American Journal of Plant Sciences, 2013, 4, 1911-1917 http://dx.doi.org/10.4236/ajps.2013.410234 Published Online October 2013 (http://www.scirp.org/journal/ajps) 1911 Comparison of Salt Tolerance in Brassicas and Some Related Species Jianjie Su1, Shu Wu1, Zhijie Xu1, Si Qiu1, Tingting Luo1, Yimin Yang1, Qitao Chen1, Yuying Xia1, Song Zou1, Bang-Lian Huang1*, Bangquan Huan g1,2* 1College of Life Science, Hubei University, Wuhan, China; 2Hubei Industrial Biotechnology Key Lab, Hubei University, Wuhan, China. Email: *414029354@qq.com, *huangbangquan@163.com Received June 30th, 2013; revised July 29th, 2013; accepted August 15th, 2013 Copyright © 2013 Jianjie Su et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. ABSTRACT In this paper the salt tolerance in Brassicas and some related species was compared. When seedlings germinated on sand cultures with liquid MS medium were considered, the relative germination rate, root length, shoot length and fresh seedling weight were significantly correlated with each other (P < 0.01), and only the relative shoot lengths were sig- nificantly different among the tested genotypes (P < 0.05); When both seedlings germinated on MS and MS plus 0.4% NaCl were considered, only the relative shoot length of seedlings germinated on MS was significantly different from that germinated on MS + 0.4% NaCl (P < 0.05), and also only the relative shoot lengths were significantly different among the tested genotypes (P < 0.01). Raphanu s sativa cv. Changfeng, B. juncea cv. JC and Brassica napus cv. ZS 10 showed low salt tolerance in terms of relative germination rate, root length, shoot length and fresh seedling weight; B. oleracea cv. JF-1, Sinapis alba cv. HN-2 showed high salt tolerance in terms of relative germination rate, root length, shoot length and fresh seedling weight. Based on our result we suggest that relative shoot length might be convenient to rank the salt tolerance but cluster analysis based on multiple parameters of relative germination rate, root length, shoot length and fresh seedling weight might be more accurate in screening for salt tolerance in Brassicas and related species. Keywords: Brassica; Related Species; Salt Tolerance; Multivariate Cluster Analysis 1. Introduction Soil salinity is an issue that affects an estimated 6% of the world’s land surface area or 12,780 million hectares (Mha) and secondary salinization from irrigation impacts an estimated 20% of irrigated land or 1474 Mha [1]. Ac- cording to the United Nations reports, 20% of agricul- tural land and 50% of world cropland are salt affected [2]. Furthermore, there is also a dangerous trend of a 10% per year increase in the saline area throughout the world [3]. China has a large area of inland saline-alkali land, ap- proximately 8.1 MHa, equivalent to 40% of the total cul- tivated land in the country [4,5]. The Brassicaceae family consists of many important field crops such as oilseed rape, the Brassica oleracea ve- getable group (cauliflower, broccoli, brussels sprouts, kale) as well as other species such as Brassica rapa and Raphanus sativus [6]. However, their growth, yield, and oil production are markedly reduced due to salinity [7]. The most common adverse effect of salinity on the crop of Brassica is the reduction in plant height, size and yield as well as deterioration of the product quality [8,9]. In previous studies, the Brassicas were usually grown in salt conditions to compare parameters such as relative germination rate, root length, fresh seedling weight or seed yield [10-17]. In the present study the seeds of Bras- sicas and related species were grown on sand cultures with and without salt to explore an efficient index to com- pare the salt tolerance in Brassicas and related species and to determine variations in degree of salt tolerance at inter- and intra-specific levels particularly at the germi- nation and seedling stages. 2. Materials and Methods 2.1. Materials Seeds of Sinapis alba (cv.HN1, HN2), Brassica carinata cv. EJ and Isatis indigotica cv. HN were provided by Pro. Dr. Zaiyun Li of Huazhong Agricultural University, *Corresponding author. Copyright © 2013 SciRes. AJPS
Comparison of Salt Tolerance in Brassicas and Some Related Species 1912 China. The Brassica napus seeds were from the Oil Crops Institute, CAAS, China. The other seeds were provided by Lab of Plant Genetics and Breeding of Hubei Univer- sity, China. 2.2. Methods Sixty healthy seeds for each genotype were inoculated on sand cultures with liquid MS and MS + 0.4% NaCl. Ger- mination rates were scored 4 - 5 days after inoculation. The germinated seeds on MS and MS + 0.4% NaCl were transferred onto sand cultures with MS and MS + 0.4% NaCl for 3 - 4 days and then root length, shoot length and fresh seedling weight were measured. The temperature was set at 25˚C with a 16-hr photo period under 2000 lx. The experiment was arranged in a completely random- ized design with three replicates. Salt tolerance indices were calculated according to Zeng et al. (2002) [18] and further used to classify genotypes using cluster analysis as described by El. Hendawy et al. (2005) [19]. Cluster analysis was carried out based on Euclidean distance of the salt tolerance indices. The cluster group rankings were obtained from the average of means of the multiple pa- rameters in each cluster group. A sum was obtained by adding the number of cluster group rankings in each ge- notype. The genotypes were finally ranked based on the sums, such that those with the smallest and largest sums were ranked respectively as the most and least tolerant genotypes in terms of relative salt tolerance. Variance ana- lyses, multiple comparisons and cluster analyses were carried out on SPSS. 3. Results 3.1. Correlations among Relative Germination Rate, Root Length, Shoot Length and Fresh Seedling Weight From Table 1 it was found that when seeds were germi- nated on sand cultures with liquid MS medium, the rela- tive germination rate, root length, shoot length and fresh seedling weight were significantly correlated with each other (P < 0.01); When seeds were germinated on sand cultures with liquid MS medium plus 0.4% NaCl, the relative germination rate was only significantly correlat- ed with shoot length (P < 0.05), the relative root length was significantly correlated with shoot length and fresh seedling weight (P < 0.01), and relative shoot length was significantly correlated with fresh seedling weight (P < 0.01). 3.2. Effect of Germination Condition on Relative Root Length, Shoot Length and Fresh Seedling Weight When only seeds germinated on MS medium were con- Table 1. Correlations among relative germination rate, root length, shoot length and fresh seedling weight. Relative germination rate Relative root length Relative shoot length Relative fresh weight Relative germination rate 0.430** 0.416** 0.454** Relative root length 0.019 0.423** 0.486** Relative shoot length0.251* 0.593** 0.804** Relative fresh weight−0.075 0.552** 0.500** Note: Left Bottom: Correlations of relative germination rate, root length, shoot length and fresh weight of seedlings germinated on MS + 0.4% NaCl; Top Right: Correlations of relative germination rate, root length, shoot length and fresh weight of seedlings germinated on MS; **P < 0.01; *P < 0.05. sidered, variance analysis indicated that the difference of relative shoot lengths among the tested genotypes was significant (P < 0.05), while that of relative root length and fresh seedling weight were not significant (data not shown). When both seeds germinated on MS and MS + 0.4% NaCl were considered, variance analysis indicated that the relative shoot length of seedlings germinated on MS was significantly different from that germinated on MS + 0.4% NaCl (P < 0.05), while that of relative root length and fresh seedling weight were not significant; the rela- tive shoot lengths were significantly different among the tested genotypes (P < 0.01), while that of relative germi- nation rate, root length and fresh seedling weight were not significant (data not shown). From Table 2 it was found that based on relative shoot length B. carinata cv. EJ, B. oleracea cv. JF-1 and Eruca sativa cv. hub12 showed high salt tolerance while Rap- hanus sativus cv. changfeng and B. napus cv. ZS 10 were identified as most salt-sensitive. 3.3. Cluster Analysis of Salt Tolerance From Table 3 it was found that R. sativa cv. changfeng, B. juncea cv. JC and B. napus cv. ZS 10 showed low salt tolerance in terms of relative germination rate, root length, shoot length and fresh weight; B. oleracea cv. JF-1, S. alba cv. HN-2 showed high salt tolerance in terms of relative germination rate, root length, shoot length and fresh seedling weight; B. carinata cv. EJ showed high salt tolerance in terms of root length, shoot length and fresh seedling weight but with very low salt tolerance in terms relative germination rate. From Figure 1 it was found that B. oleracea cv. JF-1, B. carinata cv. EJ and Sinapis alba cv. HN-2 were iden- tified as highly salt-tolerant. S. alba cv. HN-1, I. indigo- ica cv. HN and B. rapa cv. xiaobaicai were identified t Copyright © 2013 SciRes. AJPS
Comparison of Salt Tolerance in Brassicas and Some Related Species Copyright © 2013 SciRes. AJPS 1913 Table 2. Multiple comparison of relative shoot length among the tested genotypes when seeds germinated on MS were con- sidered. Genotye 1 2 3 4 5 Duncana,b R. sativus cv. changfeng 51.2267 B. napus cv. ZS 10 52.8000 52.8000 B. oleracea cv. zigan 65.7533 65.7533 65.7533 B. napus cv. ZY 821 66.6100 66.6100 66.6100 R. sativus cv. red 67.1000 67.1000 67.1000 C. abyssinica cv. hubu-1 68.2667 68.2667 68.2667 B. oleracea cv. xueyuan 68.4967 68.4967 68.4967 B. oleracea cv. qinghua 70.9833 70.9833 70.9833 B. juncea cv. JC 71.0333 71.0333 71.0333 B. napus cv. ZS 9 72.7133 72.7133 72.7133 C. abyssinica cv. hubu-2 74.0267 74.0267 74.0267 R. sativus cv. shunyuan 77.6567 77.6567 77.6567 77.6567 C. abyssinica cv. hubu-3 78.7367 78.7367 78.7367 78.7367 B. rapa cv. dabaicai 78.8000 78.8000 78.8000 78.8000 B. napus cv. ZYZ 7 81.7033 81.7033 81.7033 81.7033 R. sativus cv. nanpan 82.0167 82.0167 82.0167 82.0167 B. napus cv. ZS 11 82.4967 82.4967 82.4967 82.4967 R. sativus cv. jiujin 82.5067 82.5067 82.5067 82.5067 R. sativus cv. may 83.8400 83.8400 83.8400 83.8400 B. juncea cv. yongsheng 84.6500 84.6500 84.6500 84.6500 B. oleracea cv. GJ-1 86.2933 86.2933 86.2933 86.2933 E. sativa cv. hub11 86.4000 86.4000 86.4000 86.4000 S. alba cv. HN-1 86.6667 86.6667 86.6667 86.6667 I. indigotica cv. HN 87.1133 87.1133 87.1133 87.1133 B. rapa cv. xiaobaicai 89.6400 89.6400 89.6400 89.6400 E. sativa cv. hub10 94.9733 94.9733 94.9733 S. alba cv. HN-2 95.8467 95.8467 B. rapa cv. shiyue 97.8600 97.8600 E. sativa cv. hub12 102.1733 102.1733 102.1733 B. ol erace a cv. JF-1 117.0767 117.0767 B. carinata cv. EJ 135.7000 Note: The genotypes were classified into 5 groups. Genotypes in the same group are not significantly different from each other. as salt-tolerant, R. sativa cv. changfeng and B. napus cv. ZS 10 were identified as most salt-sensitive. 4. Discussion Although salt stress affects all growth stages of a plant, seed germination and seedling growth stages are known to be more sensitive in most plant species [8,20,21]. Huang et al. (2010) used 0.37% NaCl to compare the salt tolerance during germination and seedling growth among genotypes of B. napus, B. juncea and B. rapa [13]. Our preliminary studies also showed that seed germination, root length, shoot length and fresh seedling weight of Brassicas were inhibited by 20% - 60% when the NaCl concentration was 0.4%. Therefore we used 0.4% NaCl for seed inoculation and seedling growth to compare the salt tolerance. In earlier screening of Brassica species for salt toler- ance, the superiority of amphidiploid species B. carinata, B. juncea, and B. napus over the diploid species, B.rapa, B. nigra, and B. oleracea was proposed from different tudies [8,22-24]. It has been further suggested that the s
Comparison of Salt Tolerance in Brassicas and Some Related Species 1914 Table 3. Rankings of genotypes for their salt tolerance in terms of relative germination rate, root length, shoot length and fresh seedling weight. Genotype Given No. Rank of relative germination rate Rank of relative root length Rank of relative shoot length Rank of relative seedling weight Final genotype ranking R. sativus cv. jiujin 1 29 12 14 21 20 R. sativus cv. changfeng 2 26 29 31 31 31 R. sativus cv. shunyuan 3 7 7 20 26 13 R. sativus cv. nanpan 4 15 21 16 17 17 R. sativus cv. may 5 25 10 13 19 15 R. sativus cv. red 6 19 14 27 29 27 B. napus ZS 11 7 17 6 15 18 12 B. napus cv. ZS 9 8 20 22 22 7 18 B. napus cv. ZY 821 9 16 27 28 12 21 B. napus cv. ZYZ 7 10 24 9 17 13 14 B. napus cv. ZS 10 11 31 13 30 30 29 C. abyssinica cv. hubu-1 12 27 26 26 14 28 C. abyssinica cv. hubu-2 13 12 28 21 23 22 C. abyssinica cv. hubu-3 14 10 15 19 24 16 E. sativa cv. hub12 15 9 23 3 9 8 E. sativa cv. Hub11 16 21 30 10 25 25 E. sativa cv. Hub10 17 13 24 6 11 11 B. oleracea cv. qinghua 18 4 31 24 28 26 B. oleracea cv. zigan 19 23 16 29 16 22 B. oleracea cv. xueyuan 20 22 18 25 20 24 B. oleracea cv. JF-1 21 1 1 2 2 1 B. oleracea cv. GJ-1 22 11 11 11 4 7 B. juncea cv. yongsheng 23 5 19 12 8 8 B. juncea cv. JC 24 30 25 23 27 30 B. rapa cv. xiaobaicai 25 8 8 7 6 4 B. rapa cv. shiyue 26 18 20 4 5 10 B. rapa cv. dabaicai 27 14 17 18 22 18 S. alba cv. HN-1 28 3 5 9 15 5 S. alba cv. HN-2 29 2 2 5 3 2 B. carinata cv. EJ 30 28 3 1 1 6 I. indigotica cv. HN 31 6 4 8 10 3 salt tolerance of amphidiploids has been acquired from the A (B. rapa) and C (B. oleracea) genomes [10]. How- ever, there were also different conclusions as significant inter- and intraspecific variation for salt tolerance exists within Brassicas, which can be exploited through selec- tion and breeding for enhancing salt tolerance of the crops [9,12,13,25-27]. For example, of turnip cultivars, Shaljum desi surakh was highest in seed germination, while it was lowest in seedling shoot dry biomass pro- duction. However, Neela Shaljum having lower seed ger- mination percentage produced maximum seedling shoot dry biomass. In the same way, cv. Desi of radish with minimum seed germination had highest shoot dry weight under saline conditions [12]. Therefore efficient and ac- curate indices and criteria for identifying salt-tolerance in Brassicas and related species are needed. The multivariate cluster analysis method was recom- mended as the best criteria for the identification of salt tolerance in crop species such as rice [18,28,29], green gram [30], wheat [19], tomato [31], sugarcane [32], pea- nut [33], cauliflower [34]. As pointed out by Khrais et al. [35] and Zeng et al. [18], the advantages of using a mul- tivariate analysis in the evaluation of salt tolerance are hat it allows: 1) a simultaneous analysis of multiple t Copyright © 2013 SciRes. AJPS
Comparison of Salt Tolerance in Brassicas and Some Related Species 1915 Figure 1. Hierarchical cluster analysis of the genotypes based on multiple salt tolerance indices of seedlings germinated on MS medium (Euclidean distance). The genotypes that the numbers represent were indicated in Table 3. parameters to increase the accuracy of the genotype ranking; 2) the ranking of genotypes even when plants are evaluated at different salt levels and salt tolerance varies with salinity levels, especially when the salt toler- ance indices are averaged across salt levels; and 3) a more convenient and accurate estimation of salt tolerance among genotypes by simply adding the numbers in clus- ter group ranking at different salt levels. In the present study, when seedlings germinated on sand cultures with liquid MS medium were considered, the relative germination rate, root length, shoot length and fresh seedling weight were significantly correlated with each other (P < 0.01), and the relative shoot lengths were significantly different among the tested genotypes (P < 0.05), while the relative root length and relative fresh seedling weight were not significantly different; when seedlings germinated on liquid MS medium plus 0.4% NaCl were considered, the relative germination rate was only significantly correlated with shoot length (P < 0.05), the relative root length was significantly correlated with shoot length and fresh seedling weight (P < 0.01), and relative shoot length was significantly correlated Copyright © 2013 SciRes. AJPS
Comparison of Salt Tolerance in Brassicas and Some Related Species 1916 with fresh seedling weight (P < 0.01). When both seed- lings germinated on MS and MS plus 0.4% NaCl were considered, the relative shoot length of seedlings germi- nated on MS was significantly different from that ger- minated on MS + 0.4% NaCl (P < 0.05), suggesting that germination in salt condition had some selection effect for shoot length, while relative root length and fresh seed- ling weight were not significantly different; the relative shoot lengths were significantly different among the test- ed genotypes (P < 0.01), while the relative germination rate, root length and fresh seedling weight were not sig- nificantly different. Based on relative shoot length B. carinata cv. EJ, B. oleracea cv. JF-1 and E. sativa cv. hub12 were classified as highly salt-tolerant while R. sativus cv. Changfeng, and B. napus cv. ZS 10 were identified as most salt-sen- sitive. R. sativa cv. changfeng, B. juncea cv. JC and B. napus cv. ZS 10 showed low salt tolerance in terms of relative germination rate, root length, shoot length and fresh seedling weight; B. oleracea cv. JF-1, S. alba cv. HN-2 showed high salt tolerance in terms of relative ger- mination rate, root length, shoot length and fresh seed- ling weight; B. carinata cv. EJ showed high salt toler- ance in terms of root length, shoot length and fresh seed- ling weight but very low salt tolerance in terms relative germination rate. Based on our result we suggest that relative shoot length might be convenient to rank the salt tolerance but cluster analysis based on multiple parameters of relative germination rate, root length, shoot length and fresh seed- ling weight might be more accurate in screening for salt tolerance in Brassicas and related species. 5. 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