臺灣博碩士論文加值系統

雜草管理是蔬菜生產體系相當重要的一環。一般農夫普遍採用的管理策略包括耕犁、刈草、覆蓋及施用除草劑等。其中又以除草劑的使用最為普遍。然而，研究指出過量使用除草劑及具除草劑抗性雜草的出現，已經造成許多環境及生態的問題。有鑑於此，部分科學家致力於從化感植物中找到新的天然植毒性物質，開發新型除草劑；或是致力於將化感植物或作物，作為覆蓋植物，導入輪作或間作的系統中。咸豐草(Bidens pilosa L.)是兼具傳統可食的野菜及中草藥的化感植物，具有導入蔬菜生產系統，提升雜草防治功效的潛力。在臺灣，咸豐草共計有白花鬼針(B. pilosa L. var. pilosa)、小白花鬼針(B. pilosa L. var. minor (Blume) Sherff)及大花咸豐草(B. pilosa L. var. radiata Sch. Bip.)等3個變種。其中以大花咸豐草入侵性最強，但針對此物種的發芽光需求、自毒作用、化感物質釋出途徑、化感作用識別以及在蔬菜生產系統中的雜草控制潛力的相關研究甚少。
本研究首先進行實驗室、盆栽及田間試驗，用以評估不同pH、光環境及殘體植毒性對大花咸豐草的發芽影響，了解未來應用上如何防止該物種的衍生危害。其次，透過自田間蒐集大花咸豐草植群天然淋溶液、利用吸附劑吸收葉、莖及葉等不同部位揮發性化合物、搜集根部淋洗液及施用殘體或其水萃液試驗方法等，探討此物種之化感物質可能的釋出途徑、種類及其影響。為了評估大花咸豐草化感作用對其他植物的影響程度，以香附子(Cyperus rotundus L.)為測試植物，進行大花咸豐草殘體植毒性對香附子種間密度依賴(density-dependent)、兩物種種間競爭和大花咸豐草田間土壤與其殘體對香附子塊莖更新之影響等試驗。最後，本研究也以小白菜(Brassica rapa L. subsp. chinensis (L.) Makino)為材料，在盆栽中施用大花咸豐草殘體，探討其化感物質對小白菜生長期間的雜草防治效果，以及對小白菜生長是否會產生不良影響。
試驗結果顯示，大花咸豐草種子最適發芽pH為5-7之間，其中以胚根生長對不同pH處理較為敏感，強酸性或偏鹼性環境對其幼苗之胚根發育有明顯負面影響。光照對大花咸豐草發芽影響隨環境而異，培養皿的試驗結果顯示，發芽並無光需求，但在以土壤為介質的盆栽則有明顯照光促進發芽的現象。遠紅光處理雖可抑制種子發芽，但預處理後於黑暗中培育仍可發芽的現象，顯示種子的發芽光反應應是極低光量反應(very low fluence response, VLFR)。試驗同時發現，大花咸豐草殘體與水萃液在控制(實驗室)、半自然(溫室或生長箱盆栽)及自然(田間)狀態下均可對其種子表現出植毒性。由遠紅光預處理後於黑暗中培育的發芽受抑現象推測，殘體或水萃液中所含的酚類化感物質可能會破壞Pfr光敏素，進而抑制種子的發芽率。在化感物質釋出途徑方面，試驗結果顯示大花咸豐草雖可透過淋溶、揮發、根分泌及殘體分解等途徑釋出植毒性物質，但以殘體分解為最重要的途徑。同時，由各途徑的化感物質分析得知，淋溶或殘體分解途徑的主要化感物質應為酚類，而揮發途徑則以-蒎烯(-pinene)、莰烯(camphene)、月桂油烯(myrcene)、檸烯(limonene)及石竹烯(caryophyllene)等萜類及苯基庚三炔(phenylheptatriyne)等多炔類(polyacetylene)化感物質為主。除此之外，植毒性-密度依賴試驗結果發現，施用大花咸豐草殘體可明顯改變香附子之單株乾重-栽培密度對數關係斜率。亦即殘體對香附子生長的抑制強度隨施用量遞增，但隨栽培密度增加而遞減，顯示殘體植毒性明顯存在低密度處理，但在高密度處理中則被稀釋了。而由大花咸豐草與香附子的種間競爭試驗進一步發現，相較於無施用處理，施用活性碳處理可提高香附子的生長，推測是因為大花咸豐草所釋出的植毒性物質經活性碳吸收，降低對香附子生長的影響，顯示化感作用在大花咸豐草與相鄰植物的競爭中扮演一定程度的影響。最後，在小白菜的栽培試驗中發現，大花咸豐草殘體可以顯著抑制雜草種子發芽達5週，並且即使在施用1週後播種小白菜，亦不會對小白菜的生長造成影響。
大花咸豐草雖是高入侵性的外來植物，但在食用及醫藥方面極具發展潛力。本研究顯示，該植物殘體的化感作用具直接應用在田間防治雜草及導入蔬菜輪作生產系統的潛力。

Weed management is important for vegetable production. The often-used management strategies include plowing, mowing, mulching, and herbicide, and so many; among these strategies, herbicide utilization is the most widely used method. However, research indicated that the increased herbicide utilization and the occurrence of weeds with herbicide resistance have caused many environmental and ecological problems. Due to this reason, some researchers devoted themselves to exploring new/effective natural phytotoxic compounds from allelopathic plants while some aimed to introduce allelopathic plants/crops as the cover crop in the rotation or intercropping system. Bidens pilosa L. has been identified as an edible and medicinal herb with allelopathy, possessing the potential to enhance weed control in the vegetable production system. Three varieties, i.e. B. pilosa var. pilosa, B. pilosa var. minor, and B. pilosa var. radiata (BPr), were found in Taiwan. Among them, BPr was known as the most invasive one but few studies were conducted on its germination light requirement, autotoxicity, allelopathy, and the weed control potential in the vegetable production system.
In this study, experiments at the scales of laboratory, pot, and field were conducted to evaluate the effects of pH, light conditions, and residue phytotoxicity on BPr germination. To evaluate the possible allelopathic pathways of BPr, natural leachates, volatile compounds, root exudates, and residues of this species were collected and examined in the laboratory, pot, and field according. In addition, to understand the effects of BPr allelopathy on the neighbor plants, Cyperus rotundus L. (CR) was selected as the target plant for the density-dependent experiments, interspecies competition experiments, and several field residue/mulch application experiments. Finally, the effects of BPr residue on the weed control and growth of Chinese mustard (Brassica campestris L. ssp. chinensis (L.) Makino) were also evaluated.
The results showed that the suitable pH for BPr germination was 5 - 7, and the radicle growth was more sensitive to pH than germination. A strong acid or alkali environment had significant inhibition on radicle growth. The influence of light on BPr germination varied in different conditions, i.e., no light requirement was observed when incubated in the Petri dishes while germination was stimulated under the light when incubated in pots. Besides, although far-red light almost completely inhibited BPr germination, seeds germinated when incubated in the darkness after far-red light pretreatment. This phenomenon indicated that the light response of BPr seed germination might be a very low fluence response (VLFR). The results also demonstrated that the BPr residues and their water extracts exhibited phytotoxicity to its seeds in the experiments of controlled (laboratory), semi-natural (pots in the greenhouse or growth chamber), and natural (field) conditions. Since the germination inhibition was found in the dark incubation after far-red light pretreatment, it was suggested that the phenolics contained in the residues or water extracts might destroy the Pfr phytochrome and further inhibited the seed germination. For the research of allelochemical releasing pathways, the results illustrated that BPr could release phytotoxic substances through leaching, volatilization, root exudation, and residue decomposition; however, residue decomposition was supposed the most important pathway. According to the results of compound analysis, the main allelochemicals in leaching and residue decomposition pathways were phenolics while the terpenoids such as -pinene, camphene, myrcene, limonene, and caryophyllene, and the polyacetylene, such as phenylheptatriyne, were the main allelochemicals in volatilization pathway. In addition, the results from the density-dependent phytotoxicity experiments showed that BPr residue could alter the slope of the logarithmic relationship between dry weight per plant and the density of CR. In other words, the inhibition of the residue phytotoxicity on CR growth increased with application rate but decreased with increasing CR density. The phytotoxicity was exhibited in the low-density treatments but diluted in the high-density treatments. In the interspecies competition between BPr and CR, it was found that CR had greater growth when applied with activated carbon than that without activated carbon. It was presumed that the phytotoxins released from BPr were adsorbed by the activated carbon, and hence reduced the inhibition on the CR. Lastly, in the Chinese mustard growth experiments, it was demonstrated that the BPr residue could significantly inhibit the weed germination for five weeks while no growth reduction was found in the Chinese mustard even sowed one week after residue application.
Although BPr is a highly invasive species, it exhibits great development potential in edible and herbal medicinal purposes. In this study, it was found that the BPr residues were capable of being used for controlling weeds in the field and being introduced to the vegetable rotation systems.

口試委員會審定書 I
誌 謝 II
摘 要 IV
Abstract VI
目 錄 IX
表目錄 XV
圖目錄 XVII
附表目錄 XXII
第壹章 前言 1
第貳章 文獻回顧 6
壹、 植物化感作用(Allelopathy) 6
一、 植物化感作用的研究歷史概述 6
二、 植物化感作用的定義 12
貳、 化感物質(Allelochemical) 13
一、 酚類化感物質 13
二、 萜類化感物質 16
三、 含氮化感物質 17
四、 多炔類化感物質 19
參、 化感物質釋出途徑 20
一、 淋溶(Leaching) 21
二、 揮發(Volatilization) 23
三、 根部分泌(Root exudation) 24
四、 殘體分解(Residue decomposition) 25
肆、 化感作用試驗方法 26
一、 實驗室試驗(控制環境) 26
(一) 培養皿 27
(二) 三明治法 27
(三) 植物盒法 27
(四) 海綿法 28
二、 生長箱或溫網室內的盆栽試驗(半自然環境) 28
(一) 評估添加化感物質或殘體對受測植物的影響 29
(二) 評估化感植物對受測植物之影響 29
1. 簡單成對混植法 29
2. 添加試驗設計 30
3. 取代試驗設計 31
4. 密度依賴試驗 31
三、 田間試驗(自然環境) 33
(一) 區別化感作用的田間試驗 33
(二) 評估如何將化感植物或其殘體(含水萃液)應用於在農業生產系統 34
1. 輪作制度 34
2. 間作制度 36
3. 殘體與其水萃液 36
伍、 影響雜草種子發芽的因子 37
一、 溫度 38
二、 光環境 38
三、 pH 40
四、 化感物質 41
陸、 主要研究植物介紹 41
一、 咸豐草 41
(一) 種類、特性及分佈 41
(二) 發芽特性研究 42
(三) 化感作用研究 44
二、 蔬菜田主要雜草-香附子 44
(一) 特性及分佈 44
(二) 防治研究 45
1. 化學防治 45
2. 非化學防治 46
3. 化感作用防治 47
第參章 材料與方法 48
壹、 供試材料的預備 48
一、 大花咸豐草種子採集與調製 48
二、 大花咸豐草殘體來源與製備 48
三、 香附子塊莖採集 49
四、 供試蔬菜作物種子來源 49
貳、 土壤理化性質分析方法 49
一、 土壤反應(pH值) 49
二、 土壤電導度 49
三、 土壤質地 50
四、 土壤陽離子交換容量 50
五、 土壤礦質元素分析 50
六、 土壤中有效氮之測定 50
七、 土壤有機質含量 50
參、 光環境、酸鹼值及殘體對大花咸豐草繁衍潛力評估 51
一、 試驗一：光度及酸鹼度對大花咸豐草種子發芽之影響 52
(一) 光度及酸鹼度處理 52
(二) 發芽試驗 52
二、 試驗二：不同光因子及化感作用對大花咸豐草繁衍之影響 53
(一) 試驗二L1：不同部位殘體水萃液對大花咸豐草種子發芽與胚根發育之影響 54
1. 不同部位殘體水萃液之製備 54
2. 發芽試驗 54
(二) 試驗二L2：光因子及地上部殘體水萃液對大花咸豐草種子發芽及胚根生長之影響 55
1. 光處理 55
2. 地上部殘體水萃液之製備 56
3. 發芽試驗 56
4. 胚根量測 57
(三) 試驗二P1：光質及地上部殘體對大花咸豐草種子於盆栽中發芽之影響 57
(四) 試驗二P2：遠紅光預處理對大花咸豐草種子於盆栽中之發芽影響 58
(五) 試驗二F1：遠紅光預處理及殘體對大花咸豐草種子在田間發芽之影響 59
肆、 大花咸豐草化感作用評估 60
一、 試驗三：大花咸豐草地上部淋溶途徑之化感作用評估 60
(一) 試驗三L1：天然植群淋溶液之化感作用檢定 60
1. 雨水及植群淋溶液蒐集 60
2. 化感物質(allelochemicals)分析 62
(1) 總多酚(total phenolics)分析 62
(2) 多炔類-苯基庚三炔(phenylheptatriyne)分析 62
A. 地上部植體測定 62
B. 植群淋溶液測定 63
3. 發芽試驗 63
(二) 試驗三L2：不同部位之植體水萃液化感作用檢定 64
1. 不同部位植體淋溶液製備 64
2. 化感物質(allelochemicals)分析 64
3. 發芽試驗 64
二、 試驗四：大花咸豐草不同部位揮發物質之化感作用評估 64
(一) 不同部位之揮發性物質測定 64
(二) 大花咸豐草葉片揮發物質之化感作用檢定 66
三、 試驗五：大花咸豐草地下部化感作用評估 66
(一) 大花咸豐草根部分泌物對香附子塊莖萌芽及幼苗生長之影響 66
四、 試驗六：於半自然及自然環境中區別大花咸豐草化感作用 67
(一) 試驗六P1：密度依賴植毒性試驗 68
(二) 試驗六P2：大花咸豐草與香附子種間競爭試驗 68
(三) 試驗六F1：香附子塊莖於大花咸豐草植群田間之萌芽試驗 69
(四) 試驗六F2：大花咸豐草植群及地上部殘體覆蓋對香附子更新之影響 70
伍、 應用大花咸豐草化感作用於蔬菜生產之評估 71
一、 試驗七：添加大花咸豐草地上部殘體對小白菜於盆栽中生長之影響 71
陸、 資料處理與統計分析 71
一、 變方分析 71
二、 發芽率分析 72
第肆章 結果與討論 73
壹、 大花咸豐草種子田間發芽潛力之評估 73
一、 試驗一：光度及酸鹼度對大花咸豐草種子發芽之影響 73
(一) 大花咸豐草種子發芽率對光度及酸鹼度之反應 73
(二) 胚根胚軸發育 78
二、 試驗二：不同光因子及化感作用對大花咸豐草繁衍之影響 80
(一) 試驗二L1：不同部位殘體水萃液對大花咸豐草種子發芽與胚根發育之影響 80
(二) 試驗二L2：光因子及地上部殘體水萃液對大花咸豐草種子於控制環境中發芽及胚根生長之影響 83
1. 控制環境中種子發芽之光需求與對地上部殘體水萃液處理的反應 83
2. 控制環境中種子發芽對遠紅光預處理的反應 85
3. 控制環境中種子發芽對遠紅光處理之反應 85
4. 控制環境中胚根生長對光因子及地上部殘體水萃液的反應 86
(三) 光質及地上部殘體對種子於半自然環境中發芽之影響 88
1. 試驗二P1：半自然環境中種子發芽對光質與地上部殘體處理的反應 88
(1) 半自然環境中種子之發芽光需求與對地上部殘體處理的反應 88
(2) 試驗二P1：半自然環境中種子發芽對遠紅光處理之反應 91
2. 試驗二P2：半自然環境中遠紅光預處理對種子發芽之影響 92
(四) 試驗二F1：遠紅光預處理及地上部殘體對種子於自然環境中發芽之影響 93
1. 自然環境中種子發芽對遠紅光預處理的反應 93
2. 自然環境中種子發芽對地上部殘體處理的反應 95
貳、 大花咸豐草化感作用之評估 98
一、 大花咸豐草化感物質釋出途徑 98
(一) 大花咸豐草地上部淋溶途徑之化感作用評估 98
1. 試驗三L1：天然植群淋溶液之化感作用檢定 98
(1) 雨水與植群淋溶液酸鹼值與電導度 98
(2) 天然植群淋溶液之酚類化感物質分析 99
(3) 天然植群淋溶液之苯基庚三炔化感物質分析 100
(4) 天然植群淋溶液對萵苣種子發芽之影響 107
2. 試驗三L2：不同部位之植體水萃液化感作用檢定 110
(1) 不同部位植體水萃液之酸鹼值與電導度 110
(2) 不同部位植體水萃液之酚類化感物質分析 110
(3) 不同部位植體水萃液之苯基庚三炔化感物質分析 111
(4) 不同部位植體水萃液對萵苣種子發芽之影響 119
(二) 試驗四：大花咸豐草揮發途徑之化感作用評估 122
1. 不同部位之揮發性物質測定 122
2. 大花咸豐草葉片揮發物質之化感作用檢定 124
(三) 試驗五：大花咸豐草根部分泌途徑之化感作用評估 136
二、 試驗六：大花咸豐草化感作用之區別與地上部殘體分解途徑評估 137
(一) 試驗六P1：由植毒性-密度依賴試驗識別大花咸豐草化感作用 137
1. 香附子植株各部位之生長反應 137
2. 香附子梢根比之反應 139
3. 香附子塊莖增生之反應 139
(二) 試驗六P2：由種間競爭試驗識別大花咸豐草之化感作用 142
1. 不同競爭形式對試驗植株之生長影響 142
2. 不同競爭形式對試驗植株梢根比之影響 147
3. 不同競爭形式對香附子塊莖增生之影響 149
(三) 試驗六F1：香附子塊莖於大花咸豐草植群田間之萌芽試驗 150
(四) 試驗六F2：大花咸豐草植群及其地上部殘體對香附子更新之影響 151
參、 應用大花咸豐草化感作用於蔬菜生產系統 157
一、 試驗七：添加大花咸豐草地上部殘體對小白菜於盆栽中生長之影響 157
第伍章 結論 162
壹、 研究發現 162
貳、 研究貢獻 163
參、 研究限制 163
肆、 未來研究方向 163
參考文獻 165
附 表 187

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