軟絲與烏賊體色改變的週邊神經調控｜國立清華大學博碩士論文庫

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研究生：	施秉宏 Shih, Bing Hong
論文名稱：	軟絲與烏賊體色改變的週邊神經調控 Peripheral neural control of body patterning in oval squids and cuttlefish
指導教授：	焦傳金 Chiao, Chuan Chin
口試委員:	嚴宏洋 Yan, Hong Young 葉世榮 Yeh, Shin Rung
學位類別：	碩士 Master
系所名稱：	生命科學暨醫學院 - 系統神經科學研究所 Institute of Systems Neuroscience
論文出版年：	2016
畢業學年度：	105
語文別：	英文
論文頁數：	52
中文關鍵詞：	色素調控系統、週邊神經系統、體色改變
外文關鍵詞：	chromatophore system, peripheral neural system, body patterning
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現代頭足類能夠以快速改變體色來進行溝通及偽裝。此改變體色的能力主要是藉由神經對皮膚內的色素細胞直接進行控制來達成。目前在中樞神經系統中對色素細胞進行調控的神經傳導路徑已經被廣泛研究，然而在週邊神經系統的體色表現調控方式卻仍屬未知。藉由給予萊氏擬烏賊表皮L-麩胺酸、乙醯膽鹼、血清素、AMPA及NMDA，並觀察色素細胞的擴張與收縮，本研究證實由於環狀肌肉上AMPA受器及NMDA受器表現量的不同，因此可以藉由突觸後作用機制來分別調控黃色及棕色色素細胞的擴張，此外乙醯膽鹼則參與在兩種不同色素細胞調控的突觸前調節機制。在另外一項研究中則以虎斑烏賊為對象觀察色素調控系統的週邊神經發育，藉由觀察皮膚對L-麩胺酸與血清素的反應，本研究顯示色素細胞在胚胎發育至第26期後才會具有功能。而比較不同發育期的胚胎在三種不同方格背景(小方格低對比、小方格高對比、大方格高對比)的體色反應後發現，胚胎雖然在第27期之後可以對環境反應出破碎型體色，然而其色素細胞調控系統卻仍未發展完全以至於不能針對環境差異做出適當反應。綜合以上發現，本研究顯示體色調控的週邊神經系統發育較早，且同時具有突觸前及突觸後兩種調控色素細胞的作用機制。

Coleoid cephalopods are well known for their ability to rapidly change body patterns for communication and camouflage. This is primarily achieved by neural control of chromatophores in the skin. The neural connection of the chromatophore system in the central nervous system has been well studied. However, the specific mechanism of the peripheral neural control of chromatophores remains unknown. To determine the mechanism of the peripheral pathway, the several neurotransmitters such as L-Glutamate (L-Glu), serotonin, acetylcholine (ACh), AMPA, and NMDA were applied to the skin of oval squids (Sepioteuthis lessoniana) in vivo and the corresponding chromatophore expansion and retraction were recorded. The results showed that AMPA- and NMDA-like receptors expressed on radial muscles are involved in a postsynaptic mechanism to differentially control yellow and brown chromatophore expansions, and ACh is involved in a presynaptic mechanism of chromatophore control. In a separate study, cuttlefish embryos (Sepia pharaonis) were used to examine the peripheral development of the chromatophore system. The results showed that chromatophores are functional at embryonic stage 26 as evidence of its response to L-Glu and serotonin applications in vitro. In addition, three different checkboard backgrounds (small low-contrast, small high-contrast, and large high-contrast) were used to examine cuttlefish embryo’s ability of body patterning development. The results showed that cuttlefish could express some disruptive components at stage 27, though the full function of body pattern control was not developed completely before hatching. Taken together, this study suggests that the peripheral neural control of the chromatophore system develops early and is regulated by both pre- and post-synaptic mechanisms with various neurotransmitters.

中文摘要    i
Abstract    ii
誌謝    iv
Contents    vii
Chapter 1    Introduction    1
1.1.    Dynamic body pattern of cephalopods    1
1.2.    The elements involved in body patterns    1
1.3.    Structure of chromatophores    2
1.4.    Neurotransmitter of chromatophores    3
1.5.    Development of chromatophores and its control    6
1.6.    Specific aims    7
Chapter 2    Materials and methods    8
2.1.    Subjects    8
2.2.    Experimental design    8
2.3.    Data analysis    12
Chapter 3    Results    17
3.1.    Peripheral neural control of body patterning in oval squids    17
3.2.    Development of body patterning control in cuttlefish    20
Chapter 4    Discussion    23
4.1.    Peripheral neural control of body patterning in oval squids    23
4.2.    Neural development of body patterning in cuttlefish    26
4.3.    Functional development of body patterning in embryonic cuttlefish    27
4.4.    Conclusion    28
References    30
Figures    35
Figure 1. Side view of the experimental setup for the squid experiment    35
Figure 2. Side view of the experimental setup for the cuttlefish experiment    36
Figure 3. The experimental design for assessing body pattern control in   cuttlefish    37
Figure 4. The comparison of the MetaMorph method and ImageJ method    38
Figure 5. Chromatic components used in grading disruptive pattern of    cuttlefish    38
Figure 6. Chromatophore expansion induced by ectopic application of L-Glutamate on the skin of an oval squid    39
Figure 7. The speed of brown chromatophore expansion upon L-Glutamate application varies across different chromatophores and skins    40
Figure 8. Chromatophore retraction induced by ectopic application of serotonin on the skin of an oval squid    41
Figure 9. The speed of brown chromatophore retraction upon serotonin application varies across different chromatophores and skins.    42
Figure 10. Chromatophore expansion induced by ectopic application of acetylcholine on the skin of an oval squid    43
Figure 11. The speed of brown chromatophore expansion upon acetylcholine application varies across different skins    44
Figure 12. Chromatophore expansion induced by ectopic application of AMPA or NMDA on the skin of an oval squid    45
Figure 13. The speeds of brown and yellow chromatophore expansion upon AMPA application (100 mM) in three different samples are consistent    46
Figure 14. The speeds of brown and yellow chromatophore expansion upon NMDA application (100 mM) vary across four different samples    47
Figure 15. Expansion and retraction of brown chromatophores upon L-Glutamate and serotonin applications, respectively, in cuttlefish embryos of developmental stages 25-30    48
Figure 16. The speed of brown chromatophore response to drugs varies in different developmental stages    49
Figure 17. Cuttlefish could express some disruptive components after developmental stage 27    50
Figure 18. The functional development of disruptive pattern is not completed even after hatching for 1 week    51
Figure S1. The score of each component for cuttlefish on different backgrounds at different developmental stages    52


                                

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