Please use this identifier to cite or link to this item: http://hdl.handle.net/11455/92178
標題: 紅樹科紅樹林植物托葉、指狀腺體與木栓疣適應紅樹林之研究
Associations between Stipules, Colleters and Cork warts of Mangrove Rhizophoraceae and Adaptation to the Mangrove Environment
作者: Szu-Yang Wu
巫思揚
關鍵字: 紅樹科
水筆仔
紅茄苳
五梨跤
托葉
指狀腺體
木栓疣
葉特徵
解剖構造
下皮層
氣相層析質譜儀
烷類
Rhizophoraceae
Kandelia obovate
Bruguiera gymnorhiza
Rhizophora stylosa,stipule
colleter
cork wart
leaf trait
anatomy
hypodermis
GC/MS
alkanes
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摘要: Mangroves are the characteristic intertidal plant formations including, various species and families found along the sheltered tropical and subtropical coastlines. It is still not clear whether some particular leaf characters of mangrove Rhizophoraceae can help these plants adapting the mangrove environment. In this study, three species of mangrove Rhizophoraceae, Bruguiera gymnorhiza, Kandelia obovata and Rhizophora stylosa, were selected fort functional experiments and anatomical studies of stipules, colleters and cork warts using both plants naturally growing in the field and cultivated in pots with different salinies. Histochemical staining and GC/MS were used to analyze the composition of the exudates of colleters of K. obovata. The aims of this study were to understand the functional effects of stipules, colleters and cork warts to reveal the ways in which these structures are adaptation in the mangrove environment. In an experiment on K. obovata, it is confirmed that stipules provide physical protection to the young leaves, and that colleters located at the adaxial bases of stipules possibly provide protection for young leaves. The exudates of colleters coating the surface of young leaves may help reducing water transpiration, herbivore and pathogen attack. Newly developed leaves subdected to stipule removal developed into smaller and deformed mature leaves with obvious damage. Several leaf traits, including leaf area, fresh weight, petiole length, and stoma density, for leaves treated in this way were significantly lower than those of untreated leaves, protected by stipules during their development. The thickness of sponge tissue of the leaves with stipule removal, however, was significantly higher than that of untreated leaves. The colleters of all studied species are of standard type, which is comprised of central parenchyma cells and outer palisade-like parenchyma cells with secretory function. Only in K. obovata, a few of colleters were found to have independent tracheary elements at the stalk of colleter. This suggests that K. obovata might be a more derived taxon of this subfamily. Histochemical staining revealed that exudates of colleters of K. obovata contain lipid and starch. The results of gas chromatography mass showed that the exudates contain several alkanes. Unexpectedly, I found that the characteristics of the stipule, colleter and cork wart of studied mangroves are vared in winter and summer, but not by different salinity treatment. The colleter length is longer in winter than in summer. Moreover, cork wart density of R. stylosa is significantly higher in winter than in summer, regardless of salinity treatment. It is possibly that the winter leaves of R. stylosa increase the number of cork warts to maintain a better amount of gas exchange. The leaves of R.stylosa and K. obovata showed different responses to salinity: the thickness of hypodermis of R. stylosa increased in the environment with higher salinity, while no detectable was found anatomical change in the leaf of K. obovata. The results suggest that hypodermis of R. stylosa is a water storage tissue with ability to buffer salinity, and K. obovata may have particular physiological regulation of its root system to cope with salt. This study revealed that the characteristics of stipules, colleters and cork warts differed between summer and winter. Such seasonal variation may help these mangroves coping with lower temperature and strong monsoon in winter in Taiwan. Further study is needed to better understanding of detail mechanisms.
紅樹林為天然生長於熱帶與亞熱帶河口與潮間帶地區的植物,主要由不同科之喬木及灌木所構成之群落。而出現在紅樹科紅樹林植物葉片上的多種特殊構造是否有助於適應特殊的紅樹林環境仍然有待研究。因此本研究以紅樹科的三種紅樹林植物紅茄苳(Brugriera gymnorhiza)、水筆仔(Kandelia obovata)與五梨跤(Rhizophora stylosa)長在野外及栽植之植物為材料,進行托葉(stipule)、指狀腺體(colleter)及木栓疣(cork wart)的功能性試驗與解剖構造觀察,並使用組織化學及氣相層析質譜儀分析水筆仔指狀腺體分泌物成份。期以瞭解指狀腺體與木栓疣之解剖構造,並探討托葉、指狀腺體及木栓疣與適應紅樹林環境之可能關聯。 水筆仔在不同程度去除托葉的試驗中,結果明顯證實托葉能提供幼葉物理性的保護,而位托葉基部近軸面的指狀腺體可能提供協同保護的功能,其分泌物可能有助降低葉片的水分蒸散並利用其黏性來防止植食性昆蟲的取食。在未受托葉保護的處理組之新生葉片,其葉片較小且出現許多傷痕及扭曲變形等生長不良的狀況,包括葉面積、鮮重、葉柄長度、氣孔密度及海綿組織厚度等多項葉部特性均與具托葉保護的對照組呈顯著差異。所觀察的三種紅樹林植物之指狀腺體皆為標準型,由與指狀腺體長軸平行之中軸薄壁細胞及與中軸呈垂直排列,具分泌功能的柵狀薄壁細胞所構成。少數水筆仔指狀腺體稈的位置可發現獨立存在之管胞,但五梨跤與紅茄苳則無,顯示水筆仔屬可能具較特殊的演化位置。水筆仔指狀腺體分泌物中經組織化學測定出含有脂類與澱粉,並進一步透過氣相層析質譜儀分析出含有數種烷類。托葉、指狀腺體及木栓疣的特徵明顯受到季節的影響而有所差異,而不同鹽度之差別並不大。此三物種冬季的指狀腺體長度都明顯高於夏季,顯示指狀腺體可能具有幫助紅樹科紅樹林植物適應季節變化的功用。生長於不同鹽度樣點的五梨跤族群在冬季均有較高木栓疣密度,可能與植物在冬季較低溫時,藉由提高木栓疣密度,以維持植物體內運送至根尖分生組織的氣體量有關。五梨跤和水筆仔葉片在不同鹽度處理下反應有所不同。五梨跤葉片會透過改變下皮層厚度的方式來適應不同鹽度的環境,顯示下皮層為葉片儲存水分的構造且具有緩衝鹽分的能力。水筆仔葉片中未發現特別的組織變化,因此生理的調節機制可能是較為重要。本研究證實紅樹科紅樹林物種的托葉、指狀腺體及木栓疣等特徵會隨冬、夏兩季節不同而改變 可能有助於其適應亞熱帶地區紅樹林環境冬季的強風與低溫,然而詳細的適應機制未來仍需要更多的研究。
URI: http://hdl.handle.net/11455/92178
文章公開時間: 2015-07-15
Appears in Collections:生命科學系所

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