- 學位論文
熱馴化和甲磺酸乙酯對番茄植株生長之影響
Effect of Heat Acclimation and Ethyl Methanesulfonate on the Growth of Tomato Plants
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摘要
番茄（Solanum lycopersicum）是一種具有重要經濟意義的作物，因其良好的風味、營養價值和較短的生命週期而在全球各地種植；然而，因為番茄植株對高溫敏感，氣候變遷對番茄生產構成嚴重威脅，因此迫切需要製定有助於減輕氣候變遷對作物生產有破壞性影響的策略。甲磺酸乙酯（EMS）的化學誘變已被用來誘導不同作物中之有益突變，另外，透過將植物暴露在逐漸升高的溫度下的熱馴化處理可在許多作物中誘導出耐熱性；本研究旨在探討EMS誘變和熱馴化對不同氣候條件下番茄植株營養生長和生殖生長的影響。首先將番茄種子‘麗光’進行EMS誘變，以0.00%~2.25%不同濃度的EMS處理種子，然後將發芽的幼苗移至戶外植床上生長，直到收穫果實和種子，在為期四個月的冬季試驗中收集了不同的植物營養生長和生殖生長參數。在熱馴化試驗中，將兩片真葉之番茄幼苗暴露於四個熱馴化處理（T1-T4），然後進行致死溫度挑戰，根據恢復期間幼苗成活率、株高、莖直徑和葉片數量篩選最佳熱馴化處理；對照組、熱馴化 (HA)組和第二代 EMS 衍生植物 (EMS-M2) 在室外盆中培養直至開花和結果，在為期兩個月的夏季試驗中收集了不同的植物營養生長和生殖生長參數，然後將三組植物的葉綠素含量、脯胺酸含量和電解質滲漏進行生化分析比較，以了解高溫對植物生化特性的影響。結果顯示，隨著EMS濃度的增加，種子發芽率降低，種子發芽率在 0.25% EMS 時開始下降，在 1.75% EMS 時發芽率從80% 降至0%；與EMS濃度高於1.00%的處理相比，0.00%至1.00%較低EMS濃度處理的植株在株高、莖直徑和葉片數量方面具有更好的生長特性；第一批開花的番茄植株來自0.00%、0.25%、0.50%和1.00% EMS處理，於播種後54-55天開花；0.00%、0.50%、1.00% EMS處理在播種後75、76天最先結果； 0.00%和0.50%的EMS處理是最早在播種後123-124天果實成熟的處理；各處理間之開花數沒有顯著差異，因為所有處理中花數為每株 11.67 至 18 朵花； 0.25%-1.00%的EMS處理的果實數量較多，為每株10.07-15.18個果實，高於對照組處理的每株8.75個果實； 0.25%至1.25%的EMS處理果實最大，尺寸範圍為55至61毫米，最小的果實來自對照組植物（0.00%），果實直徑為 39 毫米；0.25%至1.50%的EMS處理產生的果實重量為60至106克，而對照組植物（0.00%）的果實重量為34克。在熱馴化實驗中，致死溫度定義為45℃暴露3小時，幼苗死亡率為100%，選擇溫度逐漸升高的T4 處理 [28℃（1 h）- 32℃（1 h）- 36℃（1 h）- 40℃（1 h）] 作為最佳熱馴化處理，因為它在存活率、株高、莖直徑和葉片數上優於其他處理。在夏季試驗期間，觀察到熱馴化植物比 EMS-M2 和對照組植物具有更好的生長參數；熱馴化植物的花數最多，每株植物有 14.78 朵花，其次是對照組植物和 EMS-M2 植物，每株植物分別有 7.98 朵和 7.40 朵花；僅在熱馴化植物中觀察到坐果，而對照組植物和EMS-M2植物不產生任何果實；熱馴化植物的總葉綠素含量最高為3.33毫克，其次是EMS-M2植物為2.13毫克，對照組植物為1.29毫克； EMS-M2和對照組植物具有較高的脯胺酸含量，分別為2.52和2.58毫克，而熱馴化植物的脯胺酸含量為1.66毫克；對照組植物的電解質洩漏率最高為 35.76%，其次是 EMS-M2 植物為 26.72%，而熱馴化植物為 18.13%。綜上所述，EMS 處理並無導致番茄植株提早開花、提早結果、提早果實成熟和更高的花數，然而，EMS 處理在冬季栽培下誘導出更多的果實數量和更高的果實品質（重量、大小），而熱馴化處理改善了夏季番茄植株的營養和生殖生長。未來EMS誘變可以與熱馴化結合，以改良在高溫條件下生長的番茄植株。
並列摘要
Tomato (Solanum lycopersicum) is a crop of major economic importance and is grown all around the globe due to its favorable flavor, nutritional value, and short life cycle. However, climate change poses a serious threat to tomato production as the tomato plants are sensitive to high temperatures. Strategies that help to mitigate the damaging effects of climate change on crop production are urgently needed. Chemical mutagenesis using ethyl methanesulfonate (EMS) have been used to induce beneficial mutations in different crops. Heat acclimation has been found to induce acquired thermotolerance in many crops through the exposure of plants to gradually increasing temperatures. This study aimed to investigate the effect of EMS mutagenesis and heat acclimation on the growth and reproduction of tomato plants under different climatic conditions. EMS mutagenesis was conducted on tomato seeds ‘麗光’ by treating the seeds with different EMS concentrations ranging from 0.00% to 2.25%. The germinated seedlings were grown in outdoor raised beds till fruit and seed harvesting. Different vegetative and reproductive parameters were collected throughout the winter trail over a four-month period. Heat acclimation was conducted by exposing two true-leaf tomato seedlings to four heat acclimation treatments (T1-T4) followed by the lethal temperature challenge. The best heat acclimation treatment was selected based on seedling survival percentage, plant height, stem diameter, and number of leaves during recovery. The control, heat acclimated (HA) and second generation of EMS-derived plants (EMS-M2) were grown in outdoor pots till flowering and fruiting. Different vegetative and reproductive parameters were collected throughout the summer trial over a two-month period. Biochemical analysis of chlorophyll content, proline content, and electrolyte leakage were compared between the three groups of plants to understand the effect of high temperatures on the plants’ biochemical properties. Results showed that the seed germination percentage decreased with the increase of EMS concentrations. Seed germination started to decline at 0.25% EMS, from 80% germination to 0% at 1.75% EMS. Plants from the lower EMS concentration treatments of 0.00% to 1.00% had better growth performance compared to EMS concentration treatments above 1.00% in terms of plant height, stem diameter, and number of leaves. The first tomato plants to flower were from the 0.00%, 0.25%, 0.50%, and 1.00% EMS treatments having flowers at 54-55 days after sowing. The EMS treatments of 0.00%, 0.50%, and 1.00% were the first to yield fruits at 75 and 76 days after sowing. The EMS treatments of 0.00% and 0.50% were the first treatments to have ripe fruits at 123-124 days after sowing. There was no significant difference in the number of flowers per plant as all the treatments had between 11.67 and 18 flowers per plant. The EMS treatment of 0.25%-1.00% had a higher number of fruits with 10.07-15.18 fruits per plant than the control treatment with 8.75 fruits per plant. The EMS treatments from 0.25% to 1.25% had bigger fruits measuring 55 to 61 mm, whereas the control treatment (0.00%) had smaller fruits of 39 mm in diameter. The EMS treatments from 0.25% to 1.50% produced fruits weighing from 60 to 106 g, while the control treatment (0.00%) had fruit weights of 34 g. The lethal temperature was defined as 45℃ for 3 h exposure with 100% seedling mortality. The T4 treatment with gradually increasing temperatures of 28℃ (1 h) – 32℃ (1 h) – 36℃ (1 h) - 40℃ (1 h) was selected as the best heat acclimation treatment as it outperformed the other treatments with high survival percentage, plant height, stem diameter, and number of leaves following the lethal temperature challenge. During the summer trial, it was observed that the heat acclimated plants had better growth parameters than the EMS-M2 and the control plants. The heat acclimated plants had the highest number of flowers, with 14.78 flowers per plant, followed by the control and EMS-M2 plants, with 7.98 and 7.40 flowers per plant, respectively. Fruit setting was only observed in the heat acclimated plants, while the control and EMS-M2 plants did not produce any fruit. The heat acclimated plants had the highest total chlorophyll content of 3.33 mg, followed by the EMS-M2 plants with 2.13 mg and the control plants with 1.29 mg. The EMS-M2 and control plants had higher proline contents of 2.52 and 2.58 mg compared to the heat acclimated plants of 1.66 mg. The highest electrolyte leakage was recorded in the control plants at 35.76%, followed by the EMS-M2 plants at 26.72% and heat acclimated plants with 18.13%. In conclusion, the EMS treatments did not induce early flowering, early fruiting, early fruit ripening, and higher flower number in tomato plants in the winter season. However, the EMS treatments induced a higher number of fruits and higher fruit quality (weight, size). Heat acclimation improved the vegetative and reproductive growth of tomato plants in the summer months. EMS mutagenesis may be combined with heat acclimation in the future for the improvement of tomato plants grown under high temperature conditions.
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