大腦視葉是頭足類的中樞神經系統中占有最大體積以及扮演重要角色的腦區，它負責處理來自眼球的視覺訊息、調控體色表現以及控制運動行為等。雖然我們對視葉的一般形態已經有基本的了解，但目前仍缺乏詳細且系統化的形質定量來描述頭足類視葉發育的形態變化。在此研究中，我們對虎斑烏賊 (Sepia pharaonis) 與萊氏擬烏賊 (Sepioteuthis lessoniana) 的大腦視葉以不同的影像技術─包括核磁共振影像、微電腦斷層掃描與組織螢光顯微影像─進行觀察。此外，一個稀有的大王魷魚 (Architeuthis dux)的大腦視葉也利用核磁共振影像方式進行定量研究，並與虎斑烏賊及萊氏擬烏賊的視葉做比較，以了解頭足類視葉在物種間的異同。
虎斑烏賊視葉的核磁共振掃描顯示在髓質中稱為「細胞小島 (cell islands)」的構造其實是連續的樹枝狀結構，而在虎斑烏賊跟萊氏擬烏賊視葉的神經構造定量中也發現，皮質跟髓質都會隨著動物從胚胎到成年的發育穩定地擴大，但是在髓質中心 (tangential zone) 的細胞小島密度卻會隨著發育持續降低，更有趣的是，在視葉中的神經細胞核會隨著發育而增大，這些發現都表示大腦視葉在動物的生命週期中會持續地生長並重組，而這些形態上的改變提供了虎斑烏賊及萊氏擬烏賊從孵化後到成體間發生視覺行為改變的有利證據。
除此之外，在大王魷魚的研究中，我們發現大王魷的視葉與眼球體積比明顯地小於虎斑烏賊及萊氏擬烏賊，但大王魷大腦視葉的視覺訊息處理區域（意即皮質）發育十分良好，相較而言，大王魷視葉的視覺行為整合中樞（意即髓質）與其他頭足類相比明顯小很多。這些結果支持了大王魷的腦結構並非與其巨大眼睛成等比例演化，且因為不需要如淺海頭足物種那樣展現複雜的視覺行為（例如：虎斑烏賊及萊氏擬烏賊的動態體色變化與視覺溝通行為），因此視葉髓質的體積相對較小。

The optic lobe is the largest brain area within the central nervous system of cephalopods and it plays important roles in the processing of visual information, the regulation of body patterning, and locomotive behavior. Despite the understanding in the general morphology of the optic lobe, a detailed and systematic characterization of its morphological development has never been reported. In the present study, the development of the optic lobe in the cuttlefish Sepia pharaonis and the oval squid Sepioteuthis lessoniana were carefully examined using various imaging techniques, including magnetic resonance imaging (MRI), micro computed tomography (micro-CT), and histological fluorescent imaging. In addition, the optic lobe from a rare sample of the second largest cephalopod, the giant squid Architeuthis dux, was also studied with MRI and compared with those of cuttlefish and oval squids.
The MRI scan of a cuttlefish optic lobe revealed that the so called “cell islands” in the medulla of the cephalopod’s optic lobe (Young, 1962; 1974) are in fact a contiguous tree-like structure. Quantification of the neural organizational development of optic lobes in both cuttlefish and oval squids showed that the cortex and the medulla expand steadily from the embryonic stage to adulthood, but the density of cell islands in the tangential zone of the optic lobe decrease continuously in parallel. Interestingly, the size of the nuclei of cells within the optic lobe increases throughout development. These findings suggest that the optic lobe undergoes continuous growth and reorganization throughout the animal’s life. These morphological changes in the optic lobe are likely to be responsible for changes in the visuomotor behavior of oval squids and cuttlefish from hatching to adulthood.
In the study of the giant squid, it was evident that the volume ratio of the optic lobe to the eye in the giant squid is much smaller than that in the oval squid and the cuttlefish. Moreover, the visual information processing area in cephalopods, the cortex, is well developed in the giant squid. In comparison, the optic lobe medulla, the visuomotor integration center in cephalopods, is much less developed in the giant squid than other species. Collectively, the results support that the brain of giant squids has not evolved proportionally for performing complex tasks when compared with shallow water cephalopod species, such as the dynamic body pattering and visual communicative behavior in oval squids and cuttlefish.

致謝 4
中文摘要 6
Abstract 7
Chapter Ⅰ. Introduction 11
1-1 Overview 12
1-2 Cuttlefish are a good model animal for studying the development of the optic lobe 14
1-3 Oval squids are also a good model animal for studying the development of the optic lobe 15
1-4 Giant squids are unique for comparing the optic lobe morphology with other cephalopod species 16
1-5 Specific aims for the present study 17
Chapter Ⅱ. Materials and Methods 18
2-1 Animals 19
2-2 Structural imaging (MRI/micro-CT) 20
2-3 Histology and image acquisition 23
2-4 Measurement and analysis 25
Chapter Ⅲ. Neural organization of the optic lobe changes steadily from late embryonic stage to adulthood in cuttlefish Sepia pharaonis 29
3-1 Results 30
3-2 Discussion 36
Chapter Ⅵ. Morphological changes of the optic lobe from late embryonic to adult stages in oval squids Sepioteuthis lessoniana 45
4-1 Results 46
4-2 Discussion 48
Chapter Ⅴ. Mismatch between the eye and the optic lobe in the giant squid 54
5-1 Results 55
5-2 Discussion 56
Chapter Ⅵ. Overall conclusion and general discussion 58
References 61
Tables 71
Figures 75
Movies 102

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