本論文探究台灣家白蟻（Coptotermes formosanus Shirake）腸道之內容物、微生物及功能，並對於木質纖維素之分解作用與效率進行評估。試驗選取野外誘引及餵食不同食材之人工養殖台灣家白蟻工蟻及兵蟻之腸道，採用DIC光學顯微鏡、影像分析儀（FS-180U）及掃描式電子顯微鏡（SEM）進行白蟻之腸道內容物觀察。另以傅立葉轉換紅外線光譜儀（FT-IR）及氣相層析質譜儀（GC-MS）分析白蟻腸道內容物之成分。本研究同時採用傳統生物及分子生物鑑定法鑑定台灣家白蟻腸道之微生物，並探討其對纖維素之分解能力。
藉由DIC光學顯微鏡觀察台灣家白蟻腸道發現，在工蟻的囊形胃內部存在著大量的共生原生動物，有Holomastigotoides sp., Pseudotrichonympha sp. 及Spirotrichonympha leidyi等3種鞭毛蟲且隨餵食材料不同，而腸道中之共生鞭毛蟲種類隨著改變。白蟻腸道SEM影像，以酒精醋酸異戊酯脫水處理者最佳，其次為冷凍乾燥處理，室內風乾處理者最差。SEM觀察結果兵蟻的囊形胃及各腸道中類似細菌的球狀顆粒略少於工蟻。推測可能因工蟻兼有餵食其他類型蟻種之任務，故其腸內需對所進食之物進行消化及代謝，因此工蟻腸道的內容物較為豐富。
藉由FT-IR分析台灣家白蟻腸道內容物成分，以代表癒瘡木基（1327 cm-1）及木質素芳香環（1510 cm-1）等吸收峰之比值相較，發現工蟻之腸道對於木質素中的癒瘡木基和紫丁香基等官能基具消化作用。進一步以GC-MS分析，台灣家白蟻工蟻之腸道內含有大量的烷類、環烷類及脂肪酸等化合物成分，隨餵食不同食材者之腸道內容物而成分各不相同，證實台灣家白蟻腸道可將木質纖維素食材分解成不同類型之化合物。
由白蟻腸道內微生物之培養，並透過傳統生物鑑定法-生化八管及分子生物鑑定法結果得知，台灣家白蟻工蟻及兵蟻腸道共同存在之菌株為：Citrobacter koseri, Enterobacter cloacae及Proteus mirabilis等革蘭氏陰性菌。由於工蟻在台灣家白蟻的生態中主要擔任攝食者的角色，故在腸道內發現之菌種較兵蟻者多。經與餵食相異食材之腸道菌相比對，發現野生白蟻腸內之優勢菌株會隨食材不同而有所變化。分析菌株纖維素分解能力之，發現野外收集或經人工養殖餵食不同食料後具纖維素分解能力之菌株台灣家白蟻工蟻腸道中皆多於兵蟻者，經以分析濾紙作為食材的台灣家白蟻工蟻腸道所分離出之菌株中，如：Vibro furnssii, Enterobacter asburiae及Escherichia hermannii，皆具纖維素分解之能力。
藉由本研究分析對於台灣家白蟻腸道之結構、蟻種間腸道存在菌株的差異及菌株在腸道的分佈，篩選出纖維素分解效率較高之優勢菌種，可應用於木質材料之生質能源的研發。

The aim of this study is to investigate the structure, microbial and functions of the guts from Formosan termite (Coptotermes formosanus Shirake) worker and soldier, we collected guts from the worker and soldier termites the wild-type and feeding with different food stuffs. The intestinal contents were observed by DIC microscope, image analyzer (FS-180U) and scanning electron microscope (SEM), then to analysis the components of termite gut contents with Fourier Transform Infrared Spectroscopy (FT-IR) and Gas Chromatography Mass Spectrometry (GC-MS). The study also uses the traditional and molecular identification method to identify microorganism from the termite's guts, and to investigate its decompose ability of cellulose.
According to the DIC microscope observation, we found a large number of the symbiotic protozoa in the worker termite's paunch. The symbiotic gut flagellates showed considerable various with different feeding material. We observed three kinds of symbiotic gut flagellates: Holomastigotoides sp., Pseudotrichonympha sp. and Spirotrichonympha leidyi. The best images of SEM photographs of termite's guts were the isoamyl acetate alcohol dehydration treatment, followed by the freeze-drying treatment and the indoor air-dried treatment. By SEM observation, the spherical particles similar to bacteria from the soldier termite's paunch and various intestinals but somewhat less in worker. We estimate that the soldier termites ate food stuffs by worker digested. Therefore a variety of the intestinal contents of worker were richer than soldier's.
The termite's guts contents analyzed by FTIR, we compared the ratio of the absorption of representative guaiacuyl group (1327 cm-1) and lignin aromatic ring (1510 cm-1) to get the gut of worker termite can digest the functional groups of guaiacuyl and syringyl lignin. Further analyses by the results of GC-MS, the intestinal of worker termite contains a lot of alkanes, cyclic alkanes and fatty acids compounds. The composition of intestinal contents from feeding with different materials termites were various. Therefore we proved that the worker termite's gut could decompose lignocellulose into different types of compounds.
With the intestinal microbial cultures, by means of traditional and molecular identification method, three common bacterial species found both in guts of worker and soldier termites: Citrobacter koseri, Enterobacter cloacae and Proteus mirabilis. Because worker termites play the role of the main feeder in ecology, the amount of bacteria species from worker's guts were more than the soldier's. And then we compared the bacteria species of the intestinal from feeding different materials termites, results showed the advantages bacteria species from wild-type termite's intestines will vary with different ingredients. In cellulolytic test, whether the bacterial species of the guts from the wild-type or farmed different materials worker termite, the bacterial species with cellulose degradation ability are more than the soldier's. Vibro furnssii, Enterobacter asburiae and Escherichia hermannii were mainly species and formed the decomposition circle on carboxymethylcellulose medium （CMC） from the worker termite’s guts by feeding with filter paper.
In this study, we had well understood about the structure of the termite's guts, the differences and distribution of the bacterial species between the worker and soldier termite's guts. In the future, the research will help to isolate the high efficiency of the cellulolytic species that apply to the developing of biomass energy from wood materials.

目錄----------------------------------------------------------------------------------------------Ⅰ
表目錄----------------------------------------------------------------------------------------Ⅴ
圖目錄----------------------------------------------------------------------------------------- Ⅷ
中文摘要-------------------------------------------------------------------------------------- XII
Abstract------------------------------------------------------------------------------------- XIV

第一章 前言----------------------------------------------------------------------------------- 1
第二章 文獻回顧----------------------------------------------------------------------------- 4
第一節 白蟻分類及分布------------------------------------------------------------------ 4
2.1.1 全球白蟻的分類及分布--------------------------------------------------------- 4
2.1.2 台灣本土白蟻的分布------------------------------------------------------------ 6
第二節 白蟻生態及危害------------------------------------------------------------------ 8
2.2.1 白蟻生態--------------------------------------------------------------------------- 8
2.2.2白蟻之危害------------------------------------------------------------------------ 11
第三節 白蟻腸道及共生之微生物----------------------------------------------------- 12
第三章 白蟻腸道之顯微觀察------------------------------------------------------------ 14
第一節 前言------------------------------------------------------------------------------- 14
第二節 材料與方法---------------------------------------------------------------------- 17
3.2 試驗材料---------------------------------------------------------------------------- 17
3.2.1台灣家白蟻----------------------------------------------------------------------- 17
3.2.2 餵食試驗材料------------------------------------------------------------------- 17
3.2.3 試驗儀器------------------------------------------------------------------------- 17
第三節 試驗方法------------------------------------------------------------------------- 18
3.3.1 白蟻之餵食處理---------------------------------------------------------------- 18
3.3.2白蟻腸道外觀及內容物觀察-------------------------------------------------- 19
第四節 結果與討論---------------------------------------------------------------------- 20
第五節 結語------------------------------------------------------------------------------- 29
第四章 白蟻腸道共生之微生物---------------------------------------------------------- 32
第一節 前言------------------------------------------------------------------------------- 32
第二節 材料與方法---------------------------------------------------------------------- 33
4.2 試驗材料--------------------------------------------------------------------------- 33
4.2.1 試驗白蟻------------------------------------------------------------------------- 34
4.2.2 餵食試驗材料------------------------------------------------------------------- 34
4.2.3 試驗用培養基------------------------------------------------------------------- 34
4.2.4 試驗試劑及緩衝液------------------------------------------------------------- 35
第三節 試驗方法------------------------------------------------------------------------- 36
4.3.1 白蟻之餵食處理---------------------------------------------------------------- 36
4.3.2 白蟻腸道採集------------------------------------------------------------------- 36
4.3.3 白蟻腸道微生物之培養------------------------------------------------------- 37
4.3.4 培養微生物之生物鑑別------------------------------------------------------- 38
第四節 結果與討論---------------------------------------------------------------------- 41
第五節 結語------------------------------------------------------------------------------- 50
第五章 白蟻腸道內容物之成分分析----------------------------------------------------- 51
第一節 前言--------------------------------------------------------------------------------51
第二節 材料與方法----------------------------------------------------------------------- 54
5.2. 試驗材料--------------------------------------------------------------------------- 54
5.2.1 試驗白蟻-------------------------------------------------------------------------- 54
5.2.2 餵食試驗用之材料-------------------------------------------------------------- 54
5.2.3 試驗儀器-------------------------------------------------------------------------- 54
第三節 試驗方法-------------------------------------------------------------------------- 54
5.3.1 白蟻之餵食處理----------------------------------------------------------------- 54
5.3.2 白蟻腸道採集-------------------------------------------------------------------- 55
5.3.3 白蟻腸道之FTIR分析-------------------------------------------------------- 55
5.3.4 白蟻腸道之氣相層析質譜儀（GC/MS）分析---------------------------- 55
第四節 結果與討論---------------------------------------------------------------------- 55
第五節 結語------------------------------------------------------------------------------- 66
第六章 白蟻腸道之分解機制------------------------------------------------------------- 68
第一節 前言------------------------------------------------------------------------------- 68
6.1.1 纖維素之結構及其分解酶----------------------------------------------------- 68
6.1.2 半纖維素之結構及其分解酶-------------------------------------------------- 70
6.1.3木質素之結構及其分解酶------------------------------------------------------ 71
6.1.4 白蟻腸道之分解機制----------------------------------------------------------- 72
第二節 材料與方法----------------------------------------------------------------------- 80
6.2 試驗材料---------------------------------------------------------------------------- 80
6.2.1 試驗之菌株----------------------------------------------------------------------- 80
6.2.2 試驗用培養基-------------------------------------------------------------------- 81
第三節 試驗方法-------------------------------------------------------------------------- 82
6.3.1 菌株之纖維素分解能力測試-------------------------------------------------- 82
6.3.2 菌株內切型葡聚糖酶活性測試----------------------------------------------- 82
6.3.3 菌株外切型葡聚糖酶活性測試----------------------------------------------- 82
第四節 結果與討論----------------------------------------------------------------------- 83
第五節 結語-------------------------------------------------------------------------------- 87
第七章 結論----------------------------------------------------------------------------------- 89
第八章 參考文獻----------------------------------------------------------------------------- 93


表目錄
頁次
表1-1 白蟻的分類與分佈------------------------------------------------------------------ 5
Table 1-1 The classification and distribution of termites

表3-1 台灣家白蟻餵食不同材料 30天後之重量損失率--------------------------- 21
Table 3-1 The average weight loss rate of the feeding materials from
Coptotermes formosanus 30 day after feeding

表4-1 分子生物鑑定法採用之引子對---------------------------------------------------- 36
Table 4-1 The primers the molecular identification of microbial species

表4-2 台灣家白蟻工蟻及兵蟻腸道單一菌落之革蘭氏染色結果------------------ 43
Table 4-2 Gram staining of the bacteria from the guts of worker and soldier
termite’s

表4-3 無氧狀態培養1天及4天下分離出之台灣家白蟻工蟻及兵蟻腸道菌株--- 44
Table4-3 Bacterial species from the guts of Coptotermes formosanus worker and
soldier on LA medium in culturing for 1 and 4 days in aerobic and
anaerobic condition

表4-4 有氧狀態培養1 d及4 d下分離出之台灣家白蟻工蟻及兵蟻腸道菌株----- 45
Table 4-4 Bacterial species in the guts of worker and soldier of Coptotermes
formosanus in aerobic condition on LA medium for 1d and 4d

表4-5有氧狀態培養第4天台灣家白蟻工蟻及兵蟻腸道分離之菌株------------- 47
Table 4-5 Bacterial species in the guts of worker and soldier of Coptotermes
formosanus in aerobic condition on LA medium


表4-6 無氧狀態培養台灣家白蟻工蟻及兵蟻腸道分離之菌株--------------------- 48
Table 4-6 Bacterial species in the guts of worker and soldier of Coptotermes
formosanus in anaerobic condition on LA medium

表4-7 餵食不同食材之台灣家白蟻腸道菌相------------------------------------------ 49
Table 4-7 The intestinal flora from the gut of Coptotermes formosanus by
after feeding different materials

表5-1台灣家白蟻及餵食不同食料者腸道之FTIR吸收峰數值比---------------- 57
Table 5-1 The value ratio of FTIR absorption peak from of the Coptotermes
formosanus guts and feeding with different material

表-2野外台灣家白蟻工蟻腸道內容物之成分分析------------------------------------ 59
Table -2 The components of intestinal contents from the wild-type worker
termite's

表5-3分析濾紙之GC-MS成分分析---------------------------------------------------- 62
Table 5-3 The GC-MS analysis of the components of the filter paper

表5-4 餵食分析濾紙台灣家白蟻工蟻腸道內容物之GC-MS成分分析----------- 63
Table 5-4 The components of intestinal contents from worker termite's feeding
with filter paper

表5-5 台灣二葉松木片之GC-MS成分分析--------------------------------------------- 64
Table 5-5 The GC-MS analysis of the components of Pinus taiwanensis

表5-6 餵食台灣二葉松木片台灣家白蟻工蟻腸道內容物GC-MS成分分析----- 65
Table 5-6 The components of intestinal contents from worker termite's feeding
with Pinus taiwanensis
表6-1 純化自白蟻或共生系統的木聚醣------------------------------------------------ 77
Table 6-1 Xylanases purified or sequenced from termites or symbiotic
microorganisms

表6-2台灣家白蟻腸道菌株--------------------------------------------------------------- 81
Table 6-2 The intestinal flora from the gut of Coptotermes formosanus

表6-3 台灣家白蟻工蟻及兵蟻腸道菌株之纖維素分解測試------------------------ 84
Table 6-3 The cellulolytic capability of bacterial species from the guts of worker
and soldier Coptotermes formosanus

表6-4 台灣家白蟻工蟻腸道所分離出具纖維素分解能力之菌株----------------- 85
Table 6-4 The cellulolytic bacterial species with cellulase activity






















圖目錄
圖2-1 各種白蟻之外觀照片----------------------------------------------------------------- 6
Fig. 2-1 The photographs of the different family termites

圖2-2 試驗流程圖---------------------------------------------------------------------------- 13
Fig. 2-2 The flow chart of research

圖3-1白蟻之腸道示意圖-------------------------------------------------------------------- 14
Fig. 3-1 The schematic diagram of the termite's gut

圖3-2 高等白蟻與低等白蟻腸道結構---------------------------------------------------- 15
Fig. 3-2 The schematic diagram of the gut between lower termites and higher
termites
圖3-3 台灣家白蟻腸道內原生動物組成之百分比------------------------------------ 16
Fig.3-3 The percentage of the protozoa in the gut of Coptotermes formosanus

圖3-4 台灣家白蟻腸道收集示意圖------------------------------------------------------ 19
Fig. 3-4 The sampling process of guts from Coptotermes formosanus

圖3-5 以影像分析儀FS-180U觀察野外台灣家白蟻工蟻及兵蟻之腸道 --------- 22
Fig. 3-5 The observation of the guts of Coptotermes formosanus worker and
soldier by FS-180U

圖3-6 餵食不同食料之台灣家白蟻工蟻及兵蟻腸道外觀----------------------------23
Fig. 3-6 The appearance of the guts from workers and soldiers Coptotermes
formosanus after feeding different materials

圖3-7 以微分干涉差顯微鏡觀察野外台灣家白蟻工蟻腸道中鞭毛蟲之觀察--- 24
Fig. 3-7 The observation of the flagellates in the guts of wild-type worker
Coptotermes formosanus by DIC microscope

圖3-8 以微分干涉差顯微鏡觀察野外台灣家白蟻工蟻腸道中鞭毛蟲-------------25
Fig. 3-8 The observation of the flagellates in the guts wild-type soldier
Coptotermes formosanus by DIC microscope

圖3-9 以微分干涉差顯微鏡觀察餵食台灣二葉松之台灣家白蟻工蟻腸道中鞭毛
蟲之觀察------------------------------------------------------------------------------ 26
Fig. 3-9 The observation of the flagellates of guts from Pinus taiwanensis-feeding
worker Coptotermes formosanus by DIC microscope

圖3-10 以微分干涉差顯微鏡觀察餵食分析濾紙之台灣家白蟻工蟻腸道中鞭毛蟲
之觀察--------------------------------------------------------------------------------- 26
Fig. 3-10 The observation of the flagellates of guts from filter papaer-feeding
worker Coptotermes formosanus by DIC microscope

圖4-1 台灣家白蟻腸道微生物之multiplex PCR電泳圖-----------------------------42
Fig. 4-1 The electrophoresis of multiplex PCR products of miciobial amplified
from the gut of Coptotermes formosanus

圖5-2 野外台灣家白蟻工蟻及餵食不同食料者之FT-IR圖----------------------- 56
Fig. 5-2 The FTIR spectrum the wild-type and worker termite's guts after
feeding different material

圖5-3 野外台灣家白蟻兵蟻及餵食不同食料者之FTIR圖------------------------ 56
Fig. 5-2 The FTIR spectrum the wild-type and soldier termite's guts after
feeding different material

圖5-4 野外台灣家白蟻工蟻腸道之GC-MS光譜圖--------------------------------- 59
Fig. 5-4 The GC-MS spectrum the wild-type worker termite's guts

圖5-5 分析濾紙之GC-MS 光譜圖----------------------------------------------------- 61
Fig. 5-5 The GC-MS spectrum the filter paper

圖5-6餵食濾紙後台灣家白蟻工蟻腸道GC-MS光譜圖---------------------------- 62
Fig. 5-6 The GC-MS spectrum worker termite's guts after feeding with filter
paper
圖5-7台灣二葉松之GC-MS 光譜圖---------------------------------------------------- 63
Fig. 5-7 The GC-MS spectrum from the Pinus taiwanensis

圖5-8餵食台灣二葉松之台灣家白蟻工蟻腸道GC-MS光譜圖-------------------- 64
Fig. 5-8 The GC-MS spectrum after feeding with Pinus taiwanensis-feeding
worker termite's guts

圖5-9 野外台灣家白蟻與餵食不同食料者之GC-MS光譜圖---------------------- 66
Fig. 5-9 The GC-MS spectrum from wild-type and feeding different
materials worker termite's guts

圖6-1 內切型葡聚醣酶水解纖維素過程------------------------------------------------ 69
Fig. 6-1 The process of cellulose hydrolysis by endo-glucanase

圖6-2 β-葡萄糖苷酶水解纖維素過程---------------------------------------------------- 69
Fig. 6-2 The process of cellulose hydrolysis byβ- glucosidase

圖6-3外切型葡聚醣酶水解纖維素過程------------------------------------------------- 70
Fig. 6-3 The process of cellulose hydrolysis by exo-glucanase

圖6-4 半纖維素水解作用------------------------------------------------------------------ 71
Fig. 6-4 Hydrolysis of hemicellulose

圖6-5 雲杉木質素的完整結構模型------------------------------------------------------ 72
Fig. 6-5 The comprehensive structural model of spruce lignin

圖6-6 低等白蟻腸道纖維素分解系統示意圖------------------------------------------- 75
Fig. 6-6 The schematic diagram of the cellulolytic systems in lower termites

圖6-7低等白蟻的木質素降解系統------------------------------------------------------- 79
Fig. 6-7 The lignocellulose-degrading systems of lower termite

圖6-8 台灣家白蟻腸道菌株之內切型葡聚醣酶--------------------------------------- 86
Fig. 6-8 The endo-glucanase of bacterial species from Coptotermes formosanus
圖6-9 台灣家白蟻腸道菌株之外切型葡聚醣酶--------------------------------------- 89
Fig. 6-9 The exo-glucanase of bacterial species from Coptotermes formosanus

圖版目錄
圖版1 相異乾燥處理之台灣家白蟻工蟻腸道SEM照相---------------------------- 28
Plate 1 The photograph of SEM from the guts of Coptotermes formosanus by
three different drying methods

圖版2經酒精醋酸異戊酯脫水處理台灣家白蟻腸道之SEM照相----------------- 29
Plate 2 The photograph of SEM from the guts of Coptotermes formosanus by
alcohol-isoamyl acetate dehydration

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