臺灣博碩士論文加值系統

摘要

本實驗利用會產生異型葉的植物異葉水蓑衣，探討環境因子對其生長與產生異型葉的影響。由實驗中發現，葉片會由葉芽組織受環境因子影響，在卵型葉與羽狀葉間逐片發生轉變，其節間長度亦會隨季節不同(夏季0.67 ± 0.046 cm、秋冬季2.29 ± 0.136 cm、冬春季2.63 ± 0.085 cm)有明顯差異(p＜0.05)，尤其夏季更短(p＜0.01)。卵型葉與羽狀葉在型態與構造上有許多明顯的差異，卵型葉的異化指數為0.92±0.007，而羽狀葉則為 0.08±0.004。卵型葉的葉片較厚(p＜0.01)、細胞內的葉綠體、表皮的氣孔都比較多(p＜0.01)，而且上下表皮長有腺毛，羽狀葉上下表皮則完全沒有腺毛存在。卵形葉的電子傳遞最大速率(ETRmax) (21.99±1.72)比羽狀葉高(14.38±0.90)，因此光合作用速率也比羽狀葉高。
實驗的調控因子包括水位、光強度、溫度與光周期。經調控因子處理後所產生的結果為新生的葉片數、新生葉片的總面積、植物莖的總長度，以及異化指數的變化量等四項。全淹與半淹水位處理後這四項結果皆具有顯著差異(p＜0.01)、光強度在正常光與加強光下除了植株生長總長度沒有差異外，其它項目都具有顯著差異(p＜0.01)。溫度對生長具有顯著影響(p＜0.01)，15℃時生長最差(新生葉片1.3±0.16片、總面積6.51±0.23 cm2、總長度7.69±0.12 cm、異化指數變量-0.01±0.008)， 30℃生長最佳(新生葉片4.0±0.27片、總面積29.03±1.66 cm2、總長度12.01±0.55cm、異化指數變量0.16±0.035)，但達到35℃時生長則受到抑制(新生葉片3.1±0.19片、總面積16.35±0.50 cm2、總長度9.98±0.38 cm、異化指數變量0.23±0.026)。隨著溫度的升高，葉片羽狀化更加明顯(p＜0.01)。光週期除了對植株生長總長度沒有差異外，其它項目都具有顯著差異(p＜0.01)，而且光週期增加新生的葉片數也隨之增加。隨著光週期的升高，葉片羽狀化更加明顯(p＜0.01)。實驗的調控因子對於植物的生長，除了植株生長總長度較不明顯外，新生的葉片數與葉片的總面積都有顯著的影響。調控因子對葉片的羽狀化有顯著的影響，尤其是在溫度方面，15℃及35℃的異化指數變量相差最大達到0.24，而正常光與加強光的異化指數變量相差最小只有0.02。

關鍵詞：異葉水蓑衣，異型葉，異化指數變量，電子傳遞最大速率。

Abstract

Aquatic plant, Hygrophila difformis, has the heterophylly and produces two different types of leaf. In this study, we plan to investigate the effect of various environmental factors on the growth and its heterophylly production of Hygrophila difformis. The result shows that the leaf buds can change with the environmental factors. The leaf morphology is gradually transformed between ovate and dissected leaves. The stem length of the plant is significant difference among three seasons (p＜0.05), when it grew in summer has the shortest stem length (p＜0.01). There are many morphological and anatomical differences of these two leaves types. The heterophylly difference index of ovate leaves is 0.92 and the dissected leaves is 0.08. The ovate leaves are thick (p＜0.01) and have more chloroplasts in cell (p＜0.01), more stomas of leaves (p＜0.01), and more gland hairs on their surface. The dissected leaves do not have any gland hairs on their surface. The maximum electronic transport rate (ETRmax) of the ovate leaves (21.99±1.72) is higher than the dissected leaves.
From the various effects of water level, light intensity, temperatures, and photoperiods show various different results. We found that full water level can cause huge differences of the quantity of new-born leaves, their total surface, stem lengths and the index of heterophylly difference. In light intensity experiment, no difference is found on the stem length, and there is significant difference in the other three characteristics (p＜0.01). In temperature experiment, there is significantly different on the growth of Hygrophila difformis (p＜0.01). The worst growth of this plant is at 15℃ and the best at 30℃, and it has limit growth at 35℃. The dissected leaves is more distinguished with the temperature increasing (p＜0.01). In photoperiods experiment, no difference is found on the stem length, and there is significant difference in the other three characteristics (p＜0.01). The dissected leaves is more distinguished with the longer of the light photoperiod (p＜0.01). The index of heterophylly difference has a largest value of 0.24 between 15℃ and 35℃, and a smallest value of 0.02 between normal light (10 µmole m-2S-1) and intensity light (20 µmole m-2S-1).

KEY WORDS: Hygrophila difformis, heterophylly, index of heterophylly difference, maximum electronic transport rate

目錄
謝辭 i
摘要 ii
Abstract iv
目次 vi
表目次 viii
圖目次 x
壹、前言 1
貳、文獻探討 4
一、葉片形狀的界定 4
二、異型葉產生之研究 4
三、異型葉光合作用效率研究 5
參、材料及方法 7
一、實驗材料 7
二、植物體器官構造之觀察比較與測量 7
三、環境因子對異葉水蓑衣生長影響之探討 12
四、異型葉光合作用速率之比較 13
肆、結果 16
一、異葉水蓑衣植株因季節所發生型態上的變化 16
二、異葉水蓑衣莖、葉因季節所發生的變化 18
(一) 節間長度的變化 18
(二)卵型葉與羽狀葉異化指數的觀察與比較 19
(三) 同一枝條卵型葉、羽狀葉葉型轉變的葉面積分析 21
三、植物體器官構造之觀察比較 23
(一)卵形葉與羽狀葉葉片厚度的比較 23
(二)卵形葉與羽狀葉葉肉細胞中葉綠體數的比較 24
(三) 水上莖與水下莖，莖內皮層所佔莖橫截面積的比例 25
(四) 葉片上下表皮氣孔數目的測量 26
(五) 葉片上下表皮氣孔面積的測量 29
(六) 葉片上下表皮腺毛數目的測量 31
四、環境因子對植株生長與異化指數變量的影響 33
(一) 水位對植株生長與異化指數變量化的影響 33
(二) 光強度對植株生長與異化指數變量的影響 35
(三) 溫度對植株生長與異化指數變量的影響 38
(四) 光週期對植株生長與異化指數變量的影響 44
五 、Diving-PAM 測定卵形葉與羽狀葉光合作用速率 50
伍、討論 51
陸、結論 56
柒、參考文獻 59
表目次
表4-1：不同生長季節節間長度之T檢定值 19
表4-2：卵形葉與羽狀葉，上下表皮單位面積氣孔數目之T檢定值 28
表4-3：卵形葉與羽狀葉，葉片上下表皮氣孔面積之T檢定值 31
表4-4：五種溫度下新生葉片數之單因子變異數分析 38
表4-5：五種溫度下新生葉片數平均差異Post Hoc檢定之Tukey HSD
多重比較 39
表4-6：五種溫度下新生葉片總面積之單因子變異數分析 40
表4-7：五種溫度下新生葉片總面積平均差異Post Hoc檢定之Tukey
HSD多重比較 40
表4-8：五種溫度下植株生長總長度之單因子變異數分析 41
表4-9：五種溫度下植株生長總長度平均差異Post Hoc檢定之Tukey
HSD多重比較 42
表4-10：五種溫度下植株葉片異化指數變量之單因子變異數分析 43
表4-11：五種溫度下植株葉片異化指數變量平均差異Post Hoc檢定
之Tukey HSD多重比較 43
表4-12：不同光週期下新生葉片數之單因子變異數分析 44
表4-13：不同光週期下新生葉片數平均差異Post Hoc檢定之Tukey
HSD多重比較 45
表4-14：不同光週期下新生葉片總面積之單因子變異數分析 46
表4-15：不同光週期下新生葉片總面積平均差異Post Hoc檢定之
Tukey HSD多重比較 46
表4-16：不同光週期下植株生長總長度之單因子變異數分析 47
表4-17：不同光週期下植株生長總長度平均差異Post Hoc檢定之
Tukey HSD多重比較 48
表4-18：不同光週期下植株葉片異化指數變量之單因子變異數分析 49
表4-19 ：不同光週期下植株葉片異化指數變量平均差異Post Hoc
檢定之Tukey HSD多重比較 49

圖目次
圖3-1：複式顯微鏡(40X)下載物台測微器(1mm) 9
圖3-2：製作已知面積的標準尺規(0.25mm2) 9
圖3-3：原圖及轉換測量區域 10
圖3-4：電腦程式操作版面 11
圖3-5：異型葉化指數a / b量測位置圖 12
圖4-1：一年中異葉水蓑衣的型態變化 17
圖4-2：異葉水蓑衣的生殖構造 18
圖4-3：不同季節節間長度變化 19
圖4-4：卵形葉轉為羽狀葉之葉型及異化指數變化 20
圖4-5：羽狀葉轉為卵形葉之葉型及異化指數變化 20
圖4-6：卵型葉與羽狀葉異化指數之比較 21
圖4-7：羽狀葉轉卵形葉，葉片面積之比較 22
圖4-8：卵形葉轉羽狀葉，葉片面積之比較 22
圖4-9：卵形葉與羽狀葉之橫切圖 23
圖4-10：卵形葉與羽狀葉，葉片中肋及厚度 24
圖4-11：卵形葉與羽狀葉葉肉細胞中的葉綠體排列圖 25
圖4-12：卵形葉與羽狀葉，葉肉細胞中葉綠體個數 25
圖4-13：水上莖與水下莖的橫切圖 26
圖4-14：水上莖與水下莖之皮層所佔莖百分比 26
圖4-15：卵形葉與羽狀葉的葉片表皮氣孔分布圖 27
圖4-16：卵形葉與羽狀葉，上下表皮單位面積氣孔數目 28
圖4-17：卵形葉與羽狀葉表皮單位面積氣孔數比值(下/上) 29
圖4-18：卵形葉與羽狀葉，葉片上下表皮之單一氣孔圖形 30
圖4-19：卵形葉與羽狀葉，上下表皮氣孔面積 30
圖4-20：卵形葉與羽狀葉，葉片上下表皮腺毛著生圖 32
圖4-21：卵形葉與羽狀葉，葉片上下表皮單位面積腺毛數目 32
圖4-22：不同水位下植株新生葉片數 33
圖4-23：不同水位下植株新生葉片總面積 34
圖4-24：不同水位下植株生長總長度 34
圖4-25：不同水位下植株葉片異化指數變量 35
圖4-26：不同光強度下植株新生葉片數 35
圖4-27：不同光強度下植株新生葉片總面積 36
圖4-28：不同光強度下植株生長總長度 37
圖4-29：不同光強度下植株葉片異化指數變量 37
圖4-30：五種溫度下植株新生葉片數 38
圖4-31：五種溫度下植株新生葉片總面積 40
圖4-32：五種溫度下植株生長總長度 41
圖4-33：五種溫度下植株葉片異化指數變量 43
圖4-34：五種光週期下植株新生葉片數 44
圖4-35：五種光週期下植株新生葉片總面積 46
圖4-36：五種光週期下植株生長總長度 47
圖4-37：五種光週期下植株葉片異化指數變量 49
圖4-38：卵形葉與羽狀葉，光合作用速率比較 50

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