臺灣博碩士論文加值系統

橈足類能將碳能從低的營養階層傳遞到高的營養階層裡 (例如魚類)，在河口與海洋食物網中扮演了重要的角色。哲水蚤屬中的安氏偽鏢水蚤 (Pseudodiaptomus annandalei, Swell 1919) 與劍水蚤屬中的短角異劍水蚤 (Apocyclops royi, Lindberg 1940) 是兩種顯著優勢的河口橈足類。過去以來這兩種橈足類都被當成是草食性的種類，也一直被養殖來當成石斑魚及其他經濟性魚類幼苗的活餌。到目前為止，這些橈足類的攝食生態學研究只獲得相當少的關注。在本論文中，調查了這兩種橈足類對於輪蟲 (Brachionus rotundiformis) 與纖毛蟲 (Euplotes sp.) 的攝食生態與掠食行為。更進一步，估計了藻類 (單一與多種藻類攝食)、動物性以及混合的食物對於牠們的繁殖力表現、存活率、胚後期發展率與族群變動之影響。此外，也研究了矽藻的多元不飽和醛 (Poly Unsaturated Aldehydes ,PUAs) 對這兩種橈足類的發育以及族群大小的影響。
在掠食效率的試驗裡，提供了纖毛蟲與三種不同生長階段的 B. rotundiformis 輪蟲 (幼生、抱卵與無抱卵成體) 給橈足類當餌食，藉以調查掠食行為的餌食選擇、對卵的掠食行為、餌食的進食率、餌食的大小選擇、飽足程度與攝食週期。在橈足類對於輪蟲與纖毛蟲的捕食消耗率研究中，也紀錄了添加小型藻類 (Isochrysis galbana) 與大型藻類 (Tetraselmis chui) 後橈足類之攝食表現。本論文亦利用快速攝影機紀錄影像來研究兩種橈足類攝食輪蟲卵、輪蟲成體與纖毛蟲的掠食行為。結果發現雌雄兩性之橈足類都能攝食輪蟲的卵、幼生、成體以及纖毛蟲。安氏偽鏢水蚤的餌食消耗速率高於短角異劍水蚤。研究結果更進一步指出，除了雄性的安氏偽鏢水蚤之外，無論橈足類的性別或是繁殖狀態，對於輪蟲幼生的餌食消耗速率皆高於輪蟲成體與纖毛蟲。另外，當藻類 I. galbana 出現時，橈足類對於餌食的進食消耗沒有影響；當藻類T. chui 出現時則會影響安氏偽鏢水蚤對餌食的消耗率。簡而論之，當食物裡存在 I. galbana 與 T. chui 時將顯著地降低短角異劍水蚤成體的攝食率。當環境裡帶卵的輪蟲數目增加時，橈足類對輪蟲卵的進食消耗率亦隨之上升。在提供 B. rotundiformis 幼生混合輪蟲成體或 Euplotes sp. 細胞的食物時，除了在高比例的纖毛蟲之條件下，短角異劍水蚤雌性對於攝食輪蟲幼生的偏好高於成體與纖毛蟲。橈足類的餌食偏好在環境中相對餌食的比例改變下會展現出轉換的現象。飽食的橈足類比飢餓狀態下的橈足類更常對輪蟲表現出棄卻的行為。安氏偽鏢水蚤的雙重食性特質，證明了相較於短角異劍水蚤，牠是更有效率的掠食者。
實驗室對攝食生態學的評估讓我們可以量化特定攝食、食物密度與食物種類對經濟性養殖足類之相對重要性，透過對攝食生態學與食物生態位階大小的認識是可靠且持續生產的先決條件。經過實驗，這兩種橈足類的食性推估屬於多元的攝食者甚於單一的草食者。因此，本論文對這兩種橈足類的繁殖力表現、存活率與胚後期發展率與族群變動在相對不同攝食組合與食物密度的關聯也進行了相關研究。當給予單一藻種如 I. galbana (I)、T. chui (T) 與 Chaetoceros sp. (C)，或是不同藻種組合如 (I+T、I+C、T+C 與 I+T+C) 的攝食下，橈足類的生產率與胚後期發展率都受到了不同程度的影響。藻類攝食對安氏偽鏢水蚤之相對影響比短角異劍水蚤更加顯著。對於測試的兩種橈足類來說，單一藻種的攝食評估結果跟混合藻種的攝食具有一樣的影響。對安氏偽鏢水蚤來說，單一的 I. galbana 藻種攝食是生殖與發育的最佳食物。對短角異劍水蚤而言，單一藻種的 I. galbana 攝食是增進繁殖最佳的食物，T. chui 則是最快產生下一個世代的最佳食物。當結合藻類 (I. galbana 或 T. chui) 與動物 (不同密度的輪蟲 - 10, 20, 40, 80, 160 and 320 ind. 60ml-1) 的不同攝食條件下，橈足類之族群成長速率會隨著輪蟲密度的增加而上升。實驗結果發現，安氏偽鏢水蚤的最佳大量養殖條件為密度 320ind. 60ml-1 並且給予 I. galbana 單一藻種的攝食。
三種矽藻的多元不飽和醛被拿來測試橈足類 (無節幼生、橈足類幼生與成體) 的感受性試驗。其中 2-trans, 4-trans decadienal (DD) 的毒性最高，還有其他兩種測試的多元不飽和醛分別是 2-trans, 4-trans haptadienal (HD) 與 2-trans, 4-trans octadienal (OD)。實驗發現，相較於短角異劍水蚤來說，安氏偽鏢水蚤對於矽藻多元不飽和醛的感受性更為敏感。同時，發育早期的橈足類 (無節幼生與橈足類幼生) 比成體對於藻類多元不飽和醛的敏感性還要更強。在存有 DD 的培養環境中，兩種橈足類的存活率、胚後期發展率與族群變動都可以看到負面的影響。
本論文提供了安氏偽鏢水蚤與短角異劍水蚤在攝食微浮游動物之效率與行為的先驅研究結果，證明了牠們的食性偏好為雜食性。本論文的研究結果證明這些橈足類的食性是相當廣泛的，暗示這兩種在許多熱帶鹹水池塘與河口終年數量豐富的橈足類在生態中扮演重要的角色，在自然的棲息地中經由掠食行為與雜食的食性形塑出群聚結構。其後，本研究提供了最佳食物養殖條件的資訊，可以應用於穩定的大量養殖橈足類供應給魚苗養殖使用。因此，本論文之研究結果除了生態上的顯著性之外，亦期對於水產之仔稚魚養殖的活餌飼育上有所貢獻。

Copepods play a central role in transferring carbon from lower trophic level to higher trophic level such as fish in the estuarine and marine food webs. The two copepods the calanoid Pseudodiaptomus annandalei (Swell 1919) and the cyclopoid Apocyclops royi (Lindberg 1940) are dominant estuarine copepod species. Both the species were considered to be herbivorous, and being cultured as live feed for grouper and other economical fish larvae. Little attention has so far been paid on feeding ecology of these copepod species. In this thesis, the feeding ecology and predation behavior of these two copepod species on rotifer Brachionus rotundiformis and ciliate Euplotes sp. were elucidated quantitatively as well as qualitatively. Their reproductive performance, survival, postembryonic developmental rates and population dynamics have been estimated on diets of phytoplankton (mono- and plurialgal), animal and combinations theirof. Moreover, the influence of Poly Unsaturated Aldehydes (PUAs) of diatom on development and population size of each copepod species were also investigated.
For predation efficiency examination, these copepods were offered ciliate and three states of B. rotundiformis (neonates, ovigerous and non-ovigerous adults) to investigate prey choice, egg predation, prey consumption rates, size selection, and satiation levels and duration. Prey consumption rates on rotifer and ciliate were also recorded in the presence and absence of algae, the smaller Isochrysis galbana and the larger Tetraselmis chui. The predation behavior of each copepod species was videotaped by Fastcam providing information about ingestion of rotifer eggs, rotifer adults and ciliates. Both males and females of each copepod species were able to ingest on eggs, neonates, adults of rotifer and ciliate. Prey consumption rates were higher in P. annandalei than in A. royi. Regardless of sex and reproductive state of the copepods, prey consumption rates were higher on rotifer neonates than on adults and ciliates except the case of male of P. annandalei. The presence of I. galbana had no effect on prey consumption; T. chui influenced the prey consumption rates in P. annandalei. However, the presence of either I. galbana or T. chui significantly reduced the prey ingestion rates in A. royi adults. The egg consumption in copepods increased with increasing numbers of egg bearing rotifers in their environment. From a combination of neonates of B. rotundiformis with either adults rotifer or cells of Euplotes sp. copepod females preferred neonates over adult rotifers and ciliate except at higher proportion of ciliate in A. royi. The prey preference of copepods exhibited switching based on relative prey proportion in the environment. Satiated copepods rejected rotifers more often than hungry individuals. Dual feeding nature of P. annandalei attested that this is more efficient predator than A. royi.
Laboratory evaluation of feeding ecology allows us to quantify relative importance of particular diet; food density and food type for commercial culture of copepod species. A thorough knowledge of feeding ecology and food niche size are prerequisite for reliable and sustained production. Both the copepods were estimated as generalist feeders rather than simple herbivorous. Therefore, the reproductive performance, survival and postembryonic developmental rates and population dynamics of each copepod species in relation to food diets and densities were also studied. With monoalgal diets, I. galbana (I), T. chui (T) and Chaetoceros sp. (C) and their different combinations (I+T, I+C, T+C and I+T+C). The reproduction and postembryonic development rates of copepods were influenced differentially by different diets. Algal diets related effects were more pronounced in P. annandalei than in A. royi. Monoalgal diet was evaluated as effective as mixed algal diet in P. annandalei. I. galbana alone has been recorded best food for reproduction and development for P. annandalei. T. chui alone was estimated to be best food for survival and fastest generation time in A. royi. Whereas the mixture of I. galbana + T. chui was found to be best diet for reproduction in A. royi. With combination of algae (either I. galbana or T. chui) and animal (different density of rotifer- 10, 20, 40, 80, 160 and 320 ind.60 ml-1) diet, the higher population growth rate was noted with increasing density of rotifer. The optimal culture condition for reliable mass culture of P. annandalei was recorded 320 ind. 60ml-1 with I. galbana.
Susceptibility of copepods (nauplii, copepodites and adults) was tested to three PUAs of diatom. 2-trans, 4-trans decadienal (DD) was recorded more toxic compared to other two tested PUAs [2-trans, 4-trans haptadienal (HD); and 2-trans, 4-trans octadienal (OD)] to copepods. P. annandalei was estimated more sensitive to PUAs compared to A. royi. Further, the early stages (nauplii and copepodites) were found more sensitive to PUAs in comparison to adults of copepods. The presence of DD in the medium negatively influenced the survival, post embryonic developmental rates and population growth rates of each copepod species.
This thesis provides first information on predatory efficiency and behavior of P. annandalei and A. royi on microplankton, which attests their omnivory feeding habit. Further, the wider food spectrum of these copepods suggests the underlying reasons of perennial abundance of these copepods in many tropical brackish ponds and estuaries. Present results also indicate the major role played by these copepods in shaping the community structure through predation and omnivory in their natural habitat. Subsequently, this thesis provides information about optimal food culture conditions to sustain reliable mass cultivation of these copepods for fish larviculture. Therefore, this thesis has ecological significance as well as aquaculture implications in live feed culture for growing fish larvae.

Acknowledgements I
摘要 III
Abstract V
Table of contents VII
List of figures X
List of tables XVIII
CHAPTER 1 General introduction 1
1.1 General overview 2
1.2 Biological model for the study 4
1.3 Thesis overview 5
CHAPTER 2 Review of literature 7
2.1 Copepods 8
2.2 Ecology and feeding behavior 9
2.3 Food effect on life history traits 12
2.3.1 Effect of food types and concentration 12
2.3.2 Effect of mixed diets 14
2.3.3 Effect of algal toxins 15
2.4 Life history attributes of Pseudodiaptomus annandalei and Apocyclops royi 18
2.4.1 Pseudodiaptomus annandalei 18
2.4.2 Apocyclops royi 21
2.5 Aquaculture importance of targeted copepods 23
CHAPTER 3 Predation: ecological and behavioral assessments 36
3.1 Introduction 37
3.2 Methods 37
3.2.1 Zooplankton sampling and culture of the experimental organisms 37
3.2.2 Experimental design 39
3.3 Results 45
PART 1 Ecological assessment 45
PART 2: Behavioral assessment 49
3.4 Discussion 50
3.4.1 Prey consumption rates 51
3.4.2 Algal effect 52
3.4.3 Egg predation 53
3.4.4 Prey size and selection 53
3.4.5 Satiation level and duration 54
CHAPTER 4 Reproduction and postembryonic developmental rates: microalgal effects 79
4.1 Introduction 80
4.2 Methods 80
4.3 Results 84
4.3.1 Reproduction 84
4.3.2 Postembryonic developmental rates 87
4.4 Discussion 90
CHAPTER 5 Functional and numerical responses: effects of animal and algal diets 107
5.1 Introduction 108
5.2 Methods 108
5.2.1 Functional responses 108
5.2.2 Numerical response 109
5.3 Results 110
5.3.1 Functional responses 110
5.3.2 Numerical response 111
5.4 Discussion 117
CHAPTER 6 Postembryonic developmental rates and population dynamics: effect of diatom toxins 135
6.1 Introduction 136
6.2 Methods 136
6.2.1 Stock solution 137
6.2.2 Experiment design 137
6.3 Results 141
6.3.1 Susceptibility of the copepods 141
6.3.2 Postembryonic developmental rates 143
6.3.3 Population size 146
6.4 Discussion 149
CHAPTER 7 Conclusions and future prospectives 183
Bibliography 187
Appendix 227

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