study was conducted to evaluate the effect of LTN on 40 tomato accessions in greenhouses with different temperature set-points for establishing optimal selection criteria to select accessions with LT tolerance.

MATERIALS AND METHODS

Tomato (Solanum lycopersicum L.) is one of important greenhouse vegetables grown all-year-round and its cultivation area and production in 2017 were 5,782 ha and 355,107 metric tones, respectively, in Korea (Korean Statistical Information Service, http://kosis.kr). Greenhouses with heating system are mandatory in the winter for tomato cultivation in Korea and the heating cost is the most important factor affecting the production cost of tomato fruits. Approximately 16% of energy cost can be reduced if temperature set-point is reduced by 2℃ in greenhouses (Elings et al. 2005). Therefore, developing tomato cultivars with LT tolerance is crucial for reducing production cost and increasing fruit quality in winter.

Temperature is one of the main environmental factors affecting vegetative and reproductive growth of tomato. In LT, which is defined as sub-optimal temperature above the chilling temperature, tomato plants exhibit reduction in both growth rate (Paul 1984; Hoek et al. 1993; Venema et al. 1999) and photosynthesis (Venema et al. 1999). LT also delays the formation of leaf, truss appearance (Hurd and Graves 1985; Khayat et al. 1985; Nieuwhof et al. 1997; Adams et al. 2001) and fruit growth (Hurd and Graves 1985; De Koning 2000; Adams et al. 2001).

Night temperature in greenhouses is more important to reduce energy cost since ambient temperature is generally lower in the night than day. Besides, high temperature during the daytime can compensate for low temperature during the night (LTN), in terms of the growth of tomato plants (De Koning 1988). The reduction in energy cost can be achieved by using covering materials such as thermal screens but the best way is simply to decrease the night temperature in greenhouses. Therefore, breeding program for tomato cultivars with LT tolerance should focus on the night temperature.

In order to establish the optimal selection criteria in tomato breeding program for LT tolerance, it is important to investigate the response of tomato accessions to LT in large scale. Several studies investigated the effect of the LT on the growth and yield of tomato; however, most of studies were conducted with less than five cultivars and only few studies with more than ten cultivars (Der Van Ploeg and Heuvelink 2005). This study was conducted to evaluate the effect of LTN on 40 tomato accessions in greenhouses with different temperature set-points for establishing optimal selection criteria to select accessions with LT tolerance.

RESULTS

Growing condition for tomato accessions

Greenhouse condition for both 10℃ and 15℃ was shown in Fig. 1. Temperature set-point at night in the first week was maintained at 15℃ in both greenhouses for adapting seedlings to the new environment. Minimum and maximum temperatures in a greenhouse with temperature set-point 10℃ were 7℃ and 33℃, respectively, and those in a greenhouse with temperature set-point 15℃ were 11℃ and 38℃ (Fig. 1a). The mimimum temperature in the greenhouse with temperature set-point 10℃ was slightly lower in the first half of growing period than the second half, probably due to unstable sensor or heating system. However, the minimum temperature after the 9th week was fully controlled, maintaining a constant temperature of approximately 10℃ (Fig. 1a). The average RH ranged from 45 to 50% in both greenhouses (Fig. 1b). Minimum and maximum RH were generally lower and higher, respectively, in the greenhouse with temperature set-point 15℃ than 10℃.

Vegetative growth of tomato accessions in LT

LT significantly affected the plant growth. The plant height in 10℃ was significantly lower than that in 15℃ in all growing period (Fig. 2a, b). Larger differences in plant height between 10℃ and 15℃, which mean susceptible to LT, were observed in T31, T01, T36 and T05 (35.3, 36.5, 36.9, and 37.3%, respectively) and smaller differences, tolerant to LT, were in T20, T10, T12 and T06 (75.4, 72.9, 69.7, and 66.5%, respectively) (data now shown). Similar to plant height, 40 and 37 accessions have short leaf length and width, respectively, in 10℃ than 15℃. Accessions T33, T10, T20 and T07 showed smallest difference in leaf length such as 89.9, 87.6, 85.9 and 82.5%, respectively (Table 2). The leaf width of T33 and T10 increased in 10℃ by 19.7 and 8.4%, respectively. However, the other 37 accessions, except for T13, showed decrease in leaf width in 10℃ (Table 2). The average stem diameter in 40 accessions also tend to be lower in 10℃ than 15℃ but 12 accessions (T10, T33, T07, T37, T27, T17, T14, T26, T21, T40, T04 and T24) in 10℃ had thicker stem diameter than those in 15℃ (Table 2).

Photosynthesis among 40 accessions significantly reduced in LT (Fig. 3). In early growing period (until 30 DAT), photosynthetic rate A was higher in 10℃ than 15℃. However, it was reversed thereafter and the difference was significant between 65 and 95 DAT (Fig. 3a). Photosynthetic rate A in both temperature conditions decreased over time during the entire growing period (Fig. 3a). Stomatal conductance and transpiration rate were also significantly higher in 15℃ than 10℃ in all growing period (Fig. 3b, c). Intercellular CO2 concentration was also higher in 15℃ but differences were not significant except for 65 DAT (Fig. 3d).

Reproductive growth of tomato accessions in LT

Flowering was significantly delayed in all accessions when grown in 10℃ (Fig. 4). On average, the flowering initiated 23.9 days later in the accessions grown in 10℃ than those in 15℃. Accessions T01 and T20 flowered earlier than others in both 10℃ (46.3 and 48.0 DAT, respectively) and 15℃ (27.8 and 33.3 DAT, respectively) (Fig. 4). Difference in days to flowering was greatest in T09 as 82% (76.0 and 41.8 DAT in 10℃ and 15℃, respectively) and smallest in T10 as 15.7% (73.5 and 63.5 DAT in 10℃ and 15℃, respectively).

The LT condition significantly influenced the number of flowers among the accessions (Fig. 5). Among 40, 25 accessions (62.5%) showed the increase in the number of flowers in 10℃ but it decreased in 15 accessions (37.5%). This could be due to the genotype specific interaction with LT. The difference in the number of flowers between 10℃ and 15℃ was largest in T14 (224.7%) and smallest in T12 (53.7%) (Fig. 4).

Fruit set tend to decrease in LT condition among the 40 tomato accessions. For instance, the fruit set decreased in 29 accessions (72.5%) whereas increased in 10 accessions (25%) (Fig. 6) and no difference was observed in a T16 (2.5%). The decrease in fruit set was genotype specific; 79.2% of accessions (19 out of 24) with cherry and medium sized fruits but 40% of accessions (6 out of 15) with large fruits had decreased fruit set ratio in 10℃ (Fig. 6). The accessions such as T03, T15, T17, T19, T25, T35 and T4 did not bear fruits on both the second and third trusses at 10℃, showing high susceptibility to LT (data not shown). Accessions T10, T14, T30 and T38 displayed highest difference in fruit set but T03, T09, T28 and T35 the smallest (Fig. 6).

Chlorophyll content of tomato accessions in LT

The chlorophyll content in all tomato accessions steadily increased during the entire growing period regardless of temperature conditions in greenhouses. Tomato accessions grown in 10℃ tend to have higher chlorophyll contents than in 15℃ but 10 accessions showed lower chlorophyll content in 10℃ than 15℃ (Fig. 7). The maximum and minimum chlorophyll contents in 15℃ at 125 DAT were 60.3 (T22) and 32.3 mg/cm2 (T01), respectively, those in LT were 6.9 (T06) and 37.5 mg/cm2 (T15), respectively. Differences in chlorophyll contents between 10℃ and 15℃ were highest in T27 (146.2%) and lowest in T11 (83.4%) (Fig. 7).

Selection of tomato accessions for LT tolerance

To select accessions with LT tolerance, seven traits were chosen such as plant height, leaf length, stem diameter, chlorophyll contents, days to flowering, the number of flowers and fruit set (Table 3), but leaf width was excluded due to high correlation with leaf length (data not shown). After selecting top four accessions (10th percentile) in each trait, an accession T10, ‘VI037163’, which has cherry type fruits (about 20 g) with brownish skin color and high sugar content, was selected for LT tolerance, based on good performance in six among a total of seven traits (Table 3).

DISCUSSION

LT has a large effect on the vegetative development in tomato. In our study, the plant growth was significantly delayed in LT in all the 40 tomato accessions (Fig. 2). Vegetative growth such as plant height, leaf length and width, and stem diameter were significantly reduced in 10℃ compared to 15℃ (Fig. 2, Table 2) although some accessions have increased stem diameter in 10℃ (Table 2). Our results are consistent with previous results that reported reduced growth rate among tomato plants grown in LTN (Smeets and Garretsen 1986; Franco 1990; Nieuwhof et al. 1991; Der Van Ploeg and Heuvelink 2005).

The difference in the growth rate between 10℃ and 15℃ might be the result of the reduced photosynthesis of tomato plants in LT because all photosynthetic parameters were higher in 15℃ than 10℃ (Fig. 3). The decrease in photosynthetic rate was also observed in wild Lycopersicon species when day temperature was reduced (Venema et al. 1999) and Fang et al. (2018) also reported reduced photosynthetic rate and stomatal conductance in tomato plants under LT stress condition. It is interesting to observe the increased chlorophyll contents among plants grown LT (Fig. 7) even though photosynthesis was significantly reduced in LT (Fig. 3). Further study necessary to clarify the relationship between chlorophyll content and photosynthesis in LT condition.

LT also affect the reproductive growth of tomato accessions. Days to flower was significantly delayed in all accessions grown in 10℃ compared to 15℃ (Fig. 4). However, the delayed flowering may be not the direct cause of LT but delayed growth of tomato plants (Fig. 2). A previous research supports our assumption, indicating that the delay in the time to flowering of tomato plants in LT is due to the reduced rate of leaf appearance (Hurd and Graves 1985).

Previously, the only increase in the number of flowers was reported in decreased day (Rylski 1979), night (Calvert 1957; Wittwer and Tuebner 1957) and root temperatures (Phatak et al. 1966). In our study, however, both the increase (25 accessions) and decrease (15 accessions) in the number of flowers were observed in 10℃ compared to 15℃ (Fig. 5). Our result suggests that the effect of LT on the number of flowers is genotype specific, showing interaction between genotypes and LT, which is still unknown (Fig. 5).

Interestingly, 72.0% of accessions (18 out of 25) with small fruits (wild, cherry and medium sized fruits) but only 40% of accessions (6 out of 15) with large fruits had decreased fruit set in 10℃ compared to 15℃ (Fig. 6), indicating a possible relationship between fruit set and fruit size. The reduced fruit set might be due to poorer quality pollen in LT (reviewed by Picken 1984) and not effects on stigma level or ovule viability (Fernandez-Munoz and Cuartero 1991).

Seven traits such as plant height, leaf length, stem diameter, chlorophyll contents, days to flowering, the number of flowers and fruit set were used in our study for the selection of tomato accessions with LT tolerance (Table 3). Selection criteria was based on the difference in those traits between 10℃ and 15℃. The largest percentage in all parameters, except for days to flower and the number of flowers, was assumed to be tolerant to LT (Table 3). For the days to flowering, accessions that have the smallest percentages were selected. For the number of flowers, accessions that were not affected by LT were selected because both the increase and decrease may be not good for the fruit yield; the increase can make plants use more energy to produce flowers and the decrease can affect the number of fruits produced. The traits chosen may be useful for tomato breeding program because they are easy to investigate in field and the data can be collected in the relatively early growing period.

ACKNOWLEDGEMENTS

This study was supported by a grant (Project No: PJ012662 “Breeding and selection of tomato lines with tolerance to abnormal temperatures stress”) from National Institute of Horticultural and Herbal Science, Rural Development Administration.

Figures
Fig. 1.

Air temperature (a) and relative humidity (b) measured during the entire growth period in greenhouses whose night temperature set-points were 10 and 15℃. LT and NT: greenhouses with night temperature set-points of 10 and 15℃, respectively.


Fig. 2.

The growth of tomato accessions in 10℃ and 15℃. (a) The tomato plants grown in greenhouses with night temperature set-points of 10℃ (left) and 15℃ (right). Photos were taken 125 days after transplanting. (b) Changes in average plant height of 40 accessions during entire growing period.


Fig. 3.

Photosynthetic rate A (a), stomatal conductance (b), transpiration rate (c) and intercellular CO2 concentration of five tomato accessions (T09, T16, T24, T28 and T37), measured 30, 65, 95 and 125 days after transplanting in greenhouses with night temperature set-point of 10℃ and 15℃.


Fig. 4.

Days to flowering among 40 tomato accessions grown in greenhouses whose night temperature set-point were 10 and 15℃.


Fig. 5.

Difference in the number of flowers between 10 and 15℃ among 40 tomato accessions. Lower and higher than 100% means the number of flowers in 10℃ was lower or higher than those in 15℃, respectively. A transverse red line indicates the 100%, no difference in the number of flowers between 10 and 15℃.


Fig. 6.

Difference in fruit set between 10 and 15℃ among 40 tomato accessions. Negative percentage means decrease in fruit set of plants grown in 10℃ greenhouse compared to 15℃.


Fig. 7.

Difference in chlorophyll contents between 10 and 15℃ among 40 tomato accessions. Lower and higher than 100% means the chlorophyll contents of plants grown in 10℃ was lower or higher than those in 15℃, respectively. A transverse red line indicates the 100%, no difference in chlorophyll content between 10 and 15℃.


Tables

Accessions used in this study and their fruit types.

Accessions Original name Fruit Type
T01 L3708 Wild
T02 Wonhong 1 Cherry
T03 Wonung 1 Cherry
T04 Tabtimdaeng T2021 Cherry
T05 B-Blocking Cherry
T06 Minichal Cherry
T07 AVTO1020 Cherry
T08 Sugar Yellow × 11AVT-2_4, F8 Cherry
T09 Wonye 9014 Cherry
T10 VI037163 Cherry
T11 Wonye 9015 Cherry
T12 Sincheonggang Cherry
T13 Tom Little Medium
T14 Power Guard Medium
T15 Sugar Yellow × CLN3125-A, F8 Medium
T16 L05945 Medium
T17 VI030494 Medium
T18 TY Strong Medium
T19 Sugar Yellow × CLN3125-E, F8 Medium
T20 B-Strong Medium
T21 12AVT-22 × Bacchus, F2 Medium
T22 Tom Heart Medium
T23 IT 032935 (CL80-0-2-0) Medium
T24 12AVT-14 × Dafnis, F4 Medium
T25 Tomate RASTEIRO RIO GRANDE Medium
T26 IT 032964 (CL185-0-1-0) Large
T27 AVTO1314 Large
T28 Sinheuksu Large
T29 Tamseure Large
T30 AVTO0101 Large
T31 Pink Top F3 Large
T32 Avemaria Large
T33 Tamnara Large
T34 K151771 Large
T35 Dafnis Large
T36 T1-Mobir (Uzbekistan) Large
T37 Tomate SUPER MARMANDE Large
T38 SV0244TG Large
T39 - Large
T40 IT 247072 Large

Values are mean ± standard deviation. Comparison of the heading date in 2014 (HD), grain length (GL), grain width (GW), 1000 grain weight and grain fertility of wild-type and wspl. Data show mean and SD of biological replicates (n > 10) and statistically calculated by Student’s t-test (NS,

*, and ** stand for not statistically significant at the 0.05 probability level, significant at the 0.05 and 0.01 probability levels, respectively).


Effects of low temperature on leaf length and width, and stem diameter among 40 tomato accessions.

Accessions Leaf length (cm) Difference (%) Leaf width (cm) Difference (%) Stem diameter (mm) Difference (%)



10℃ 15℃ 10℃ 15℃ 10℃ 15℃
T01 19.0 ± 1.6 33.3 ± 1.5 57.1 19.5 ± 1.6 29.8 ± 1.0 65.4 11.6 ± 0.3 12.7 ± 1.1 91.3
T02 18.3 ± 1.1 45.8 ± 3.4 40.0 17.6 ± 1.1 48.5 ± 3.9 35.7 15.2 ± 0.5 15.8 ± 0.3 96.2
T03 23.5 ± 4.5 44.7 ± 1.7 52.6 20.3 ± 3.3 43.7 ± 1.5 46.5 12.0 ± 0.8 16.2 ± 0.9 74.1
T04 30.0 ± 2.4 43.3 ± 2.1 69.3 31.0 ± 4.0 40.0 ± 3.5 77.5 13.2 ± 0.1 12.9 ± 0.1 102.3
T05 23.0 ± 1.1 36.5 ± 1.8 63.0 25.6 ± 1.4 42.8 ± 2.9 59.8 17.7 ± 0.3 20.2 ± 0.9 87.6
T06 31.5 ± 1.1 49.5 ± 2.5 63.6 33.6 ± 1.7 43.3 ± 2.5 77.6 15.4 ± 1.4 16.0 ± 0.5 96.3
T07 28.3 ± 2.2 34.3 ± 3.4 82.5 27.8 ± 2.5 34.8 ± 8.4 79.9 16.0 ± 0.9 14.5 ± 1.7 110.3
T08 30.0 ± 0.7 45.5 ± 4.6 65.9 28.6 ± 0.5 40.0 ± 3.2 71.5 13.7 ± 0.4 13.7 ± 0.5 100
T09 21.3 ± 2.2 40.5 ± 2.1 52.6 25.8 ± 2.9 39.0 ± 2.0 66.2 14.0 ± 0.7 16.0 ± 0.9 87.5
T10 29.6 ± 1.7 33.8 ± 1.6 87.6 29.8 ± 1.8 27.5 ± 2.9 108.4 15.9 ± 0.8 12.5 ± 1.1 127.2
T11 23.2 ± 3.0 40.0 ± 2.9 58.0 24.7 ± 4.6 39.0 ± 2.4 63.3 14.9 ± 0.7 18.5 ± 1.8 80.5
T12 24.5 ± 0.7 40.0 ± 3.0 61.3 27.4 ± 1.3 50.5 ± 4.3 54.3 17.5 ± 0.9 18.8 ± 0.9 93.1
T13 23.9 ± 0.6 29.5 ± 1.9 81.0 31.0 ± 1.0 31.0 ± 1.6 100.0 13.7 ± 0.3 15.2 ± 1.3 90.1
T14 29.1 ± 1.2 43.3 ± 1.1 67.2 29.5 ± 1.9 42.0 ± 3.0 70.2 15.0 ± 0.6 14.2 ± 0.0 105.6
T15 28.6 ± 2.2 38.8 ± 2.8 73.7 28.5 ± 1.4 42.5 ± 3.6 67.1 13.5 ± 0.4 16.4 ± 2.0 82.3
T16 29.3 ± 2.6 40.8 ± 1.9 71.8 27.5 ± 1.9 35.3 ± 1.9 77.9 15.3 ± 0.8 16.3 ± 1.0 93.9
T17 18.5 ± 2.9 36.8 ± 2.6 50.3 16.3 ± 2.0 34.3 ± 1.5 47.5 14.1 ± 0.1 13.1 ± 0.9 107.6
T18 23.6 ± 1.5 47.5 ± 4.3 49.7 27.1 ± 3.2 49.3 ± 2.2 55.0 17.4 ± 0.1 18.7 ± 1.0 93
T19 29.0 ± 1.5 37.3 ± 2.9 77.7 38.0 ± 2.1 46.0 ± 3.1 82.6 14.8 ± 1.6 16.8 ± 0.8 88.1
T20 26.9 ± 1.5 31.3 ± 1.1 85.9 29.5 ± 1.7 33.0 ± 2.3 89.4 17.6 ± 0.7 18.6 ± 1.1 94.6
T21 29.8 ± 2.1 49.3 ± 2.7 60.4 35.5 ± 3.5 54.8 ± 6.0 64.8 15.6 ± 0.6 14.9 ± 0.4 104.7
T22 25.4 ± 1.1 40.5 ± 0.6 62.7 29.8 ± 1.6 39.0 ± 1.7 76.4 14.0 ± 0.7 15.7 ± 0.6 89.2
T23 28.4 ± 1.7 34.5 ± 1.5 82.3 30.9 ± 2.2 37.5 ± 5.5 82.4 16.3 ± 0.4 16.6 ± 0.9 98.2
T24 39.0 ± 0.6 60.0 ± 2.0 65.0 42.0 ± 3.1 50.0 ± 0.8 84.0 15.0 ± 0.4 14.8 ± 0.4 101.4
T25 30.7 ± 2.4 39.5 ± 2.1 77.7 35.7 ± 5.4 40.8 ± 0.5 87.5 17.8 ± 0.8 19.6 ± 1.1 90.8
T26 29.6 ± 1.5 49.5 ± 4.7 59.8 30.8 ± 1.2 51.0 ± 4.8 60.4 19.1 ± 0.2 18.1 ± 1.3 105.5
T27 34.6 ± 0.4 44.5 ± 3.2 77.8 38.8 ± 3.5 47.5 ± 4.4 79.6 19.5 ± 0.3 17.9 ± 0.6 108.9
T28 28.8 ± 1.2 41.0 ± 1.4 70.2 33.0 ± 1.0 46.5 ± 2.1 71.0 15.4 ± 0.9 16.4 ± 1.8 93.9
T29 34.5 ± 1.6 45.0 ± 1.8 76.7 39.5 ± 1.2 43.5 ± 1.3 90.8 17.7 ± 0.2 18.8 ± 0.8 94.1
T30 26.5 ± 1.4 42.0 ± 1.8 63.1 28.0 ± 1.8 35.5 ± 2.3 78.9 17.3 ± 1.1 17.5 ± 0.7 98.9
T31 30.3 ± 0.7 44.0 ± 2.1 68.9 36.5 ± 2.3 47.3 ± 2.6 77.2 17.2 ± 1.3 19.2 ± 1.4 89.6
T32 26.3 ± 3.1 46.3 ± 1.7 56.8 32.5 ± 3.3 49.8 ± 2.3 65.3 14.1 ± 0.9 16.2 ± 0.2 87
T33 32.1 ± 1.7 35.7 ± 0.9 89.9 39.5 ± 2.7 33.0 ± 1.0 119.7 16.4 ± 0.6 13.8 ± 1.6 118.8
T34 29.7 ± 2.4 43.3 ± 3.4 68.6 32.0 ± 3.5 41.5 ± 3.0 77.1 17.3 ± 1.5 18.3 ± 1.6 94.5
T35 24.1 ± 1.6 51.8 ± 2.3 46.5 31.0 ± 1.4 60.0 ± 1.5 51.7 17.7 ± 0.4 20.3 ± 0.5 87.2
T36 20.8 ± 1.3 41.8 ± 2.3 49.8 27.9 ± 2.1 47.0 ± 2.6 59.4 17.3 ± 0.3 18.8 ± 0.7 92
T37 28.5 ± 1.8 39.3 ± 3.1 72.5 29.8 ± 2.6 37.3 ± 2.6 79.9 18.9 ± 0.4 17.2 ± 1.4 109.9
T38 29.8 ± 3.0 54.0 ± 0.6 55.2 34.3 ± 2.1 53.8 ± 5.4 63.8 14.8 ± 0.8 21.5 ± 1.5 68.8
T39 33.5 ± 0.7 54.5 ± 2.1 61.5 37.5 ± 2.9 52.8 ± 2.0 71 18.0 ± 1.2 21.0 ± 1.0 85.7
T40 29.3 ± 2.4 42.8 ± 2.5 68.5 30.0 ± 1.5 46.5 ± 3.3 64.5 17.1 ± 0.6 16.6 ± 0.4 103

± Means standard errors (n = 3).


Selection criteria for accessions with low temperature tolerance and selected accessions for 7 traits.

Traits Selection criteria (difference between 10 and 15℃ ) Selected accessions (10th percentile)
Plant height The largest percentage T20, T10, T12, T06
Leaf length The largest percentage T33, T10, T20, T07
Stem diameter The largest percentage T10, T33, T07, T37
Chlorophyll contents The largest percentage T27, T21, T10, T06
Days to flowering The smallest percentage T10, T33, T14, T25
The number of flowers No difference T16, T27, T35, T32
Fruit set ratio The largest percentage T14, T38, T10, T30

± Means standard errors (n = 3).


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