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  • 學位論文

沿海拔高度植物葉部形態及穩定同位素變化趨勢

Variations in leaf morphology and stable isotope ratio of widely-distributed plant species along altitudinal gradients

指導教授 : 袁孝維
共同指導教授 : 高文媛(Wen-Yuan Kao)

摘要


台灣擁有許多廣布種植物,從低海拔到高海拔地區、從潮濕到乾旱環境皆有分布。為因應不同海拔高度的環境變化,這些植物的外觀形態與生理反應是否有所不同?這些改變是否有助於其適應不同的環境?此為本研究探討之重點。 本研究藉由野外調查,採集生長於不同環境條件下的廣布種植物--臺灣二葉松(Pinus taiwanensis)、台灣赤楊(Alnus japonica)、五節芒(Miscanthus floridulus)、車前草(Plantago asiatica)--的健康葉片,帶回實驗室分析。採集樣點位於臺灣的中部橫貫公路(台十四線、台十四甲線、台八線),以武嶺為分界,分成東、西部。東部的總海拔高度差約為3000 m,西部約為2500 m,相鄰兩樣點間之海拔高度差為300-500 m。本研究分成三個部份:(1) 比較四種試驗植物在不同海拔樣點之穩定碳、氮同位素比值(δ13C、δ15N)與碳、氮含量百分比(C%、N%);(2) 比較不同月份五節芒與車前草在不同海拔高度其生理、形態及δ13C值之差異;(3) 探討有毛型與無毛型的車前草在地理分布及生理、形態上是否有所差異。 第一部份的實驗於2009年11月進行採樣,根據δ13C值可判斷四種植物只有芒草是C4植物,其他三種皆為C3植物。車前草是三種C3植物中δ13C值最低者,顯示其長期水分使用效率低於台灣赤楊及台灣二葉松,或是其所使用的碳源本身的δ13C值原本就比較低。台灣赤楊的δ15N值明顯低於其他物種,可能是受到其共生根瘤菌之影響。四種植物的各項測值與海拔高度間之均無顯著之線性相關。 第二部份的實驗於2009年11月和2010年7月進行採樣,分析後發現:在7月時,無論東、西部的車前草族群其葉片之δ13C值均隨海拔高度上升而增加,而在高海拔地區的車前草族群,7月所採集之葉片其δ13C值高於在11月時所採集的葉片,其原因可能是7月時葉片的單位葉面積乾重(LMA)增加所造成;因車前草的LMA在7月較高,可能使CO2不易通過葉表皮導致其δ13C值升高。無論東部或西部的芒草族群,芒草的Narea均與隨海拔高度上升而增加,推測是為了增加光合作用之羧化反應。而芒草的δ13C與LMA呈顯著正相關,顯示LMA是影響芒草δ13C的主要原因。 第三部份的結果顯示:除了白柵鐵門樣點外,其他各海拔高度的樣點上均有有毛型、無毛型及邊緣有毛型之車前草植株。比較無毛型及有毛型之車前草葉片其葉綠素含量、LMA、總氣孔密度、δ13C、Narea及Carea,發現均無差異。由結果推測表皮毛對於減少水分散失並無顯著功能,其功能有待進一步研究。

並列摘要


Widely-distributed plants might have mechanisms enabling them to survive in a wide range of climatic and environmental conditions. The purpose of this research was to find out whether the morphological and physiological characteristics of four plant species change with the altitudinal environments and seasons, and what are the significance of these changes to the plants. I collected healthy leaves of populations of four native widely-distributed plants along altitudinal gradients on the eastern side and western side of Central Taiwan. The four species were: Pinus taiwanensis, Alnus japonica, Miscanthus floridulus, and Plantago asiatica. The elevation intervals between adjacent sample sites were around 300-500 m. I collected leaves in Nov. of 2009, and Jul. of 2010. This study include three parts: (1) To compare leaf stable carbon (δ13C) and nitrogen isotope ratio (δ15N), and carbon (C%), and nitrongen (N%) content of populations of the four species in different altitudes. (2) To compare δ13C, δ15N, carbon (Carea) and nitrogen (Narea) content per leaf area and morphological characteristics, including leaf mass per area (LMA) and stomatal density of M. floridulus and Pla. asiatica along altitudinal gradient between two different seasons. (3) To compare the geographical distribution, and morphological traits between Pla. asiatica populations with glabrous and pubescent leaf form. According to the δ13C analysis, M. floridulus is a C4 plant, the other three species are C3 plants. The results of the first part also showed that the δ15N of A. japonica and the δ13C of M. floridulus had positive correlations with altitudes. Among three C3 species, Pla. asiatica had the lowest δ13C, indicating its long-term water use efficiency was lower than that of A. japonica and Pin. taiwanensis, or the δ13C of its carbon sources were lower. The δ15N values of A. japonica were significantly more negative than other three species, I speculated it is caused by the impact of its symbiotic rhizobium. Results of the second part revealed that leaves of Pla. asiatica populations at high altitudes had significantly higher δ13C in July than in Nov.. And a positive correlation between δ13C and altitudes was found in samples collected in July 2010. The main reason why Pla. asiatica had higher δ13C in July 2010 and in higher altitudes might due to its increases in LMA. The LMA of Pla. asiatica in July was higher than that in November, indicating that it was harder for CO2 to diffuse through the leaves in July. A positive correlation between leaf δ13C and altitude was found in western population of M. floridulus, Narea of M. floridulus also showed a generally positive correlation with altitude while LMA didn’t, it suggested that the increase of Narea was not caused by LMA , but due to more allocation of N into photosynthetic machinery. A significant correlation between δ13C and LMA was found, which suggests that LMA might be the main reason causing the variation of δ13C in M. floridulus. The results of the third part showed that Pla. asiatica individuals with glabrous, pubescent, and marginal pubescent distributed in all sample sites except for the site White Iron Gate. Further analysis revealed that there were no significant differences between glabrous and pubescent leaves in chlorophyll content, LMA, SD, δ13C、Narea and Carea.According to the result, the appearance of trichomes may have no significant effects on reducing water loss. Further study is needed to understand the functions of trichomes on leaves of Pla. asiatica.

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