烏賊缺乏親代照顧的行為，其幼體孵化後直接面對龐大的掠食壓力，在這樣的情況下烏賊幼體需要孵化後立刻能使用有效的防禦技巧抵禦掠食者。虎斑烏賊(Sepia pharaonis)和歐洲常見烏賊(Sepia officinalis)的成體形態類似，然而兩者在卵莢的形態上有很大的差異；前者為透明的卵莢，後者則為黑色卵莢。根據前人研究，歐洲常見烏賊在胚胎發育晚期感官系統已發育完全且具備接收環境因子的能力，烏賊胚胎會利用其胚胎經驗改變孵化後的食物選擇偏好，但烏賊胚胎時期暴露掠食者因子後對其孵化後行為的影響仍然未知。虎斑烏賊的卵莢形態提高了卵被掠食的潛在風險，但也可能幫助虎斑烏賊的胚胎利用視覺偵測環境中的變化，增加孵化後的存活率。本研究旨在了解虎斑烏賊和歐洲常見烏賊胚胎時期的感覺經驗是否會影響孵化之後的禦敵行為，並比較兩者之間的差異。
本研究將烏賊卵於胚胎期間暴露在高掠食風險的環境，利用不同的掠食者暴露形式(視覺/嗅覺/視覺與嗅覺)，比較各組別孵化後初級及次級防禦行為的差異。結果顯示烏賊胚胎時期暴露在高掠食風險環境對於烏賊卵的孵化時間、孵化後幼體的初級及次級防禦行為皆有影響；在虎斑烏賊和歐洲常見烏賊胚胎時期的感覺經驗上，虎斑烏賊胚胎時期視覺經驗可能比嗅覺經驗重要，在歐洲常見烏賊上則是其胚胎時期接收的視覺經驗和嗅覺經驗同等重要，這樣的差異也對應到兩個物種間卵莢形質的不同，顯示烏賊卵的形質會影響其胚胎時期所接受的感官訊息比重。

Cuttlefish lack parental care, and hatchlings face high predatory risk immediately after hatching. Thus, cuttlefish hatchlings need to equip the anti-predatory ability as soon as they hatch. Pharaoh cuttlefish Sepia pharaonis and European common cuttlefish Sepia officinalis have similar morphological appearances, but their egg characteristics are distinctly different. Pharaoh cuttlefish produce eggs with transparent capsules, and European common cuttlefish produce eggs with a black envelop which is stained by ink. Previous studies have shown that sensory systems of European common cuttlefish are fully developed in late embryonic stages, and they can use these sensory systems to receive information from the environment. Moreover, European common cuttlefish can alter their food preference by their embryonic visual experience. However, there is little known about the effect of embryonic experience to predatory cues on postnatal behaviors in cuttlefish. Compared with European common cuttlefish, eggs of pharaoh cuttlefish are more vulnerable to predators, but the transparency of eggs may provide embryos an opportunity of collecting visual cues from the environment and thus increase their survival rate. The present study was aimed to understand whether cuttlefish’s embryonic experience of predators affects their postnatal defensive behaviors, and to compare the difference between pharaoh cuttlefish and European common cuttlefish.
During the embryonic exposure period, all eggs of cuttlefish were separated into seven groups: one control group and six cue exposure groups. Embryos in three groups were exposed to the predatory cues (visual cue only, chemical cue only, and combination of visual and chemical cues). Similarly, Embryos in the other three groups were exposed to the cues of non-predator. Cuttlefish were tested their primary and secondary defensive behaviors after they hatched. The result showed that early exposure to predatory cues altered the cuttlefish hatching period and postnatal defensive behavior. However, effects of embryonic sensory information were different in pharaoh cuttlefish and European common cuttlefish. Specifically, visual information is more important than chemical information for pharaoh cuttlefish embryos, and chemical information may even reduce these effects. In contrast, visual information and chemical information are equally important for European common cuttlefish embryos. These results may explain the functional difference between egg appearance of two species.

摘要 i
Abstract ii
致謝 iv
Table of Contents vii
List of Figures viii
Chapter 1 Introduction 1
Chapter 2 Materials and Methods 6
2.1 Subjects 6
2.2 Experimental design 9
2.3 Data analysis 14
Chapter 3 Results 15
3.1 Experiment 1 15
3.2 Experiment 2 18
Chapter 4 Discussion 20
References 26
Figures 30
Supplementary figures 44

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