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Chapter 4: Cortical Organization
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Visual processing streams
Striate cortex: simple feature detection But how are objects spatially represented in the visual system? Are signals representing different objects in a scene go to different places in the brain?
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Retinotopic map 外界物體刺激網膜的空間位置,與物體的大腦皮質表徵,具有空間的對應性
Coortical magnification Figure 4.1 A person looking at a tree, showing how points A, B, C, and D are imaged on the retina and where these retinal activations cause activity in the brain. Although the distances between A and B and between C and D are about the same on the retina, the distance between A and B is much greater on the cortex. This is an example of cortical magnification, in which more space is devoted to areas of the retina near the fovea. Figure 4-1 p78
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大腦空間分佈地圖 側膝核(lateral geniculate nuclceus, LGN)
retinotopic map網膜光刺激部位與LGN的激發部位之間具有空間分佈對應性
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皮質放大因子(cortical magnification factor):中央小窩只占網膜的0.01%,但在視皮質卻佔了8-10%的區域
Figure 4.2 The magnification factor in the visual system. The small area of the fovea is represented by a large area on the visual cortex. Figure 4-2 p78
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Brain Imaging Techniques
運用大腦造影技術瞭解皮質放大 Positron emission tomography (PET) Functional agnetic resonance imaging (fMRI)
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物體在手中 物體放在手中操弄 操弄物體 Figure 4.3 A person in a brain scanning apparatus.
Figure 4-3 p79
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所以對於細節的區辨能力除與眼球特性有關外,也與腦中地圖有關
人類視皮質放大因子 有個別差異 所以對於細節的區辨能力除與眼球特性有關外,也與腦中地圖有關 Figure 4.4 (a) Red and blue areas show the extent of stimuli that were presented while a person was in an fMRI scanner. (b) Red and blue indicate areas of the brain activated by the stimulation in (a). Figure 4-4 p79
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Figure 4. 5 Demonstration of the magnification factor
Figure 4.5 Demonstration of the magnification factor. A person looks at the red spot on the text on the left. The area of brain activated by each letter of the text is shown on the right. The arrows point to the letter a in the text on the left, and the area in the brain activated by the a on the right. Figure 4-5 p80
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皮質的柱狀(column)結構 Orientation columns Ocular dominance columns
Location columns Hypercolumns by Hubel and Wiesel 對落在某個網膜位置的視覺訊息作完整的分析
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Figure 4. 7 Orientation columns
Figure 4.7 Orientation columns. All of the cortical neurons encountered along track A respond best to horizontal bars (indicated by the red lines cutting across the electrode track). All of the neurons along track B respond best to bars oriented at 45 degrees. Figure 4-7 p81
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Figure 4.8 If an electrode is inserted obliquely into the cortex, it crosses a sequence of orientation columns. The preferred orientation of neurons in each column, indicated by the bars, changes in an orderly way as the electrode crosses the columns. The distance the electrode is advanced is exaggerated in this picture. Figure 4-8 p81
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Orientation columns Also ocular dominance columns
Figure 4.9 A location column that contains the full range of orientation columns. A column such as this, which contains a full array of orientation columns, was called a hypercolumn by Hubel and Wiesel. A column such as this receives information about all possible orientations that fall within a small area of the retina. Also ocular dominance columns Figure 4-9 p81
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How Do Feature Detectors Respond to a Scene?
Tiling location columns working together to cover the entire visual field Each location column is focused on a specific area of the scene
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Figure 4. 10 (a) A scene from the Pennsylvania woods
Figure 4.10 (a) A scene from the Pennsylvania woods. (b) Focusing in on part of a tree trunk. A, B, and C represent the parts of the tree trunk that fall on receptive fields in three areas of the retina. Figure 4.11 (a) Receptive fields for the three sections of the tree trunk from Figure 4.10b. The neurons associated with each of these receptive fields are in different location columns. (b) Three location columns in the cortex. Neurons that fire to the tree trunk’s orientation are within the orange areas of the location column. Figure 4-10 p82 Figure 4-11 p82
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Figure 4.12 The yellow circles and ellipses superimposed on the forest scene each represent an area that sends information to one location column in the cortex. The way these location columns cover the entire receptive field is called tiling. Figure 4-12 p82
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訊息流的路徑—是什麼?在哪裡?如何作? Two parallel streams in extrastriate cortex (Ungerleider & Mishkin, 1982) Figure 4.14 The monkey cortex, showing the what, or ventral, pathway from the occipital lobe to the temporal lobe, and the where, or dorsal, pathway from the occipital lobe to the parietal lobe. The where pathway is also called the how pathway. Figure 4-14 p84
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物體區辨作業 地標區辨作業 Figure 4.13 The two types of discrimination tasks used by Ungerleider and Mishkin. (a) Object discrimination: Pick the correct shape. Lesioning the temporal lobe (shaded area) makes this task difficult. (b) Landmark discrimination: Pick the food well closer to the cylinder. Lesioning the parietal lobe makes this task difficult. Figure 4-13 p83
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Where pathway? What pathway? 運用切除(ablation)法
測量知覺—切除(破壞,冷凍)腦區域—訓練動物,再測量知覺 切除部分頂葉(parietal lobe)與側葉(temporal lobe) 結果頂葉切除猴子對於地標區辨產生困難,側葉切除猴子對於物體區辨產生困難 Where pathway? What pathway? 背側 腹側 19
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背側與腹側路徑的細節
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Where: locating objects, the dorsal pathway M cell in retina Magno
Where: locating objects, the dorsal pathway M cell in retina Magno. layers in LGN V1V2V3MTParietal What: identifying objects, the ventral pathway P cell in retina Parvo. Layers in LGNV1 V2 V4 IT Information flows in both directions Pathways are interconnected
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Streams for Information About What and How
What and how stream (Milner & Goodale, 1995) the dorsal stream provides information about how to direct action toward objects (action pathway) 神經生理證據 猴子頂葉有看見物體並試圖觸及物體時才有反應的神經元 神經心理學(Neuropsychology)證據 運用雙重分離(double dissociation)邏輯 (功能)分離(dissociation) 某個功能存在,而另一個功能受損
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單一分離(single dissociation) 某甲產生「功能X存在,而功能Y受損」,顯示功能X與Y有不同,但未必彼此完全獨立的機制
雙重分離(double dissociation) 某甲產生「功能X存在,而功能Y受損」,某乙產生「功能Y存在,而功能X受損」,顯示功能X與Y有不同的機制,且彼此也完全獨立
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Patient D.F. (Milner & Goodale, 1995)
一氧化碳中毒導致腹側路徑受損 visual form agnosia 無法把卡片match細縫的方位 但可以「寄信」 Figure 4.16 Performance of D.F. and a person without brain damage on two tasks: (a) judging the orientation of a slot; and (b) placing a card through the slot. See text for details. Figure 4-16 p85
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單一 __分離 雙重分離— 顯示「方位判斷」和「視覺/動作協調」為不同機制 顯示「方位判斷」和「視覺/動作協調」為__機制
有其他有相反狀況的病人 獨立
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Behavior of People Without Brain Damage
Evidence for the separation of perception and action in non-brain-damage subjects. Ganel et al. (2008) length estimation task grasping task
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(a) The size illusion used by Ganel and coworkers (2008) in which line 2 looks longer than line 1. The numbers were not present in the display seen by the subjects. (b) The two vertical lines from (a), showing that line 2 is actually shorter than line 1. (c) Subjects in the experiment adjusted the space between their fingers either to estimate the length of the lines (length estimation task) or to reach toward the lines to grasp them (grasping task). The distance between the fingers is measured by sensors on the fingers. (d) Results of the length estimation and grasping tasks in the Ganel et al. experiment. The length estimation task indicates the illusion, because the shorter line (line 2) was judged to be longer. In the grasping task, subjects separated their fingers more for the longer line (line 1), which was consistent with the physical lengths of the lines. Figure 4-17 p86
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視覺處理的模組化(Modularity)
模組(module) 某個腦結構中多數神經元選擇性地處理具有某種特性的訊息,那麼這個腦結構為處理此種特性的一個模組 Fusiform face area (FFA) 臉孔辨識的模組
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Figure 4.18 How a neuron in a monkey’s temporal lobe responds to a few stimuli. This neuron responds best to a circular disc with a thin bar (a). Figure 4-18 p87
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Figure 4.19 Size of response of a neuron in the monkey’s IT cortex that responds to face stimuli but not to nonface stimuli. (R Figure 4-19 p87
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Figure 4. 20 Results of the Tsao et al
Figure 4.20 Results of the Tsao et al. (2006) experiment in which activity of neurons in the monkey’s temporal lobe was recorded in response to faces, other objects, and a scrambled stimulus. Figure 4-20 p88
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臉孔失認症(prosopagnosia)
LH (Farah, 2000) WJ(McNeil & Warrington, 1993)
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Figure 4.21 (a) The parahippocampal place area (PPA) is activated by places (top row) but not by other stimuli (bottom row). (b) The extrastriate body area (EBA) is activated by bodies (top), but not by other stimuli (bottom). Figure 4-21 p88
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Figure 4.22 fMRI responses of the human brain to various types of stimuli: (a) areas that were most strongly activated by houses, faces, and chairs; (b) all areas activated by each type of stimulus. Figure 4-22 p89
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The visual system is organized both spatially and functionally Spatial
Retinotopic map Becomes weaker as moving towards higher cortical regions Functional What/where, how Modules for specific objects
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Where Vision Meets Memory
MTL structures are extremely important in memory H.M. hippocampus
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Figure 4.23 (a) Location of the hippocampus and some of the other structures that were studied by Quiroga and coworkers (2005). (b) Some of the stimuli that caused a neuron in the hippocampus to fire. Figure 4-23 p90
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Figure 4.24 Activity of a neuron in the MTL of an epilepsy patient as he remembered the things indicated below the record. A response occurs when the person remembered The Simpsons TV program. Earlier, this neuron had been shown to respond to viewing a video clip of The Simpsons. Figure 4-24 p90
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經驗 神經元按照視覺環境特性發展成為最能適應環境刺激的模式—experience-dependent plasticity, eg., visual word form area eg., selective rearing eg., expertise related activation in FFA
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Figure 4. 25 (a) Greeble stimuli used by Gauthier
Figure 4.25 (a) Greeble stimuli used by Gauthier. Participants were trained to name each different Greeble. (b) Brain responses to Greebles and faces before and after Greeble training. Figure 4-25 p91
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