陸柏憲1,* 詹庭筑2 賴永昌3
摘要
陸柏憲、詹庭筑、賴永昌。2020。甘藷誘變品系耐鹽性之研究。台灣農業研究 69(4):298–
311。
本試驗以甘藷 (Ipomoea batatas) 誘變品系 (P、13、17、87、89) 與對照品種「台農 57 號」、「紅心尾」為 材料，分別於組織培養、盆栽及田間三方面進行鹽分逆境試驗，分析甘藷於鹽分逆境下之外表型性狀及生理反 應，以評估參試品系 (種) 之鹽分耐受性。試驗結果顯示，不同誘變品系之耐鹽指數 (salt tolerance index; STI)
因不同性狀而異，且隨鹽分濃度增加而下降；組培苗期性狀於0.5% 鹽分濃度下表現穩定，當鹽分濃度達 1.0%
時，甘藷生長明顯受到抑制，其中誘變品系以13、89 號之根長表現最佳。此外，盆栽試驗結果顯示，甘藷
以150 mM NaCl 處理下塊根重量表現穩定，以誘變品系 87 號之 STI 為最高，且甘藷塊根累積大量 Na+及麥 芽糖含量增加，推測為參試品系具較強耐鹽性之原因。進一步評估田間試驗之結果，鹽分逆境對甘藷地上部
之負面影響較輕微，而對根部有較大之負面效應，導致甘藷產量降低，但減產程度因不同品系 (種 ) 而異，
其中以「紅心尾」、P 及 87 號產量顯著較高。綜合所有試驗結果，在鹽分逆境條件下，甘藷誘變品系 P、87
號於整個栽培期可正常生長且塊根產量較高，顯示兩品系有較高之鹽分耐受性，可供台灣未來育種選用之耐 鹽甘藷雜交親本。
關鍵詞：甘藷、誘變品系、耐鹽性。
投稿日期：2020 年 8 月 7 日；接受日期：2020 年 9 月 28 日。
* 通訊作者：[email protected]
1 農委會農業試驗所嘉義農業試驗分所農藝系助理研究員。台灣 嘉義市。
2 農委會台中區農業改良場作物改良課助理研究員。台灣 彰化縣。
3 農委會農業試驗所嘉義農業試驗分所農藝系研究員兼系主任。台灣 嘉義市。
DOI:10.6156/JTAR.202012_69(4).0004
前言
作物栽培生產過程中，生物性或非生物性 逆境常是限制作物生長發育的環境因子，其中 因土壤鹽化所造成的影響屬於非生物性逆境，
嚴 重 限 制 現 今 農 業 發 展 與 土 地 利 用。 依 據 美 國 農 業 部 (United States Department of Agri-culture; USDA) 訂定之分類標準，土壤 pH 值
< 8.5、飽和抽出液電導度 (electrical conduc-tivity; EC) ≥ 4 dS m-1時，稱之為鹽土 (saline soil) (United States Department of Agriculture 1975; Zaman et al. 2018)。全球土壤鹽化面積 根據不同研究者的估算而略有差異，Massoud (1981) 指 出 在 全 球 1.5 × 109 ha 已 開 墾 的 可
耕地中，約有3.4 × 108 ha 為鹽土。另根據聯 合 國 糧 農 組 織 (Food and Agriculture Organi-zation of the United Nations; FAO) 之 統 計，
全 球 約 有9.32 × 108 ha 的土壤受鹽分所影響 (Abrol
et al. 1988)。此外，根據 FAO 之官方
網站指出，全球可耕及不可耕地中，近2.3 × 108 ha 的水田中約有 4.5 × 107 ha 受鹽分影響；近1.5 × 109 ha 的旱田中約有 3.2 × 107 ha 的土 壤 受 鹽 分 影 響 (Food and Agriculture Organi-zation of the United Nations 2000)。就台灣土 壤而言，受鹽分影響之土壤多分布在西部沿海 地區，面積約有2.5 × 104 ha，主要由於海水 倒灌及過量施用肥料所致 (Chen et al. 2015)。
值得注意的是，由於灌溉農業等人類活動所造 研究報告
臺灣農業研究69(4)-04 陸柏憲.indd 298
臺灣農業研究69(4)-04 陸柏憲.indd 298 2020/12/17 上午 10:52:592020/12/17 上午 10:52:59
成之土壤次生鹽鹼化作用 (secondary (reactive oxygen species; ROS) 清除系統等 (Taji
et al. 2002; Ueda et al. 2003; Medeiros et al.
2014)；另一則是基於作物於鹽分逆境下表現 之典型農藝性狀，如發芽率、存活率、相對生 長 率、 株 高 及 產 量 等 (Zeng et al. 2002; Kat-erji
et al. 2003; Parida & Das 2005; Grieve et al. 2012)。Grieve et al. (2012) 評估不同作物
在鹽分逆境下之相對產量，並劃分成四種相對 耐鹽等級：耐鹽 (tolerant)，如大麥 (Hordeumvulgare L.)、棉花 (Gossypium hirsutum L.)、
燕麥 (Avena sativa L.) 等；中度耐鹽 (moder-ately tolerant)，如高粱 [Sorghum bicolor (L.) Moench]、大豆 (Glycine max L.)、小麥
(Trit-icum aestivum L.) 等； 中 度 感 鹽 (moderately
sensitive)，如甘藷 [Ipomoea batatas (L.) Lam.]、玉 米 (Zea mays L.)、花生 (Arachis hypogaea L.)、馬鈴薯 (Solanum tuberosum L.) 等；感鹽 (sensitive)， 如 水 稻 (Oryza sativa L.)、 芝 麻 (Sesamum indicum L.)、 菜 豆 (Phaseolus
vul-garis L.) 等。
甘藷為旋花科 (Convolvulaceae) 之一年生 草本作物，主要以莖蔓進行扦插繁殖，與馬鈴 1994; Lai et al. 2008)，但以往之育種程序中 並無特別針對耐鹽特性進行選拔評估，以至較 甲基磺酸乙酯 (ethyl methane sulfonate; EMS) 應用於水稻誘變育種，已獲得耐鹽、耐旱等耐 非生物逆境之水稻種原 (Wang et al. 2002; Wu 2013; Chou et al. 2015)，顯示以誘變方式改變 作物特性、擴大作物遺傳變異，以獲得目標性
(‘Tainung No. 73’; ‘TN73’) 雜交種子所獲之誘 變品系 (P、13、17、87、89) 為材料，分別於
300 台灣農業研究 第69 卷 第 4 期 2% (v/v) 次氯酸鈉水 (hypochlorous acid) 消毒 20 min 後，以無菌水清洗 3 次。將消毒完成 之節間接種於MS (Murashige & Skoog 1962) 基 本 鹽 類 培 養 基 (含 30 g L-1蔗 糖、30 g L-1 Na-alginate，pH 5.7 ± 0.1)。接種後之甘藷瓶 苗置於溫度25℃ ± 2℃、光照週期 16 h 之環境 (glucose)、果糖 (fructose)、蔗糖 (sucrose) 及 麥芽糖 (maltose)。試驗方法參考 Picha (1985) 之步驟，取5 g 之甘藷泥加入 80% 乙醇，混合 均勻後置入80℃熱水中，隔水加熱萃取 3 次。
萃取液過濾後定量至10 mL，再以 0.45 μm 拋 棄式針筒過濾器過濾後，進行高效液相層析儀 (high performance liquid chromatography; HPLC) 分析 (pump model No. LC1150, GBC Scientific Equipment, Dandenong, Australia)。使用 reverse phase C18 分析管柱 (Model No. LiChrolut RP-18, Merck, Darmstadt, Germany)，流動相 (mobile phase) 組成為 80% 的氰甲烷 (acetonitrile) 與 系 (P、13、17、87、89)、「紅心尾」及 ‘TN57’，
剪取先端苗作為試驗材料，扦插於台南市七股 區之鹽化土壤進行試驗。該田區土壤經電導度 計 (Spectrum 2265 FSTP, Spectrum Technolo-gies, Aurora, IL, USA) 與 pH/EC 計測定土壤
耐鹽指數 (salt tolerance index; STI) 試驗中各性狀之STI 值為鹽分處理組與對 照組之比值，數值越高則代表耐鹽性越佳，其 運算公式：STI = NaCl – treated value/control value。
試驗設計
組 織 培 養 及 盆 栽 試 驗 採 完 全 隨 機 設 計 (complete randomized design; CRD)， 田 間 試 驗採完全隨機區集設計 (randomized complete block design; RCBD)。試驗所得數據計算平均 值 與 標 準 誤 (standard error of mean; SEM)，
並以套裝軟體RStudio 進行變方分析 (analysis of variance; ANOVA)。 若 處 理 因 子 達 顯 著 差
302 台灣農業研究 第69 卷 第 4 期
表1. 不同NaCl濃度對甘藷品系 (種) 組織培養苗之影響。 Table 1. Effect of different concentrations (0.0, 0.5 and 1.0%) of NaCl on in vitro sweet potato traits. TreatmentVariety and mutant lines of sweet potato ‘TN57’P13178789 (A) Plant height (cm) 0.0% NaCl14.75 ± 0.65 bAz 15.25 ± 0.93 bA14.38 ± 0.72 bA17.56 ± 1.02 aA15.11 ± 0.82 bA11.00 ± 0.53 cAB 0.5% NaCl17.06 ± 1.23 aA17.24 ± 1.11 aA15.63 ± 1.43 aA16.75 ± 1.10 aA14.90 ± 4.05 aA12.56 ± 0.74 aA STIy 1.161.131.090.950.991.14 1.0% NaCl4.30 ± 1.69 bcB3.61 ± 1.36 bcB10.80 ± 2.59 aA6.59 ± 2.24 abcB2.85 ± 0.89 cB8.88 ± 1.90 abB STI0.290.240.750.380.190.81 (B) Shoot length (cm) 0.0% NaCl3.03 ± 0.18 bA4.90 ± 0.28 aA3.95 ± 0.44 bA4.93 ± 0.34 aA3.91 ± 0.40 bA3.78 ± 0.24 bA 0.5% NaCl4.00 ± 0.66 aA4.24 ± 0.18 aA3.98 ± 0.14 aA4.86 ± 0.33 aA4.13 ± 0.42 aA4.60 ± 0.21 aA STI1.320.861.010.991.051.22 1.0% NaCl1.81 ± 0.12 bB1.68 ± 0.22 bB1.84 ± 0.14 bB2.53 ± 0.24 aB1.81 ± 0.21 bB2.84 ± 0.38 aB STI0.600.340.470.510.460.75 (C) Root length (cm) 0.0% NaCl11.73 ± 0.52 aA10.35 ± 1.06 aA10.43 ± 0.78 aA12.64 ± 1.14 aA11.20 ± 0.72 aA7.23 ± 0.54 aA 0.5% NaCl13.06 ± 1.19 aA13.00 ± 0.96 aA11.65 ± 1.46 aA11.89 ± 1.11 aA10.98 ± 3.75 aA7.96 ± 0.74 aA STI1.111.261.120.940.981.10 1.0% NaCl4.05 ± 2.01 aB3.61 ± 1.79 aB9.03 ± 2.61 aA3.88 ± 2.05 aB1.14 ± 0.76 aB5.60 ± 1.42 aA STI0.350.350.870.310.100.78 (D) Fresh weight (g) 0.0% NaCl0.26 ± 0.03 cB0.49 ± 0.04 abA0.53 ± 0.09 abB0.57 ± 0.06 aB0.37 ± 0.04 bcA0.59 ± 0.07 aA 0.5% NaCl0.52 ± 0.09 bcA0.57 ± 0.07 bcA0.74 ± 0.05 abA0.86 ± 0.12 aA0.34 ± 0.10 cA0.63 ± 0.12 abA STI1.991.171.401.510.931.05 1.0% NaCl0.12 ± 0.02 bcB0.10 ± 0.01 bcB0.22 ± 0.05 abC0.16 ± 0.03 abcC0.09 ± 0.02 cB0.27 ± 0.07 aB STI0.470.210.410.280.240.46 z Mean ± standard error. Means within each row (in small letter) and within each column (in capital letter) followed by the same letter(s) are not significantly different at 5% level by least significant difference (LSD) test. y STI: salt tolerance index.
臺灣農業研究69(4)-04 陸柏憲.indd 302
臺灣農業研究69(4)-04 陸柏憲.indd 302 2020/12/17 上午 10:53:002020/12/17 上午 10:53:00
苗扦插於盆栽中進行耐鹽試驗。試驗分成控制 150 mM NaCl 處理組為 1.88、1.36、11.96 及 0.00 mg L-1 (低於偵測極限 )。誘變品系 13 號 理組為1.18、1.08、9.49 及 0.12。綜合前述結 果，‘TN57’、「紅 心 尾」、P、13、87 號 葡 萄
304 台灣農業研究 第69 卷 第 4 期
Type of saccharide Saccharide content (mg L-1 )
(A)
glucose fructose maltose sucrsoe
glucose fructose maltose sucrsoe 89
Fig. 1. Changes of the soluble sugar content of sweet potato mutant tubers under 150 mM NaCl. Vertical bars indi-cate standard error of mean. (A) ‘TN57’; (B) ‘Hong Xin Wei’; (C) P; (D) 13; (E) 17; (F) 87; and (G) 89.
臺灣農業研究69(4)-04 陸柏憲.indd 304
臺灣農業研究69(4)-04 陸柏憲.indd 304 2020/12/17 上午 10:53:022020/12/17 上午 10:53:02
表2. 150 mM NaCl處理對甘藷品系 (種) 塊根重量之影響。 Table 2. Effect of 150 mM NaCl on sweet potato tuber weight. Tuber weight (g)Variety, line and mutant lines of sweet potato ‘TN57’‘Hong Xin Wei’P13178789 0 mM NaCl 564.5 ± 6.5 aAz560.3 ± 15.3 abA376.8 ± 83.0 bcA501.9 ± 43.7 abcA321.9 ± 33.9 cA345.1 ± 86.9 cB506.0 ± 62.4 abcA 150 mM NaCl596.5 ± 50.5 abA678.5 ± 53.5 aA314.5 ± 27.5 bcA512.0 ± 63.0 abcA249.8 ± 56.3 cA506.5 ± 16.9 abcA376.8 ± 100.2 abcA STIy1.061.210.831.020.781.470.74 z Mean ± standard error. Means within each row (in small letter) and within each column (in capital letter) followed by the same letter(s) are not significantly different at 5% level by least significant difference (LSD) test. y STI: salt tolerance index. 表3. 150 mM NaCl處理對甘藷品系 (種) 塊根鈉離子含量分析。 Table 3. The sodium ion content of sweet potato mutant tubers under 150 mM NaCl.
Sodium ion (mg 100 g
-1 ) Variety, line and mutant lines of sweet potato ‘TN57’‘Hong Xin Wei’P13178789 0 mM NaCl28.5 ± 8.5 abBz 9.8 ± 0.1 bB 25.0 ± 0.2 abB 22.5 ± 6.2 abB 42.4 ± 17.3 aB14.2 ± 1.8 bB16.6 ± 2.7 abB 150 mM NaCl113.4 ± 0.4 bA89.8 ± 7.6 bA126.0 ± 30.3 bA116.9 ± 36.1 bA272.5 ± 27.8 aA139.2 ± 29.0 bA148.2 ± 45.3 bA STIy3.989.195.055.206.439.808.91 z Mean ± standard error. Means within each row (in small letter) and within each column (in capital letter) followed by the same letter(s) are not significantly different at 5% level by least significant difference (LSD) test. y STI: salt tolerance index. 表4. 甘藷品系 (種) 於田間試驗與鹽土試驗之農藝性狀表現。 Table 4. The agronomy traits of sweet potato mutants under normal and saline soil field.
Cultivation environment and crop season
TraitVariety, line and mutant lines of sweet potato ‘TN57’‘Hong Xin Wei’P13178789 Normal soil fieldStem weight (kg plot-1 )z 10.6 ± 0.2 dy -13.5 ± 1.7 cd14.7 ± 3.3 bcd18.0 ± 1.6 ab20.4 ± 2.6 a17.4 ± 1.8 abc Fall Crop 2018 Yield (kg plot-1)19.2 ± 0.4 ab-22.8 ± 2.0 a16.4 ± 1.6 bc14.0 ± 0.9 c16.1 ± 2.0 bc14.6 ± 0.6 bc Harvest index0.64-0.630.530.440.440.46 Saline soil fieldStem weight (kg plot-1)5.3 ± 0.2 b10.6 ± 1.8 b15.0 ± 1.6 ab23.7 ± 3.5 a5.4 ± 0.4 b27.7 ± 5.9 a15.8 ± 5.4 ab Fall Crop 2019 Yield (kg plot-1 )8.6 ± 0.3 abc12.3 ± 2.7 ab11.9 ± 4.3 ab5.0 ± 0.2 c4.4 ± 0.3 c13.9 ± 3.0 a6.2 ± 0.2 bc Harvest index0.620.540.440.170.450.330.28 z The experiment was replicated three times with plot size of 6.0 m × 3.3 m. The sweet potato was planted in 1.10 m × 0.25 m of row and plant spacing. y Mean ± standard error. Means within each row (in small letter) followed by the same letter(s) are not significantly different at 5% level by least significant difference (LSD) test.
臺灣農業研究69(4)-04 陸柏憲.indd 305
臺灣農業研究69(4)-04 陸柏憲.indd 305 2020/12/17 上午 10:53:022020/12/17 上午 10:53:02
306 台灣農業研究 第69 卷 第 4 期 離體篩選 (in vitro screening)。Dasgupta et al.
(2008) 以 15 個不同甘藷種原為材料，接種於 低 排 序：89、13、‘TN57’、17、P、87；另由 於甘藷生產較注重塊根，若由根長性狀之耐鹽
甜菜鹼 (betaine)、山梨糖醇 (sorbitol) 等，降 低 細 胞 滲 透 勢， 以 利 細 胞 能 繼 續 吸 收 水 分 而 恢復正常生長 (Ashraf & Harris 2004; Bartels
& Sunkar 2005; Parida & Das 2005; Arzani 2008)。Wu et al. (2013) 以不同甜菜 (Beta vulgaris
節之緣故；但進一步檢視不同醣類含量增減不
Bartels & Sunkar 2005; Grieve et al. 2012)。
一般植物利用以下三種機制避免過量Na+之危 害：(1) 根部限制對 Na+之吸收；(2) 透過 Na+/ H+反向運輸蛋白 (antiporter) 將 Na+隔離於液 胞中；(3) 透過 Na+/H+反向運輸蛋白將Na+反 向排出細胞 (Bartels & Sunkar 2005; Grieve et
al. 2012; Yu et al. 2016)。根據 Yu et al. (2016)
(phytoremediation) 以改良鹽土 (Boonsaner &Hawker 2012; Jesus et al. 2015)。 本 文「紅 心 麥 [Triticum turgidum L. subsp. durum (Desf.) Husn.] 品系為材料 (包含耐鹽與感鹽品系 )，
308 台灣農業研究 第69 卷 第 4 期
Salt-affected Soils and Their Management. Food and Agriculture Organization of the United Nations.
Rome, Italy. 131 pp.
Arzani, A. 2008. Improving salinity tolerance in crop plants: A biotechnological view. In Vitro Cell. Dev.
Biol. Plant 44:373–383.
doi:10.1007/s11627-008-9157-7
Ashraf, M. and P. J. C. Harris. 2004. Potential biochem-ical indicators of salinity tolerance in plants. Plant Sci. 166:3–16. doi:10.1016/j.plantsci.2003.10.024 Bartels, D. and R. Sunkar. 2005. Drought and salt
tol-erance in plants. Crit. Rev. Plant Sci. 24:23–58.
doi:10.1080/07352680590910410
Boonsaner, M. and D. W. Hawker. 2012. Remediation of saline soil from shrimp farms by three different plants including soybean [Glycine max (L.) Merr.]. J.
Environ. Sci. Health A 47:558–564. doi:10.1080/10 934529.2012.650559
Chandler, S. F., K. Y. Paek, E. C. Pua, E. Ragolsky, B.
B. Mandal, and T. A. Thorpe. 1988. The effective-ness of selection for salinity tolerance using in vitro shoot culture. Intl. J. Plant Sci. 149:166–172.
doi:10.1086/337704
Chen, Z. S., Z. Y. Hseu, and C. C. Tsai. 2015. The Soils of Taiwan. Springer. Dordrecht, Germany. 127 pp.
doi:10.1007/978-94-017-9726-9
Chou, S. Y., I. C. Tsai, W. J. Chang, and H. S. Wu.
2015. Salt tolerant characters of salt tolerant rice mutant CNY1031130. Crop Environ. Bioinform.
12:183–197. (in Chinese with English abstract) doi:10.30061/CEB.201512_12(4).0003
Dasgupta, M., M. R. Sahoo, P. C. Kole, and A. Mukherjee.
2008. Evaluation of orange-fleshed sweet potato (Ipomoea batatas L.) genotypes for salt tolerance through shoot apex culture under in vitro NaCl me-diated salinity stress conditions. Plant Cell Tissue Organ Cult. 94:161. doi:10.1007/s11240-008-9400-2 Food and Agriculture Organization of the United Nations.
2000. Extent and causes of salt-affected soils in participating countries. http://www.fao.org/ag/AGL/
agll/spush/topic2.htm#top (visit on 03/01/2020) Grieve, C. M., S. R. Grattan, and E. V. Maas. 2012. Plant salt
tolerance. p.405–459. in: Agricultural Salinity Assess-ment and ManageAssess-ment. 2nd ed. (Wallender, W. W. and K. K. Tanji, eds.) ASCE. Reston, VA. 1124 pp.
Houshmand, S., A. Arzani, S. A. M. Maibody, and M.
Feizi. 2005. Evaluation of salt-tolerant genotypes of durum wheat derived from in vitro and field exper-iments. Field Crops Res. 91:345–354. doi:10.1016/
j.fcr.2004.08.004
Hunsche, M., C. Lankes, H. Hoffstall, and G. Noga.
2010. Vegetative performance, leaf water potential, and partitioning of minerals and soluble sugars:
Traits for ranking the NaCl-tolerance of tomato genotypes? Plant Growth Regul. 62:151–162.
doi:10.1007/s10725-010-9500-y
Jesus, J. M., A. S. Danko, A. Fiúza, and M. T. Borges.
2015. Phytoremediation of salt-affected soils: A
臺灣農業研究69(4)-04 陸柏憲.indd 308
臺灣農業研究69(4)-04 陸柏憲.indd 308 2020/12/17 上午 10:53:022020/12/17 上午 10:53:02
review of processes, applicability, and the impact of climate change. Environ. Sci. Pollut. Res. Intl.
22:6511–6525. doi:10.1007/s11356-015-4205-4 Katerji, N., J. W. van Hoorn, A. Hamdy, and M.
Mast-rorilli. 2003. Salinity effect on crop development and yield, analysis of salt tolerance according to several classification methods. Agric. Water Manag.
62:37–66. doi:10.1016/S0378-3774(03)00005-2 Kitayama, M., T. Samphumphuang, R. Tisarum, C.
Theer-awitaya, K. Cha-um, M. Takagaki, and S. Cha-um.
2020. Calcium and soluble sugar enrichments and physiological adaptation to mild NaCl salt stress in sweet potato (Ipomoea batatas) genotypes. J. Hor-tic. Sci. Biotechnol. doi:10.1080/14620316.2020.17 49532
Lai, Y. C., Z. T. Li, T. L. Jeng, and W. H. Tsai. 2008.
Breeding of the new sweet potato variety, Tainung No. 73. J. Taiwan Agric. Res. 57:279–294. (in Chinese with English abstract) doi:10.6156/
JTAR/2008.05704.05
Li, L. and C. H. Liao. 1994. The achievement and pros-pect of sweet potato breeding in Taiwan. p.11–28.
in: Proceeding of a Symposium on Root Crop Yield Improvement, Processing and Utilization. No-vember 25, 1993. Chiayi, Taiwan. Taiwan. Chiayi Agricultural Experiment Branch, TARI. Taichung, Taiwan. (in Chinese with English abstract)
Massoud, F. I. 1981. Salt affected soils at a global scale and concepts for control. Food and Agriculture Or-ganization of the United Nations. Rome, Italy. 21 pp.
Medeiros, C. D., J. R. C. Ferreira Neto, M. T. Oliveira, R. Rivas, V. Pandolfi, É. A. Kido, J. I. Baldani, and M. G. Santos. 2014. Photosynthesis, antioxi-dant activities and transcriptional responses in two sugarcane (Saccharum officinarum L.) cultivars under salt stress. Acta Physiol. Plant. 36:447–459.
doi:10.1007/s11738-013-1425-4
Mukherjee, A. 2001. Effect of NaCl on axillary shoot pro-liferation in sweet potato. Ann. Trop. Res. 23:1–10.
Murashige, T. and F. Skoog. 1962. A revised medium for rapid growth and bio assays with tobacco tissue cultures. Physiol. Plant. 15:473-497. doi:10.1111/
j.1399-3054.1962.tb08052.x
Oldeman, L. R., R. T. A. Hakkeling, and W. G. Sombroek.
1991. World map of the status of human-induced soil degradation: An explanatory note. 2nd ed. In-ternational Soil Reference and Information Center.
Vienna, Austria. 35 pp.
Picha, D. H. 1985. HPLC determination of sugars in raw and baked sweet potatoes. J. Food Sci. 50:1189–
1190. doi:10.1111/j.1365-2621.1985.tb13045.x
Parida, A. K. and A. B. Das. 2005. Salt tolerance and salinity effects on plants: A review. Ecotoxical.
Environ. Saf. 60:324–349. doi:10.1016/j.eco-env.2004.06.010
Qadir, M., E. Quillérou, V. Nangia, G. Murtaza, M.
Singh, R. J. Thomas, P. Drechsel, and A. D. Noble.
2014. Economics of salt-induced land degradation and restoration. Nat. Resour. Forum 38:282–295.
doi:10.1111/1477-8947.12054
Shahid, S. A. 2013. Irrigation-induced soil salinity under different irrigation systems- Assessment and man-agement: Short technical note. Clim. Change Out-look Adapt. Intl. J. 1:19–24.
Taji, T., C. Ohsumi, S. Iuchi, M. Seki, M. Kasuga, M. Ko-bayashi, K. Yamaguchi-Shinozaki, and K. Shinozaki.
2002. Important roles of drought- and cold-induc-ible genes for galactinol synthase in stress toler-ance in Arabidopsis thaliana. Plant J. 29:417–426.
doi:10.1046/j.0960-7412.2001.01227.x
Tester, M. and R. Davenport. 2003. Na+ tolerance and Na+ transport in higher plants. Ann. Bot. 91:503–527.
doi:10.1093/aob/mcg058
Ueda, A., M. Kanechi, Y. Uno, and N. Inagaki. 2003.
Photosynthetic limitations of a halophyte sea aster (Aster tripolium L.) under water stress and NaCl stress. J. Plant Res. 116:63–68. doi:10.1007/s10265-002-0070-6
United States Department of Agriculture. 1975. Soil Taxonomy: A Basic System of Soil Classification for Making and Interpreting Soil Surveys. Soil Con-servation Service and United States Department of Agriculture. Washington, DC. 754 pp.
Vargas, R., E. I. Pankova, S. A. Balyuk, P. V. Krasilnikov, and G. M. Khasankhanova. 2018. Handbook for Saline Soil Management. Food and Agriculture Organization of the United Nations and Lomonosov Moscow State University. Moscow, Russia. 132 pp.
Vijayan, K., S. P. Chakraborti, and P. D. Ghosh. 2003. In vitro screening of mulberry (Morus spp.) for salinity tolerance. Plant Cell Rep. 22:350–357. doi:10.1007/
s00299-003-0695-5
Wang, C. S., T. H. Tseng, and C. Y. Lin. 2002. Rice biotech research at the Taiwan Agriculture Re-search Institute. Asia Pacific Biotech 6:950–956.
doi:10.1142/S0219030302001969
Wu, G. Q., N. Liang, R. J. Feng, and J. J. Zhang. 2013.
Evaluation of salinity tolerance in seedlings of sugar beet (Beta vulgaris L.) cultivars using proline, sol-uble sugars and cation accumulation criteria. Acta Physiol. Plant. 35:2665–2674. doi:10.1007/s11738-013-1298-6
Wu, Y. P. 2013. Drought tolerant rice lines by
screen-臺灣農業研究69(4)-04 陸柏憲.indd 309
臺灣農業研究69(4)-04 陸柏憲.indd 309 2020/12/17 上午 10:53:022020/12/17 上午 10:53:02
310 台灣農業研究 第69 卷 第 4 期
ing an ‘IR64’ mutant stock. J. Taiwan Agric. Res.
62:195–208. (in Chinese with English abstract) doi:10.6156/JTAR/2013.06203.01
Yu, Y., T. Xu, X. Li, J. Tang, D. Ma, Z. Li, and J. Sun.
2016. NaCl-induced changes of ion homeostasis and nitrogen metabolism in two sweet potato (Ipo-moea batatas L.) cultivars exhibit different salt tol-erance at adventitious root stage. Environ. Exp. Bot.
129:23–36. doi:10.1016/j.envexpbot.2015.12.006 Zair, I., A. Chlyah, K. Sabounji, M. Tittahsen, and H.
Chlyah. 2003. Salt tolerance improvement in some wheat cultivars after application of in vitro selection
pressure. Plant Cell Tiss. Org. Cult. 73:237–244.
doi:10.1023/A:1023014328638
Zaman, M., S. A. Shahid, and L. Heng. 2018. Guideline
Zaman, M., S. A. Shahid, and L. Heng. 2018. Guideline