2,327 research outputs found
Bizonoplast a unique chloroplast in the epidermal cells of microphylls in the shade plant Selaginella Erythropus (Selaginellaceae)
No full textCopyright © 2007 Botanical Society of America, Inc.Chiou-Rong Sheue, Vassilios Sarafis, Ruth Kiew, Ho-Yih Liu, Alexandre Salino, Ling-Long Kuo-Huang, Yuen-Po Yang, Chi-Chu Tsai, Chun-Hung Lin, Jean W. H. Yong and Maurice S. B. K
Mongo Breeding in Taiwan
No full text台灣芒果栽培品種，早期皆以引進國外優良品種推廣試種為主，並未進行育種工作。直至1969年起，農業試驗所鳳山分所開始進行自然雜交與實生選拔，高雄區農業改良場亦於1994年開始進行芒果之育種工作。育種目標為選育具早熟或晚熟、不時花、開花著果穩定、中型果、纖維少、果肉厚、不易生理劣變、抗（耐）病及貯運等特性之品種。育種方法多以自然雜交授粉，再從單胚型品種之後裔進行實生選拔。育種程序包括雜交後裔的培育、品系選拔、品系比較試驗、區域試種、植株性狀檢定及品種權申請等。實生品系選拔以進行果實品質分析為初選項目，優選品系除了調查果實性狀之外，並進一步評估產量、耐病性及地方適應性等。目前鳳山分所已於1985年培育出’台農1號’及’台農2號‘等2個種；高雄場亦於2008年育成’高雄3號夏雪‘。
The mongo cultivars in Taiwan were introduced excellent varieties from foreign countries and nobreeding program was conducted in early time. Fengshan Tropical Horticultural Experiment Branch began to carry out selections of open pollination and seeding until 1969. Kaohsiang District Aghcultural Research and Extension Station (Kaohsiang DARES) performed mongo breeding from 1994. The breeding objects were focus on early mature, Iate mature, off-season, stable blooming and frUitbear, middle focus size, less pulp fiber, high pulp rate, non-physiological disorder, disease resistance/tolerance and well storage ability. The breeding method was open pollenated then selected the progenies of mono-embryonic varieties. The breeding processes contain progenies cultivation, line selection, line comparetest, local test, DUS test and variety right application. The preliminary trait of seeding lines selection is focus on fruit quality. Fruit characteristics of selected Iines were the yield, diseases tolerance, and local adaption were evaluated. So far, Fengshan Tropical Horticultural Experiment Branch released 'Tainung No.l' and 'TainungNo. 2' in 1985. Kaohsiang DARES released 'Kaohsiang No.3' (Shia Sheue) in 2008
Laporan Kegiatan: Kunjungan singkat ke hutan bakau di Pulau Mendanau dan Belitung
Full text linkKunjungan singkat ini bertujuan untuk melacak keberadaan Ceriops australis dari suku Rhizophoraceae di Pulau Mendanau dan Belitung. Satu specimen yang berasal dari pulau ini yang diidentifikasi sebagai Ceriops tagal (Bilinton Island: Teijsmann s. n. [1875] (BO)), diidentifikasi ulang oleh C-R Sheue sebagai Ceriops australis (Sheue et al. 2009). Studi ini diperlukan untuk mengetahui fitogeografi Ceriops australis yang umum ditemukan di bagian Timur Indonesia, dan bermanfaat pada upaya konservasi jenis ini. Dari pengamatan di beberapa bagian Pulau Mendanau dan Pulau Belitung, kecuali Belitung bagian Selatan, tidak dijumpai Ceriops australis. Satu jenis Bruguiera cylindrical (Rhizophoraceae) dijumpai di Pulau Mendanau sehingga jumlah jenis bakau di pulau tersebut menjad 19 jenis. Populasi jenis ini jarang dan tersebar. Hutan bakau di Pulau Mendanau masih terjaga baik dengan keragaman substrat termasuk lumpur, pasir, batu, dan batu karang. Pulau ini menjadi lokasi studi yang baik untuk mengamati asosiasi antara keragaman bakau dan lingkungannya. Masyarakat di Pulau Mendanau seyogyanya bangga memiliki hutan bakau yang masih terjaga baik. Sedikit disayangkan dijumpai sampah di hutan bakau di sekitar beberapa kampong nelayan. Pendidikan dan pengelolaan sampah yang baik diharapkan akan menjaga fungsi ekosistem bakau secara optimal
How Shih-chi Was Wr itten to Follow Chun-chiou
No full text《史記》是中國的正史鼻祖、散文大宗，但司馬遷卻自謂其作史接周、孔，「繼春秋」。因此，《史記》與《春秋》的關係，遂變成一個重要、有爭論且難解的老問題。本研究計畫約縮研究範圍，直指問題核心，主要探討司馬遷所說的是哪一部《春秋》？何人所作？以及《史記》究竟如何「繼春秋」？此一問題之探，需要對《史記》與《春秋》同具深入的理解，特別是春秋公羊。Shih-chi has been known as the origin of
Chinese standard histories and a classic of
prose for a long time. But its author Symachian
claimed that the root of what he did
was the spirits of Zhou-gung (周公) and
Confucius, and his book was a follower of
Chun-chiou( 春秋). Therefore the relation
between Shih-chi and Chun-chiou has
constantly been an important open question
without any identical answer. Now in this
plan, only the keys of all difficult problems
are aimed at so that actual progress would be
made. Essentially three questions will be
discussed one after another: Which Chunchiou
is the one to be followed by Shih-chi?
Who is the author of the Chun-chiou? How
was Shih-chi written to follow it? I think a
successful research about these questions will
deeply depend on a comprehension of Shihchi
as well as of Chun-chiou, especially the
interpretations of Chun-chiou from Gungyang(
公羊) school
Leaf Morphology of Selaginella P. Beauv. and its Taxonomic Significance in Taiwan
No full textMicrophyll morphology of 16 Taiwanese species of Selagienlla was observed under light microscopy and scanning electron microscopy. Features studied include microphyll arrangement, epidermal cell morphology, stomal distribution, shape arrangement and distribution silica body on microphyll surface. Microphyll are dimorphic and 4-ranked arrangement, with ventral microphylls larger than the dorsal microphylls. The epidermal cells at dorsal side of dorsal microphylls and at ventral side of ventral microphylls are similar, which are tetragonal or oblong in shape and sinuolate or sinuate in anticlinal cell walls. The epidermis at ventral side of dorsal microphylls and dorsal side of ventral microphylls are similar, which are sub-square or rectangular in shape and straight or sinuolate in anticlinal walls. Stomata are mainly distributed on dorsal side of dorsal microphylls and ventral side of ventral microphylls. Four distribution patterns of are found on microphyll silica body, which are microphyll margin, midrib, homogeneity and nil patterns. Four types of arrangement of silica body on epidermal cells were recognized, namely single row, multi-row, mixed and globulate types. In Taiwan, the characters appeared in epidermal cell morphology and stomal distribution of microphyll are useful for species identification in Selaginella, while those of distribution at patterns and arrangement types of silica bodies on epidermal cells are valuable for the species identification under subgenus
Associations between Stipules, Colleters and Cork warts of Mangrove Rhizophoraceae and Adaptation to the Mangrove Environment
No full textMangroves 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。摘要 ......i
Abstract......ii
中文表目錄 ......vi
中文圖目錄 ......viii
英文表目錄 ......x
英文圖目錄 ......xiii
第一章、前言 ......1
第二章、托葉和指狀腺體形態與水筆仔指狀腺體化學成分分析 ......5
一、前言 ......5
二、材料與方法 ......12
三、結果 ......16
1. 指狀腺體形態 ......16
2. 水筆仔指狀腺體分泌物成分分析 ......27
四、討論 ......32
第三章、水筆仔托葉與指狀腺體功能性試驗 ......36
一、前言 ......36
二、材料方法 ......38
三、結果 ......42
1. 托葉及葉片生長 ......42
2. 葉特徵 ......52
3. 葉片結構......69
四、討論 ......77
第四章、托葉和葉片特徵之鹽度試驗......81
一、前言 ......81
二、材料方法 ......85
三、結果 ......87
1. 葉片結構 ......87
2. 指狀腺體與木栓疣......98
四、討論......115
第五章、引用文獻......11
Toward a Motivation Model of Pragmatics/ Rong Chen.
No full textIn English.With the "discursive turn" has come a distrust - a complete rejection by some - of theories that seek deeper reasons for surface phenomena. Rong Chen argues that this distrust, with its accompanying overemphasis on specificity and fluidity of linguistic meaning and social values, is unwarranted and unhelpful. Drawing on insights from social theories and various strands of pragmatics, he proposes a motivation model of pragmatics (MMP), contending that language use can be adequately, coherently, and elegantly studied via the motivation behind it in its varied and dynamic contexts. The model, with its well-laid out components, is then applied to (im)politeness research, cross-cultural pragmatics, diachronic pragmatics, discourse and genre analysis, conversation analysis, identity construction, and the study of metaphor, sarcasm, parody, and lying. MMP is thus a framework aimed at accounting for fluidity with stable notions, specificity with general principles, and differences with similar underlying factors. As such, the book should appeal to students of pragmatics, (im)politeness, conversation analysis, sociolinguistics, applied linguistics, communication, sociology, and psychology.Frontmatter -- Foreword -- Contents -- List of figures -- List of tables -- Chapter 1 Pragmatics then and now -- Chapter 2 A motivation model of pragmatics (MMP) -- Chapter 3 MMP and (im)politeness -- Chapter 4 MMP and cross-/intercultural variation -- Chapter 5 MMP and diachronic pragmatics -- Chapter 6 MMP and discourse -- Chapter 7 MMP and metaphor -- Chapter 8 MMP and the non-literal -- Afterword -- References -- Appendix -- Subject index -- Author index1 online resource (XIII, 333 p.)
Leaf traits and growth of the Taiwan palm Phoenix hanceana Naudin in different habitats
No full textTaiwan date palm, Phoenix hanceana Naudin (Arecaceae) is a native palm widely distributed in lowland Taiwan. Compared to most palm trees growing in tropical regions, P. hanceana found in subtropical habitats is a chilling tolerant indicator plant of its family. This species can serve as a valuable indicator to study the effects of global warming and changing environments and to provide a better understanding of how these changes affect the future populations of species in Arecacae and their growth strategies. In this study, P. hanceana from seven selected study sites in Taiwan (including western and eastern coastal areas and an outlying island), and cultivated individuals with different treatments, were used to monitor. The aim was to explore the association between growth strategy and leaf functional traits of P. hanceana and compare sapling growth responses. In addition, the tree age modeling and age estimation of local populations of P. hanceana were first reported.
Results showed that the plants growing in eastern Taiwan (Nantian, Guanshan and Baxiandong) had slower leaf growth rates. Among these sites, the plants growing in Baxiandong with relative shaded environments and those in the 21st- 25th plots of Guanshan had higher values of mature leaf length, leaf petiole length, leaf life span, leaf area and specific leaf area (SLA), but smaller leaflet frequency. The plants growing at Tongxiao, Dadu, Ludao (island) and Dulan had higher leaf growth rates. The two western sites (Dadu and Tongxiao) had experienced long term disturbance from human activity and fire, and gave the highest leaf dry matter content and low specific leaf area. Stepwise regression predicted that annual mean temperature, cumulative days of temperature < 12℃ per year, annual max temperature and min temperature were the main factors significantly related to leaf traits. A similar result was found from Canonical Correspondence Analysis (CCA). The seven study sites can be classified as coming from three geographic regions: eastern coastal, Guanshan, and western coastal regions. Among the classified ranks of the plant trunk height, there were no significant differences between leaf traits and trunk heights in western coastal and in Guanshan regions, but the leaf growth rate and leaf area increased with trunk height of the plants growing in eastern coastal region and the 14-16th plots of Guanshan. No matter leaf growth rate or green leaf number of any nursery saplings increase as trunk height increases. Those plants cultivated in eastern Taiwan treated with [sunny + moist] had a higher growth rate and green leaf number; while the plants treated with [sunny + water logging] had the shorter leaf life span than the others. The seasonal growth surveys during 2011 to 2013 showed that the highest leaf growth rate appeared in the summer of 2012, while the slowest one was in the winter of 2012. In summer and in fall of 2012, leaf growth rate was faster than those of the previous year. This study revealed that P. hanceana has different growth strategies in response to different environments and seasonal changes, and these responses may affect the trunk height. Moreover, temperature is the most important factor for palm growth. Light shapes the leaf. Precipitation promotes leaf growth, and the rainy days affect the leaf growth strategies: to increase the leaf number or to elongate leaf length. However, if precipitation is excessive, and the trunk bases are soaked, gas exchange rates are reduced, thus reducing leaf longevity.
Based on the growth modeling of P. hanceana constructed in this study, the young tree will not form a significant trunk until the growth of the 79th leaf. It is estimated that the ages of the existing populations of P. hanceana in Taiwan are at least 94- 273 years. Estimation of the oldest trunk age allows inference of the minimum local population survival time. In summary, these results of growth strategies and plant age composition of this species provide valuable insights for a better understanding of plant adaptive plasticity and for future conservation.台灣海棗（Phoenix hanceana Naudin）為廣泛分布於台灣低海拔的原生棕櫚科（Arecaceae）植物。相對於一般的棕櫚科為熱帶植物，分布至亞熱帶地區的台灣海棗是本科植物可耐臨界低溫的指標物種，對於日趨嚴重的全球暖化及氣候劇烈動盪，值得探討其生長適應及族群延續策略。本研究以監測生長於台灣本島東西岸、離島等七個地區不同生育地的台灣海棗成株，並且加入人工種植的植株，探討其生長策略與葉功能特性的相關性。此外，本研究亦首次建構海棗樹齡的推算模式，並推估各地族群存活或形成的時間。
結果顯示生長在東部南田、關山及八仙洞地區的台灣海棗有顯著較緩慢的葉增加速率；其中較具遮陰的八仙洞及關山第21?25樣區，有顯著較大的成熟葉長度、葉柄長度、葉面積、比葉面積值及較小的小葉排列頻度。而生長在通霄、大肚、綠島及都蘭地區的台灣海棗具較快的葉增加速率，其中以長期受到干擾的西部大肚及通霄地區有較大的葉乾含量及較小的比葉面積。逐步回歸分析顯示年平均溫度、日平均溫度低於12℃的年累積天數百分比、年最高溫度及年最低溫度為主要影響葉特性的因子，與CCA第一軸有相似的結果。依結果大致可將調查的樣區劃分為三大地理區域：東部濱海、關山地區、西部濱海。而依植株高度區分的七個等級中，西部及關山地區均無任一葉特性在高度分級間有顯著差異，僅在東部及關山第14?16樣區之葉增加速率及葉面積隨著高度分級而增加。人工栽種的幼株也隨植株高度的增加，其葉增加速率及綠葉宿存數量均隨之增加。此外，在東部全日光、給水環境的人工處理組尤具明顯較快的葉增加速率及較多的綠葉宿存數量；然而在全光照、泡水組中的葉壽命明顯較其他處理組短。在二年（2011?2013年）葉特性的調查中，最快的葉增加速率出現於2012年夏季，而同年的冬季則最慢；葉增加速率及綠葉宿存數量在2012年夏季及秋季均較前一年大。推測季節變化可能和溫度及雨量豐沛與否有關，另外成熟葉長度及葉伸長速率均受到雨量因子的影響。以上得知台灣海棗植株在不同環境、桿高及季節變換均會採取不同的生長策略，溫度為重要影響生長的因子，光線改變葉片之形態塑形，雨量除促進生長外，降雨天數的多寡會影響增加葉數量或增長葉片長度而影響葉片的生長策略，但泡水導致換氣差而降減葉壽命。
經由生長觀測推算模式得知，台灣海棗約長出79片葉子才形成明顯的桿，並推估現存族群約有94?273年，由最大樹齡可推斷各族群在該地近期存活的時間。綜合了解此物種的生長策略及其樹齡結構，對台灣海棗族群的可塑性、未來動向和保育工作具有重要意義。摘要 i
Abstract iii
目錄 v
表目錄 viii
圖目錄 ix
第一章、 前言 1
一、棕櫚科海棗屬分類及分布狀況 1
二、葉生長特性與因應不同環境的策略 3
三、棕櫚生長型的特性與權衡效益 5
四、研究目的 6
第二章、 材料與方法 7
一、 研究區的概況 7
(一) 樣區地理位置 7
(二) 生育環境概況 10
(三) 氣象站資料與生態氣候圖 14
二、 環境因子蒐集與紀錄 16
(一)微氣候環境監測 16
(二) 地形因子的監測 17
三、台灣海棗植株調查方法 17
(一) 每木調查及葉量測 19
(二) 物候的監測 19
(三) 葉物理特性測量 20
(四) 特性值的計算 21
四、 人工栽培與處理試驗監測 24
五、 資料處理與統計分析 25
(一) 不同樣區、桿高的葉特性差異 26
(二) 環境、季節變化對葉特性的影響 26
(三) 季節變換或採集時間葉特性的差異 28
(四) 人工栽培中不同處理（生育地）的生長差異 29
(五) 樹齡的推算與相關關係檢定 29
(六) 葉伸長的生長模式 29
第三章、 結果 31
一、不同樣區與莖桿高分級間葉特性的差異 31
(一) 不同樣區間葉特性的差異 33
(二) 不同環境對葉特性的影響 41
(三) 樣區生育地、環境因子與台灣海棗葉特性的關係 44
(四) 三大地理區域與植株高度分級的葉特性差異 51
二、季節變換對植株高度分級間葉特性的差異 58
(一) 濱海與內陸樣區的葉特性在不同植株高度分級間之差異 59
(二) 濱海樣區葉特性在不同高度分級及季節間之差異 59
(三) 不同採集季節及高度分級間葉特性之差異 62
三、不同樣區間葉特性對季節變換的反應 63
(一) 濱海與內陸樣區其葉特性在不同季節之變化 65
(二) 解析濱海樣區葉特性季節變化的多樣性 67
(三) 季節變換與葉特性之關係 69
四、人工栽種植株在不同生育地生長的生長差異 71
(一) 不同生育地、高度分級、季節間葉特性的差異 71
五、樹齡的推算 76
(一) 小苗生長及模式 76
(二) 桿高與樹齡的關係 80
(三) 族群存在與樹齡組成的時間推估 85
六、葉伸長的生長模式 85
第四章、 討論 89
一、環境因子對葉特性的影響 89
二、不同植株高度葉特性生長策略 93
三、季節變換對葉特性的反應 94
四、植株成長及樹齡 96
第五章、 參考文獻 98
附錄 105
附錄1、每一生長葉特性在不同樣區間的差異。 105
附錄2、每一結構及產能葉特性在不同樣區間的差異。 107
附錄3、每一樣區的環境因子資料。 109
附錄4、產能葉特性環境因子與植株高度分級間的關係。 111
附錄5、產能葉特性樣區與環境因子的CCA雙序圖。 112
附錄6、支撐結構葉特性環境因子與植株高度分級間的關係。 113
附錄7、支撐結構葉特性樣區與環境因子的CCA雙序圖。 114
附錄8、樣區與環境因子的DCA雙序圖。 115
附錄9、在不同樣區與植株高度分級間東部濱海（2、4、6、8?13樣區）的台灣海棗對12個葉特性值的差異。 116
附錄10、在不同樣區與植株高度分級間關山地區（14?16，21?25樣區）的台灣海棗對12個葉特性值的差異。 117
附錄11、在不同樣區與植株高度分級間關山14?16樣區的台灣海棗對12個葉特性值的差異。 119
附錄12、在不同樣區與植株高度分級間西部濱海的台灣海棗對12個葉特性值的差異。 121
附錄12、在不同樣區與植株高度分級間西部濱海的台灣海棗對12個葉特性值的差異。 122
附錄13、不同植株桿高的台灣海棗植株與綠葉宿存數量的關係。 122
附錄14、不同植株桿高的台灣海棗植株與葉柄長度的關係。 123
附錄15、不同植株桿高的台灣海棗植株與小葉排列頻度的關係。 123
附錄16、不同植株桿高的台灣海棗植株與葉壽命的關係。 124
附錄17、不同植株桿高的台灣海棗植株與葉綠素含量的關係。 124
附錄18、不同植株桿高的台灣海棗植株與葉厚度的關係。 125
附錄19、不同植株桿高的台灣海棗植株與葉厚度的關係。 125
附錄20、每一樣區在不同季節間的季葉增加速率差異。 127
附錄21、每一樣區在不同季節間的季綠葉宿存數量差異。 128
附錄22、每一樣區在不同季節間的季平均葉壽命差異。 130
附錄23、在每樣區的大、中、小植株平均葉增加速率的差異。 13
Diversity of silica bodies on microphylls of Selaginellaceae
No full text矽元素是地表中僅次於氧的第二豐富元素。矽酸鹽礦物經風化遇水溶解形成矽酸（silicic acid, H4SiO4），矽酸被植物吸收利用成生物矽，並以矽質體（phytolith）的方式存在於植物體內。其中，木賊和被子植物中的禾本科與莎草科已被證實植物體內具大量的矽沉積，而苔蘚、石松及蕨類等若干物種亦具有不同形態的矽晶體。卷柏科植物的矽晶體雖有零星的報導，但對其了解仍十分有限。本研究首次針對卷柏小葉表面的矽晶體特徵進行多樣性研究，希望能對本科植物矽晶體的多樣性、分類價值和此特徵和親緣或環境關係的關聯有進一步的瞭解。
卷柏科（Selaginellaceae, 石松植物門）僅有一屬––卷柏屬（Selaginella），目前卷柏屬植物約有800種。本研究的材料取自美洲、歐洲、亞洲、澳洲和夏威夷等地的76個卷柏植物，已接近本科全世界物種數的十分之一，且涵括了七個亞屬。將各種成熟的卷柏葉片經清洗後，以掃描式電子顯微鏡觀察矽晶體於葉表的形態與分布類型，且輔以徒手切片確認矽晶體的沉積位置與形態，並使用原子力顯微鏡（atomic force microscope, AFM）製作出矽晶體於葉表的3D模擬圖。為了解矽晶體中的矽元素含量，以X射線能量散布分析儀（energy dispersive X-ray spectrometer, EDS）對矽晶體做位點分析並製作出葉表的元素含量分布圖。結合卷柏科植物以葉綠體rbcL建構的親緣關係樹，以了解矽晶體和親緣關係的關聯。
研究結果發現74種卷柏葉表具有矽晶體沉積，此顯示矽晶體為卷柏植物普遍存在之特徵。卷柏矽晶體沉積的形態依國際矽晶體命名法規（International Code for Phytolith Nomenclature, ICPN），共命名10種形態，分別為乳突狀、微乳突狀、小顆粒狀、薄板狀、圓錐形、腎形、圓平台形、針形、隆凸狀矽晶體和邊緣長細胞細胞壁之矽累積。
以EDX分析卷柏的小葉顯示葉表的結晶突起物確實為矽晶體，且大部分的矽晶體所含的矽元素量皆高於10%且大致穩定；但微乳突狀、薄板狀和乳突狀矽晶體的矽含量則較低。其中邊緣長細胞細胞壁之矽累積的矽含量最高且矽含量差異也最大（14.98-38.77%），乳突狀矽晶體的平均矽含量最低（0.93-5.08%）。
矽晶體於一個細胞的排列方式有八種，於葉表的分布類型可分成九種。這些特徵與物種相關，且多數物種背葉的腹面具較少或無矽晶體分布。其中，沿著細胞邊緣排列的圓錐形矽晶體僅於復甦卷柏中發現。卷柏葉表常見的六種細胞類型（等徑細胞、長形細胞、邊緣長細胞、異形細胞、毛茸和少數的保衛細胞）均可發現矽元素沉積，但其中保衛細胞者並無特定及明顯晶體形態。
生長在開闊地的陽性卷柏（多為非背腹性之枝條和同型葉）僅發現針形、圓錐形和薄版狀矽晶體，而生長於森林下層或陰暗處的卷柏 (均為背腹性枝條和異型葉) 則有高度多樣性的矽晶體形態（十種）。本研究發現植株的生長形態（匍匐或直立）並不影響矽晶體特徵差異，但意外發現生長於環境溼度極高的卷柏多具有葉緣氣孔。
本研究中，僅Selaginella亞屬之物種（Selaginella deflexa）和部分Ericetorum亞屬之物種（S. gracillima）不具有矽晶體，結合卷柏親緣關係樹發現此兩種卷柏皆為基本群的較原始物種，因此可推測最古老的卷柏可能不具有矽晶體。另外，亦發現親緣關係較接近的物種通常具有相似的矽晶體形態，但矽晶體的葉表分布類型卻與親緣關係無明顯的相關性。
本研究證實不同物種卷柏的矽晶體形態與分布皆具有其獨特性，且矽晶體特徵於各物種中皆具有穩定性。因此，卷柏葉表之矽晶體特徵具有極高的分類價值，且可顯示出古老的卷柏科對矽元素吸收沈積在3億5千萬年的演化過程中的變化。卷柏矽晶體的形態似與植物生長的光環境有關（陽性卷柏多為圓錐形，而陰性卷柏則有極多樣的矽晶體形態），此有趣的結果顯示矽晶體的光學效應值得更進一步探討。Silicon (Si) is the second abundant element on the earth surface. Silicate minerals can be weathered and release silicic acid (H4SiO4). Plants take up silicic acid and precipitate it as biogenic silica structures named phytoliths. It has been shown that Equisetum, Poaceae and Cyperaceae have been proved that they can precipitate Si inside cells or on cell walls. Different forms of silica bodies were also found in bryophyte, lycophyte, and ferns. We still have poor understanding and very little information of silica bodies of Selaginellaceae, although a few species had been studied. This is the first systematic study of silica bodies on microphylls of Selaginellaceae. The aims are to explore the diversity of silica bodies, to evaluate their taxonomic value, and to understand find the relationships between silica bodies and phylogeny in Selaginellaceae.
Selaginella is the only genus in Selaginellaceae (Lycophyta) which comprises ca. 800 species in the world. In this study, 76 Selaginella taxa, nearly one-tenth species of Selaginella collected from America, Europe, Asia, Australia, and Hawaii were studied. Healthy and mature leaves (microphylls) were detached and cleaned before various microscopic observations. The forms and patterns (on leaves surface) of silica bodies were observed by scanning electron microscope. Free hand sections were conducted to confirm the deposition site and forms of silica bodies. The 3D models of silica bodies were constructed by atomic force microscope (AFM), and the content of composition elements of silica bodies were measured by an energy dispersive X-ray spectrometer (EDS)Last, the relationships between silica bodies and the phylogenetic tree of Selaginellaceae constructed from chloroplast rbcL sequences were assessed.
A total of 74 out of the 76 species of Sellaginella studied were found to have silica bodies found on their leaf surfaces. The results show that silica bodies are one of the general features of Selaginella. Following to the International Code for Phytolith Nomenclature, 10 forms of silica bodies in Selaginella were identified. They are papillae, light-protruded papillae, granular, thin-tabular, conical, reniform, rondel, acicular, sinuate carinate and marginal long cell wall with silicon.
The results of EDS proved that the peglike projections on leaves of Selaginella are silica bodies. Most of silica bodies contain more then 10% of silicon, but silicon contents of papillae, light-protruded papillae, and thin-tabular silica bodies are less than 10%. Among them, the silicon content of granular silica bodies varies greatly (0.05-22.32%), and papillae silica bodies have the lowest content of silicon (0.93-5.08%).
There are eight arrangements of silica body on one epidermal cell, and nine patterns of silica bodies on Selaginella leaves were delineated. Generally very few silica bodies or no silica bodies distributed on the ventral side of dorsal leaves in most Selaginella species with anisophylls. The pattern of conical silica bodies scattered along cell margins of an epidermal cell were only can found in resurrection Selaginella. Silica bodies can be found in all of six common cell types (isodiametric cell, elongated cell, marginal long cell, idioblast, trichome, and few guard cells) of Selaginella, but guard cells only deposit silicon (without specific projections).
For the Selaginella species living in sun exposed environments (mostly non-dorsiventral, isophyllous), acicular, conical and thin-tabular silica bodies are their major silica bodies. In contrast, the Selaginella species (all dorsiventral leaves and anisophllous) growing in the understorey of forests (less sunlight) have high diversity of forms of silica bodies (10 forms). There are no relations between the character of silica bodies and plant morphology (creeping or erect) of Selaginella. Unexpectedly, an unusual stomatal type, marginal stomata, was commonly observed in the species living in extreme humid environments (water fall, stream and wet cliff).
The result shows that there are no silica bodies in the subgenus Selaginella (Selaginella deflexa) and some of the subgenus Ericetorum (S. gracillima). Since these they are the basal group of this family, it implies that the ancestors of Selaginella might not have silica bodies. In addition, the species with close relationship usually contain similar forms of silica bodies, but there are no relations between patterns of silica bodies (on leaves) and phylogeny of Selaginella.
This study has revealed high diversity of silica bodies of Selaginella and confirmed these traits are stable and species-specific. The characters of silica bodies have high taxonomic value to be an aid for solving the identification challenge of Selaginellaceae. Moreover, the change of ability of silicon deposition of Selaginella species during the past 350 million years it is uncovered. A close relation between the forms of silica bodies and light environments implies that the possible optical effects of silica bodies of Selaginella are worthy to explore.摘要 i
Abstract iii
目錄 v
表目錄 ix
圖目錄 x
第一章 前言 1
一、卷柏屬之介紹3 1
二、卷柏特徵介紹 1
三、矽元素與矽晶體 4
四、研究目的 7
第二章 材料與方法 8
一、研究材料 8
二、研究方法 10
1. 葉片材料取樣與準備 10
2. 矽晶體形態觀察 10
3. 矽晶體元素分析 12
第三章 結果 13
一、矽晶體特徵 13
1. 葉表矽晶體分布類型 13
2. 卷柏葉表具矽晶體的細胞形態 14
3. 卷柏葉表細胞的表面紋理和矽晶體排列方式 15
4. 矽晶體種類 16
5. 矽晶體之元素分析 22
卷柏科矽晶體特徵 小目錄 38
二、卷柏科矽晶體特徵 40
1. Selaginella anceps (Panama) 41
2. Selaginella arbuscula (Hawaii) 44
3. Selaginella arenicola (Florida, USA) 47
4. Selaginella aristata (Taiwan) 膜葉卷柏 49
5. Selaginella arizonica (Arizona, USA) 52
6. Selaginella arthritica (Panama) 54
7. Selaginella articulata (Panama) 57
8. Selaginella australiensis (Australia) 60
9. Selaginella bisulcata (India) 63
10. Selaginella bombycina (Panama) 66
11. Selaginella boninensis (Taiwan) 小笠原卷柏 69
12. Selaginella caulescens (cultivated, UK; native, East Asia) 72
13. Selaginella chrysoleuca (Panama) 75
14. Selaginella ciliaris (Taiwan and Singapore) 緣毛卷柏 78
15. Selaginella cupressina (Indonesia) 81
16. Selaginella deflexa (Hawaii) 84
17. Selaginella delicatula (Taiwan and China) 全緣卷柏 86
18. Selaginella devolii (Taiwan) 棣氏卷柏 89
19. Selaginella diffusa (Panama) 92
20. Selaginella doederleinii (Taiwan and China) 生根卷柏 95
21. Selaginella douglasii (Oregan, USA) 98
22. Selaginella erythropus (USA) 101
23. Selaginella euclimax (Panama) 104
24. Selaginella eurynota (Panama) 107
25. Selaginella exaltata (Panama) 110
26. Selaginella flagellate (Panama) 113
27. Selaginella flexuosa (Panama) 116
28. Selaginella gracillima (Australia) 119
29. Selaginella haematodes (Panama) 121
30. Selaginella helvetica subsp. pseudonipponica (Taiwan) 擬日本卷柏 124
31. Selaginella heterostachys (Taiwan and China) 姬卷柏 127
32. Selaginella hieronymiana (Indonesia) 130
33. Selaginella horizontalis (Panama) 133
34. Selaginella huehuetenangensis (Panama) 136
35. Selaginella intermedia (Singapore) 139
36. Selaginella involvens (Taiwan) 密葉卷柏 142
37. Selaginella kraussiana (cultivated, UK; native, South Africa) 145
38. Selaginella labordei (Taiwan and China) 玉山卷柏 148
39. Selaginella lepidophylla (USA) 151
40. Selaginella longipinna (Australia) 154
41. Selaginella lutchuensis (Taiwan) 琉球卷柏 157
42. Selaginella martensii (cultivated, UK; native, Mexico and Central America) 160
43. Selaginella minima (Panama) 163
44. Selaginella moellendorffii (Taiwan and China) 異葉卷柏 166
45. Selaginella mollis (Panama) 169
46. Selaginella monospora (India) 172
47. Selaginella nipponica (Taiwan) 日本卷柏 175
48. Selaginella oregana (Olmpic National Park,USA) 178
49. Selaginella pallescens (Panama) 180
50. Selaginella pilifera (USA) 183
51. Selaginella plana (Indonesia) 186
52. Selaginella porelloides (Panama) 189
53. Selaginella porphyrospora (Panama) 193
54. Selaginella pulcherrima (cultivated, UK; native, Mexico) 196
55. Selaginella remotifolia (Taiwan and China) 疏葉卷柏 199
56. Selaginella repanda (Taiwan) 高雄卷柏 202
57. Selaginella revolute (Panama) 205
58. Selaginella rupincola (Arizona, USA) 208
59. Selaginella salazariae (Panama) 210
60. Selaginella sertata (Panama) 213
61. Selaginella simplex (Panama) 216
62. Selaginella stauntoniana (Taiwan) 擬密葉卷柏 219
63. Selaginella stellata (Panama) 222
64. Selaginella tamariscina (Taiwan) 萬年松 225
65. Selaginella uliginosa (Australia) 228
66. Selaginella umbrosa (Panama) 230
67. Selaginella uncinata (Taiwan and China) 翠雲草 233
68. Selaginella underwoodii (Arizona, USA) 236
69. Selaginella vogelii (Australia) 238
70. Selaginella wallacei (Oregan, USA) 241
71. Selaginella wallichii (China) 243
72. Selaginella willdenowii (Australia; native, Thailand, and Malesia) 246
73. Selaginella wolffii (Panama) 249
74. Selaginella sp. new1 (Panama) 252
75. Selaginella sp. new2 (Panama) 255
76. Selaginella sp. new3 (Panama) 258
三、 卷柏植物葉表之矽晶體的綜合比較 261
1. 各種矽晶體的矽含量比較 261
2. 卷柏葉表之矽晶體的大小與分布類型 263
3. 卷柏小葉之矽晶體分布類型示意圖（76種） 265
4. 小型和大型圓錐形矽晶體的分布位置 270
5. 圓錐形矽晶體於等徑細胞表面的排列方式 271
6. 圓錐形與隆凸狀矽晶體於長形細胞表面的排列方式 271
7. 薄板狀矽晶體的多變形態 273
8. 卷柏小葉之六種毛茸形態 274
9. 乳突狀矽晶體的變異性 275
10. 不同矽晶體於同一張SEM照片中的亮度差異 276
11. 零星氣孔和氣孔帶附近的矽晶體形態 277
12. 卷柏之葉緣氣孔 278
13. 卷柏葉表之異形細胞 279
14. 復甦卷柏的復水情況與矽晶體特徵 280
四、台灣之卷柏特徵檢索表 282
第四章 討論 286
一、 卷柏屬植物之矽晶體形態種類和分布類型 288
1. 卷柏屬植物之矽沉積方式與其他科植物的比較 288
2. 矽晶體於卷柏葉表的分布類型 294
3. 矽晶體於細胞表面的排列方式 296
4. 卷柏屬不同矽晶體之3D模型與矽元素含量比較 299
5. 卷柏屬矽晶體與植物特性和環境的關係 302
二、 卷柏矽晶體與親緣之間的相關性 305
1. 卷柏七個亞屬間的矽晶體形態差異 305
2. 卷柏每個亞屬內的矽晶體形態與分布差異 311
3. 新舊大陸卷柏間矽晶體的差異 312
第五章 結論 314
參考文獻 31
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