虎斑烏賊的視覺認知能力：視覺分辨、視覺類化及視覺填補

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研究生：	林宜蓉 Lin, I-Rong
論文名稱：	虎斑烏賊的視覺認知能力：視覺分辨、視覺類化及視覺填補 Visual Cognition in Cuttlefish Sepia Pharaonis: Discrimination, Generalization, and Amodal Completion
指導教授：	焦傳金 Chiao, Chuan-Chin
口試委員:	陳建中楊恩誠
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：	2012
畢業學年度：	100
語文別：	英文
論文頁數：	59
中文關鍵詞：	頭足類動物、認知能力、操作制約
外文關鍵詞：	cephalopod, cognitive ability, operant conditioning
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頭足類動物（包括章魚、魷魚及烏賊）擁有複雜的神經系統，且被認為是具有相當程度認知能力的無脊椎動物。頭足類大腦內之神經細胞數量高達五億個，其中負責處理視覺訊息的腦區佔三分之二以上，加以具有空間解析度良好的單眼結構，因此頭足類動物的視覺系統非常發達。頭足類動物高度倚賴視覺，舉凡日常生活中之生存行為、至複雜的偽裝及溝通行為皆與視覺密切相關，因此視覺在頭足類動物認知功能的展現上，應扮演著極為重要之角色。在頭足類大腦中，視覺訊號究竟如何被處理、感知層面的訊息是如何被建立，尚需要更多實驗來檢視。本篇研究希望藉由行為實驗來探究他們的視覺認知能力，期許能以此作為釐清頭足類動物視覺運作機制之洞見。
我們利用操作制約的方式訓練虎斑烏賊分辨魚、蝦圖案，所使用之實驗裝置主要功能是分隔出兩個獨立區域，烏賊必須進入放置特定圖案之區域或以觸手攻擊此圖案，才可獲得食物作為獎賞，而通過訓練的條件是連續兩天之選擇正確率皆達80%。烏賊通過訓練後，需接受一檢視其視覺分辨能力的測驗，在此測驗中，撤換未與獎賞作連結之圖片，改為螃蟹圖案。隨後，進行一連串的測驗來檢視其視覺類化能力，測驗用圖片為魚、蝦圖案之改編圖案(例如：縮小型、素描型…)。最後檢視其視覺填補能力，測驗用圖片為部分遮蔽的魚、蝦圖案。
實驗結果顯示，烏賊可以成功地分辨魚、蝦及螃蟹圖案，能對魚、蝦圖案之縮小型、素描型與剪影做出正確反應，且可辨識部分遮蔽的魚、蝦圖案。因此我們認為虎斑烏賊具有視覺的分辨、類化及填補能力。

Modern cephalopods are the most intelligent invertebrates with keen vision which is important for them to move rapidly for successful predation, camouflage for danger avoidance and communicate in complex ways with their body patterns. Therefore, vision might impact the cognition of cephalopods to some degree, but knowledge about how visual information is processed in the perceptual level is still not completely understood. In this work, we investigated the intelligence developing in a different way from vertebrates by observing the cognitive behaviors of cuttlefish. Results from this research are expected to give insights into the visual processing mechanism of cuttlefish.
Cuttlefish (Sepia pharaonis) were trained to discriminate figures of shrimp and fish. Training for discrimination was based on operant conditioning and was given in the home tanks of each animal. Experimental apparatus was composed of two separated regions, and cuttlefish had to enter the rewarded area where the rewarded figure was presented or even to strike the rewarded figure with their tentacles for food. After cuttlefish achieved the learning criteria, a discrimination task was conducted to confirm learning. The non-rewarded figure was replaced with a crab figure in the discrimination test. Several transfer tests utilizing novel stimuli which are modified versions of training figures followed up the discrimination test. The outcome of visual processing in cuttlefish was examined in these transfer tasks.
The results show that cuttlefish are capable of discriminating figures of fish, shrimp and crab. Furthermore, they do generalize the small-scale, sketch, and silhouette of the training figures, and respond properly, though the response strength is lower than that in training and individual differences exist in performance. In addition, cuttlefish also recognize the fragmented image of the training figures. These findings suggest that cuttlefish have the ability of discrimination, generalization, and amodal completion.

Table of Contents
摘要    i
Abstract    ii
誌謝    iii
Table of Contents    v
List of Tables    vii
List of Figures    viii
Chapter 1    Introduction    1
1.1    Brain and cognitive functions in cephalopods    1
1.2    Visual cognition in cephalopods    2
1.2.1    Visual discrimination    4
1.2.2    Visual generalization    6
1.2.3    Visual completion    7
1.3    Specific aims    9
Chapter 2    Materials and Methods    11
2.1    Subjects    11
2.2    Apparatus    12
2.3    Visual stimuli    13
2.4    Experimental procedure    14
2.4.1    Discrimination training    14
2.4.2    Discrimination Task    15
2.4.3    Transfer Tasks    16
2.5    Scoring    17
2.6    Data analysis    17
Chapter 3    Results    18
3.1    Discrimination training    18
3.2    Visual discrimination    18
3.3    Visual generalization    19
3.4    Amodal completion    23
Chapter 4    Discussion    25
4.1    Visual association learning in cuttlefish    25
4.2    Prey preference and visual discrimination    26
4.3    Object perception and visual generalization    27
4.4    Visual completion and visual generalization    31
4.5    Individual difference exists in visual cognition of cuttlefish    32
4.6    Conclusions    33
References    34
Tables and Figures    41

List of Tables
Table 1. Number of trials for each cuttlefish to reach criteria.    41
Table 2. The statistical results of individual cuttlefish in all tasks.    42
Table 3. The statistical results of all cuttlefish in the transfer tasks.    43

List of Figures
Figure 1. Examples of visual completion.    44
Figure 2. The experimental apparatus.    45
Figure 3. Visual stimuli used in the discrimination training and discrimination task.    46
Figure 4. Visual stimuli used in the generalization tasks.    47
Figure 5. Visual stimuli used in the amodal completion tasks.    48
Figure 6. Learning curves of cuttlefish in the discrimination training.    49
Figure 7. Cuttlefish can discriminate among figures of fish, shrimp, and crab.    50
Figure 8. Cuttlefish can generalize across different prey sizes.    51
Figure 9. Cuttlefish can generalize from the original figures to the sketched figures.    52
Figure 10. Cuttlefish may generalize from the original figures to the reduced-contrast figures.    53
Figure 11. Cuttlefish cannot generalize from the original figures to the contoured figures.    54
Figure 12. Cuttlefish can generalize from the original figures to the black silhouette.    55
Figure 13. Cuttlefish cannot generalize from the original figures to the white silhouette.    56
Figure 14. Cuttlefish can amodally complete the posteriorly occluded figures.    57
Figure 15. Cuttlefish can amodally complete the anteriorly occluded figures.    58
Figure 16. Cuttlefish can amodally complete the partially occluded figures.    59

                                

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