莎草科是世界開花植物中的大科之一，許多物種生長於溼地水生環境，其光合作用包含C3、C4路徑型式，甚至擁有C3與C4路徑間的中間型。多樣性的光合作用型式在莎草科植物解剖、生態、生理與演化上的相關性及意義均值得探討。本研究以葉部或桿解剖及穩定碳同位素確認台灣產莎草亞科的光合作用類型。應用ArcGIS配合Excel將氣候資料與不同光合作用類型、族的物種比率及物種數行簡單相關與迴歸分析，並繪製主題圖。此外，亦調查莎草亞科物種的棲地環境。
結果顯示共49種5亞種9變種為C4路徑，22種4亞種2變種為C3路徑，11種未檢驗。計算出台灣產莎草科C4物種比率占35.2%，在世界其他地區莎草科光合作用路徑的研究中，C4物種比率較高，因台灣位處副熱帶、熱帶氣候區。相關與迴歸分析的結果C4物種比率與溫度因子的相關程度皆較雨量因子高，且有正因果關係，雨量因子則無。在海拔高度與C4比率並無因果關係，應受因變數n值小影響。但在C3與C4物種數上，皆與海拔高度為高度負因果關係，顯示無論光合作用類型為何，莎草亞科物種為熱帶、副熱帶親合。在族比率的分析中，飄拂草族迴歸上皆不顯著，而莎草族與溫度因子有正因果關係，莞族有負因果關係，但程度皆較C4比率為小。因此莎草亞科光合作用類型受氣候因子的影響較分類群大。海拔高度則與莎草族有負因果關係，符合前人研究。
莎草亞科棲地以溼地水生為主(68.5%)，雖C4路徑比率於溼地水生中較低(59.7%)，但中溼環境亦有21.9%，相較禾草與雙子葉C4植物適應於較乾燥環境，水分使用效率似乎並非莎草科演化出C4類型的關鍵。另一方面，C3路徑適應於濕地水生棲地(88.9%)，除說明水分取得對C3路徑的重要性外，可能暗示C4植物由溼地水生的C3路徑演化後，逐漸適應於不同棲地，且分化出不同物種。
本研究確認莎草亞科光合作用類型，並初步探討台灣產莎草亞科與環境因子的關係，未來研究除繼續檢驗薹亞科物種外，應用地理資訊系統結合生態環境資料庫與數學模型更精確揭櫫影響莎草科C3、C4路徑分布的因子，更深入探討整個莎草科各類光合作用型與生態上的意義。

Cyperaceae is the large flowering plant family in the world. There are many species growing in wetland and aquatic areas, and their photosynthesis include C3 and C4 pathways, and even have C3-C4 intermediates. It is worth to explore the diversity of the photosynthetic mechanisms of Cyperaceae in plant anatomy, ecology, physiology and evolution of their relation and significance. In this study, photosynthetic pathways of Cyperoideae in Taiwan were recognized by leaves or culms anatomy and stable carbon isotope ratios. Applying ArcGIS and Excel softwares, simple correlation and linear regression analysis were used to investigate the correlations of climatic data and latitude with species percentages, species numbers of different photosynthetic pathways and 3 tribes. All these correlations are displayed by the theme maps. In addition, we also investigated the habitats of Cyperoideae.
The results showed that 49 species, 5 subspecies and 9 varieties displayed C4 pathway, and 22 species, 4 subspecies and 2 varieties belonged to C3 pathway. The C4 percentage was 35.2% of Cyperaceae in Taiwan. It was higher when compared with other regional studies of photosynthetic pathways of Cyperaceae in the world, because Taiwan is located in the subtropical and tropical climate districts. The results of correlation and regression showed that the correlations of the C4 percentage with temperature factors are higher than rainfall factors. It was positively correlated with temperature factors, but not with the rainfall factors. The C4 percentage was not correlated with altitude, due to the small number of independent variable. However, the numbers of C3 and C4 were strongly and negatively correlated with altitude, showing that no matter what types of photosynthesis, Cyperoideae possess tropical and subtropical affinities. The analysis of tribe percentage, Abildgaardieae showed no correlated with any environmental factors, Cypereae was positively correlated with temperature factors, but Scirpeace was negatively correlated. Their coefficients of regression were smaller than that of C4 percentage. Therefore, the climatic factors are much to do with photosynthetic pathways than taxa. And Cypereace is negatively correlated with altitude in line with previous studies.
Most species in Cyperoideae were found in wet and aquatic areas (68.5%). The percentage of C4 plants was lower in wet and aquatic areas (59.7%), but there was 21.9% in mesic areas, compared with the C4 grasses and dicots that tended to be more abundance in arid areas. It appears that water use efficiency is not the key point of C4 evolution in Cyperaceae. On the other hand, C3 pathway adapted to wet and aquatic areas (88.9%). It may explain that the water availability is crucial and suggests C4 plants evolved from C3 pathway in wet and aquatic areas, then gradually have been adapting to different habitats, and differentiation of species.
This research recognizes photosynthetic pathways of Cyperoideae, and preliminary investigates the correlations between Cyperoideae and the environmental factors in Taiwan. More information still need further survey especially the Caricoideae species, and the application of Geographic Information System to establish ecological environmental databases and mathematical models for understanding the effects of environmental factors on the distribution of C3 and C4 pathways.

摘要……………………………………………………………………………Ⅰ
英文摘要………………………………………………………………………III
致謝……………………………………………………………………………V
表目錄…………………………………………………………………………VIII
圖目錄…………………………………………………………………………IX
壹、前言………………………………………………………………………1
貳、文獻回顧…………………………………………………………………3
一、莎草科與光合作用研究簡介………………………………………3
二、莎草科的光合作用研究……………………………………………6
三、穩定碳同位素比值與光合作用……………………………………10
四、物種分布與環境因子……………………………………………………11
参、材料與方法………………………………………………………………13
一、材料蒐集…………………………………………………………………13
二、實驗方法…………………………………………………………………13
肆、結果………………………………………………………………………18
一、葉部/桿解剖 ………………………………………………………18
二、穩定碳同位素比值………………………………………………………18
三、莎草族小穗排列…………………………………………………………21
四、環境因子…………………………………………………………………21
五、棲地類型與物種分布……………………………………………………30
伍、討論………………………………………………………………………32
一、光合作用路徑判定………………………………………………………32
二、環境因子分析……………………………………………………………34
三、棲地環境…………………………………………………………………42
四、檢討與建議………………………………………………………………43
陸、結論………………………………………………………………………45
柒、參考文獻…………………………………………………………………47

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