在世界各海域都發生水母大量增生的問題，且此現象可能隨著全球暖化、水污染及過漁的問題惡化而日益嚴重。水母在固著的底棲階段(水螅體)時會以無性的出芽生殖方式產生新的水螅體並以橫裂生殖方式產生橫裂體進而成長為自由浮游的碟狀體，而此碟狀體可逐漸發育成為能行有性生殖的浮游水母體。很多環境因子都會影響水母的無性生殖，包含食物、溫度、鹽度及光等。本研究是將源自於台灣西南部大鵬灣的海水水母 (Aurelia aurita L.)樣品所培育的水螅體，分兩階段飼養在不同的溫度、光強度及鹽度中，以探討這些環境因子對海月水母無性生殖的影響。在第一階段實驗中，水螅體在十二小時照光-十二小時黑暗的光週期下，分別以三種溫度 (20, 25, 30℃)及三種光強度 (372, 56, 0 lux)共九種組合下進行飼養。觀察發現，水螅體在較高的溫度及較強的光強度之下有較多的出芽生殖，並產生較多新的水螅體。同時，較高的溫度亦縮短了橫裂生殖所須要的時間，並在此實驗中提高了每日新生碟狀體的量。碟狀體在無性生殖中所佔的比例在較高的溫度及較強的光強度下都提高，但存活率在較高的溫度的環境中較低。由無性生殖的觀測結果得知，中間至高溫的環境可造成較快的生殖並且產生較多的水母，然而過高的溫度則可能會因為死亡率大增而有反效果。在第一階段的實驗中，光強度相較於溫度對無性生殖的影響較小，只對加速橫裂生殖有顯著影響，然而飼養在最低溫-全黑的水螅體完全沒有橫裂生殖，顯示溫度和光強度可能有加乘的影響。在本次實驗中，溫度和光強度在水螅體的存活時間及預估水母未來產量上皆有顯著的交互作用，因此推估在自然環境中此二環境因子的交互作用對水螅體的橫裂生殖是重要的。在第二階段實驗中，水螅體則在完全黑暗的環境中，以三種溫度 (27, 31, 35℃)及三種鹽度 (25, 30, 35)共九種組合下進行飼養。觀察發現，水螅體在較冷的溫度及較低的鹽度之下有較多的出芽生殖，產生較多新的水螅體。碟狀體在無性生殖中所佔的比例在較高的溫度提高，而存活率在較高的鹽度環境中較低。綜合兩實驗的結果來比較溫度、光強度和鹽度三種環境因子對海月水母水螅體的無性生殖的影響可得知，溫度的影響最為明顯。水螅體在特定的溫度範圍內，會隨著溫度上升而縮短橫裂生殖所須要的時間，並且提高橫裂生殖在無性生殖中的比例；但是持續地升溫則反而會因為死亡率提高及出芽生殖下降而減少最後水母的產量。雖然第二階段全黑暗的實驗溫度整體上較第一階段有光的實驗溫度高，但第二階段實驗中的橫裂生殖卻比第一階段的少很多；因此，光存在與否對無性生殖的影響可能比光強度還重要。鹽度對水螅體無性生殖的影響無溫度或光來得明顯，只在出芽生殖上有較顯著影響，並在存活時間及碟狀體佔總無性生殖的比率上有顯著影響。

Jellyfish blooms create problems worldwide, which may increase with global warming, water pollution, and over-fishing. Benthic polyps (scyphistomae) asexually produce buds and small jellyfish (ephyrae), and this process may determine the population size of the large, swimming scyphomedusae. Environmental factors that affect the asexual reproduction rates include food, temperature, salinity, and light. In the present study, polyps of Aurelia aurita (L.), originated from Tapong Bay, southwest Taiwan, were studied in different combinations of temperatures (T), light intensities (L), and salinities (S). In the T (20, 25, 30°C) × L (372, 56, and 0 lux) experiment which was with a 12 h light-12 h dark photoperiod, production of new buds decreased with warmer temperatures and stronger light intensity. Warm temperatures accelerated strobilation and increased the daily production of ephyrae. The proportion of ephyrae to total asexual reproduction (new buds + ephyrae) increased dramatically in warmer temperatures and stronger light. Survival period was reduced at the highest temperature. Strobilation did not occur at the lowest temperature in darkness. All measures of total asexual reproduction indicated that medium to high temperatures would lead to faster production of more jellyfish; however, continuous high temperatures might result in high polyp mortality. Light intensity affected asexual reproduction less than did temperature, only significantly accelerating the strobilation rate. Because the interactive effects of light and temperature were significant for polyp survival time and the production of jellyfish per polyp, combined light and temperature effects are likely important for strobilation in situ. In the T (27, 31, 35°C) × S (25, 30, and 35) experiment which was in dark environment, production of new buds decreased with higher temperatures and salinity. The proportion of ephyrae to total asexual reproduction (new buds + ephyrae) increased with warmer temperatures, but survival period was reduced at the highest salinity, and strobilation was substantially reduced, even though the temperature was warmer compared to the T × L experiment. Salinity affected asexual reproduction less than did temperature, only significantly affecting production of new buds, and slightly affecting survival period and the proportion of ephyrae to total asexual reproduction. According to these two experiments, warmer temperature may accelerate strobilation in light condition and lead to better yield of swimming jellyfish, however continuously warm temperature would reduce the yield by decreasing budding and higher mortality. Complete dark led to much less strobilation, especially at low temperatures, suggesting that the existence of light might be more important than light intensity. The effects of salinity on asexual reproduction were not as conspicuous as that of temperature and light.

Contents
Chapter One Introduction……………………………………………………………… 1
Chapter Two Materials and methods ………………………………………………….. 6
Experiment one: Temperature × light intensity……………………. 6
Experiment two: Temperature × salinity…………………………... 8
Statistical analysis…………………………………………………. 9
Chapter Three Results…………………………………………………………………… 10
Experiment one: Temperature × light intensity……………………. 10
Survival ratio and survival period…………………………….. 10
Budding rates…………………………………………………. 11
Strobilation ratios and rates…………………………………… 11
Ephyra production…………………………………………….. 12
Experiment two: Temperature × salinity…………………………... 13
Survival ratio and survival period…………………………….. 13
Budding rates…………………………………………………. 14
Strobilation ratios and rates…………………………………… 15
Ephyra production…………………………………………….. 16
Chapter Four Discussion……………………………………………………………….. 18
Interaction effects of temperature and light………………………... 18
Effects of temperature……………………………………………… 18
Effects of light……………………………………………………... 21
Comparisons of tropical and temperate Aurelia spp……………….. 22
Effects of salinity…………………………………………………... 23
Interactive effects of temperature and salinity……………………... 24
Effects of short term starvation……………………………………. 26
Future research……………………………………………………... 27
Conclusion………………………………………………………… 29
Reference …………………………………………………………………………… 30

Tables .................................................................................................................... 40
Table 1. Experimental conditions in combinations of temperature and light intensity during the experiment as measured with Onset data loggers……………………………………………………..

40
Table 2. The effects of temperature and light intensity on polyps were tested in combinations of temperature and light intensity............
41
Table 3. The proportion of polyps that died without strobilating in combinations of temperature and light intensity..........................
42
Table 4. The proportions of polyps strobilating in combinations of temperature and light intensity.....................................................
43
Table 5. Ephyra production per day and the proportion of total asexual reproduction of each polyp that resulted in combinations of temperature and light intensity.....................................................

44
Table 6. The effects of temperature and salinity on polyps were tested in combinations of temperature and salinity.....................................
45
Table 7. The proportion of polyps strobilating in combinations of temperature and salinity...............................................................
46
Table 8. Ephyra production per day and the proportion of total asexual reproduction of each polyp resulted in combinations of temperature and salinity...............................................................

47











Figures .................................................................................................................... 48
Figure 1. Average survival periods, pre-strobilation periods and strobilation periods, in combinations of temperature and light intensity........................................................................................

48
Figure 2. Percentages of polyps that died without strobilating in combinations of temperature and light intensity........................
49
Figure 3. The cumulative numbers of new buds in combinations of temperature and light intensity...................................................
49
Figure 4. The cumulative numbers of strobilating polyps in combinations of temperature and light intensity........................
50
Figure 5. The cumulative numbers of ephyrae in combinations of temperature and light intensity...................................................
50
Figure 6. The potential yield of ephyrae in combinations of temperature and light intensity.......................................................................
51
Figure 7. Average survival periods, pre-strobilation periods and strobilation period, in combinations of temperature and salinity........................................................................................

52
Figure 8. The cumulative numbers of new buds in combinations of temperature and salinity..............................................................
53
Figure 9. The cumulative numbers of strobilating polyps in combinations of temperature and salinity...................................
53
Figure 10. The cumulative numbers of ephyrae in combinations of temperature and salinity............................................................
54
Figure 11. The potential yield of ephyrae in combinations of temperature and salinity............................................................
54






Appendix …………………………………………………………………………… 55
Appendix 1: The row data of temperature and light intensity experiment..............................................................................
55
Appendix 2: The row data of temperature and salinity experiment..............................................................................
60

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