本研究針對南海北部海域的單細胞固氮生物之時空分布，及影響其分布之環境因子進行探討。研究方法以全細胞免疫化學染色法，利用固氮酵素抗體之探針，將具有固氮酵素抗原的細胞呈色，以利計數，並依體型及外形分成四類，即直徑1-2 μm 之圓形細胞(1-2 μm C)、直徑>2 μm 之圓形細胞(>2 μm C)、直徑1-2 μm 之橢圓形細胞(1-2 μm R)，與>2 μm 之橢圓形細胞(>2 μm R)。探測期間自2010 年的5月至12 月，包含1455 航次(春季)、1487 航次(夏季)與950 航次(冬季)三航次。探測區域自北緯21º至22º，東經116.5º至122.5º，橫跨南海北部之陸棚、陸坡與海盆區，以及黑潮上游。夏季航次之採樣為颱風過後一週進行，其結果可能與無颱風影響的夏季狀況不同。除了自然界的分布之外，夏季在陸棚與海盆測站，冬季在陸坡測站，採集南海表水，分別添加磷、添加鐵及同時添加磷鐵，進行48 小時的培養實驗。
結果發現南海北部的單細胞固氮生物，表水細胞密度(20.4-23.8×104 cells L-1)以春季和夏季略高於冬季；水柱0-100 m 累計生物量(14.8-16.7×109 cells m-2)以夏季和冬季略高於春季，但統計上季節間細胞密度的差異皆不顯著，在四個海區之間的差異亦不顯著。四類細胞中又以1-2 μm C 的細胞密度最高，佔表水與水柱累計總細胞數的47%與49%，>2 μm C 者次之(佔39%與33%)，1-2 μm R (6%與8%)和>2 μm R (8%與9%)細胞密度較低。1-2 μm C 表水細胞密度(9.7-11.5×104 cells L-1)在夏季與春季略高於冬季；>2 μm C 表水細胞密度(7.0-11.1×104 cells L-1)以春季最高，夏季與冬季相當；1-2 μm R (0.8-1.8×104 cells L-1)和>2 μm R (1.3-2.4×104 cells L-1)表水細胞密度皆以夏季高於冬季和春季。表水單細胞固氮生物在季節間變化，除了1-2 μm R 在夏季顯著高於春季與冬季之外(p<0.05)，其他各類者皆不顯著。>2 μm C 表水細胞密度和表水硝酸鹽濃度、N/P 均呈顯著負相關，與水體分層程度之間呈顯著正相關；1-2 μm C 與>2 μm C 兩類圓形單細胞固氮生物均與大氣落塵量之間呈顯著正顯著；顯示硝酸鹽越低、磷酸鹽或是鐵濃度越高時，>2 μm C 生物量越高。營養鹽添加培養實驗結果，顯示在夏季時，>2 μm C 單細胞固氮生物同時受到磷和鐵缺乏之限制，>2 μm R 抑是如此，而1-2 μm R 於夏季可能受到鐵缺乏之限制。
垂直分布而言，屬於圓形的1-2 μm C 和>2 μm C 細胞密度在水柱上層(0-50 m)較高，隨水深遞減，上層:中層(50-100 m):下層(100-150 m 或200 m)細胞密度之比例分別為1.4:1:0.7 與1.6:1:0.7，可能與水柱上層較低的硝酸鹽有關。而1-2 μm R
的比例為1.2:1:0.9，為四類單細胞固氮生物中垂直分布較均勻者。>2 μm R 之比例
為1.4:1:1.0，相較於前三者，其在水柱下層的比例偏高，推測與其能適應較低光照
或/且同時需要磷與鐵有關。

This study investigated the spatial and temporal distribution of unicellular nitrogen-fixing organisms (diazotrophs) in the northern South China Sea (SCS). The diazotrophs contain nitrogenase that can be detected by whole-cell immunocytochemical method using the antibody of nitrogenase, and distinguished and counted readily under a microscope. From present study, the unicellular diazotrophs were classified to four categories according to their sizes and shapes, coccoid cells with diameters of 1-2 μm (1-2 μm C), coccoid with diameters of >2 μm (>2 μm C), rod shape with diameters of 1-2 μm (1-2 μm R), and rod shape with diameters of >2 μm (>2 μm R). All of the field work were conducted between May and December of 2010 during three cruises, including CR1455 (spring), CR1487 (summer) and CR950 (winter). The summer cruise was carried out one week after the passage of a typhoon, which could be different from summer cruise without typhoon event. Sampling stations were located between 21°-22°N and 116.5°-122.5°E covering the shelf, slope and basin of the northern South China Sea. One station was in the upstream Kuroshio. In addition to the field data, some nutrient enrichment experiments were conducted on board the ship, including one on the shelf and one on the basin stations during the summer cruise, and one on the slope station during the winter cruise. Surface water from each station were enriched with including phosphate, iron, phosphate and iron simultaneously, respectively and were compared with unamended control. All trearments were incubated on deck for 48 hours.
The cell densities of the unicellular diazotrophs on surface water (20.4-23.8×104 cells L-1) were somewhat higher in spring and summer than in winter. Water column (0-100 m) integrated abundances (14.8-16.7×109 cells m-2) were slightly higher in summer and winter than spring. However, their seasonal differences were not significant after statistical tests. Among the four categories, 1-2 μm C was the most abundant group, occupying 47% and 49% of total unicellular diazotrophs in the surface water and the whole water column, respectively. The second abundant group was >2 μm C (39% and 33%), and the least abundant groups were 1-2 μm R (6% and 8%) and >2 μm R (8% and 9%). The surface cell density of 1-2 μm C (9.7-11.5×104 cells L-1), 1-2 μm R (0.8-1.8×104 cells L-1) and >2 μm R (1.3-2.4×104 cells L-1) were all highest in summer. The surface density of >2 μm C (7.0-11.1×104 cells L-1) was however highest in spring. Statistical test, however, showed an unsignificant seasonal differences, except that 1-2 μm R which was significantly (p<0.05) higher in summer than winter and spring. The cell density of >2 μm C was negatively correlated to both surface nitrate concentration and the N/P ratio, and pasitively correlated to stratification of water column. Cell densities of two coccoid diazotrophs (1-2 μm C and >2 μm C) both were positively correlated to the dust influx. This suggested that high abundance of >2 μm C occurred at low nitrate concentration and/or high phosphate/iron concentratio. The results of the nutrient enrichment experiments farther support these biological features. As contrast to unamended control, the abundances of >2 μm C and >2 μm R were enriched when phosphate and iron were added simultaneously in summer. Similarly, iron enrichment also stimulated the growth of 1-2 μm R. Vertical distribution of the four diazotrophic groups were relatively different.
The cell density of the coccoid diazotrophs (1-2 μm C and >2 μm C) decreased with depth, with ratios of their densities in the surface (0-50 m): middle (50-100 m): bottom layers (100-150 or 200 m) vertical density profile being 1.4:1:0.7 and 1.6:1:0.7, respectively. High densities in the surface water column might associate with the low nitrate concentration. By contrast, the vertical distribution of 1-2 μm R was relatively homogenous with a vertical cell density profile of 1.2:1:0.9. Among the diazotrophs, >2 μm R showed a highest ratio in the bottom layer, as their vertical cell density profile being of 1.4:1:1.0. The phenomenon implicated that >2 μm R, compared with other diazotrophic groups, probably requires lower illumination and/or higher abundances of phosphate and iron.

論文審定書 i
誌謝 ii
中文摘要 iii
英文摘要 v
目錄 viii
圖次 x
表次 xii
第一章 前言 1
第二章 文獻回顧 4
2.1 南海之水文環境 4
2.2 固氮生物與固氮酵素 10
2.3 單細胞固氮生物之分類 11
2.4 單細胞固氮生物的重要性與其在各海域之分布 18
2.5 影響單細胞固氮生物分布之環境因子 24
第三章 材料方法 31
3.1 水文資料測定 31
3.2 單細胞固氮生物採集 32
3.3 營養鹽添加培養實驗 38
3.4 其他水質資料 38
3.5 數據分析 40
第四章 結果 42
4.1 春季1455 航次 42
4.2 夏季1487 航次 48
4.3 冬季950 航次 50
4.4 季節間之變化趨勢 53
4.5 單細胞固氮生物與環境因子之間的關係 60
4.6 營養鹽添加培養實驗 62
第五章 討論 64
5.1 單細胞固氮生物之分布 65
5.2 垂直分布 72
參考文獻 76

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