视觉注意的时间结构和动态神经机制
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摘要
把有限的心理和神经资源选择性地根据当前环境或任务来分配给某些而不是所有过程,是通过注意来完成的。以往的大部分有关视觉注意的研究都集中在注意是如何在空间维度和特征维度进行分配的问题上,而近来的一系列研究则从时间动态这个角度揭示了视觉注意中的一些新现象和新机制。本文将从以下四个方面综述时间这个基本维度在视觉注意中的表现、作用及其神经机制:(1)注意对外部刺激动态结构的追踪(注意动态理论),(2)注意的内秉动态特性,(3)基于时间组织的多物体加工,(4)视觉动态和经典注意现象的相互关系。我们认为,注意的时间组织和动态结构是注意中非常重要和基本的方面,对其的研究将为注意理论的一些长久以来的争论和难点提供新的角度和解决方案。本文进一步提出了视觉注意研究领域亟待探索和研究的新方向和问题。
Attention shapes what we see and what we act upon by allocating limited resources to certain parts of visual display in a selective and adaptive manner. While most previous studies in visual attention mainly focused on the attentional distribution over space or features, recent studies have revealed that temporal dynamics also plays a crucial function in visual attention. This paper reviews the representation, function and neural mechanism of temporal dynamics in visual attention from the following four aspects:(1) Tracking dynamic structure of external stimulus by attention;(2) Intrinsic dynamic characteristics of attention;(3) Time-based multiple object representation;(4) Relationship between visual dynamics and classical attentional phenomena. We propose that the dynamic structure and temporal organization are fundamental to visual attention, and the research on it might provide new solutions to many unresolved issues in visual attention research.
Attention shapes what we see and what we act upon by allocating limited resources to certain parts of visual display in a selective and adaptive manner. While most previous studies in visual attention mainly focused on the attentional distribution over space or features, recent studies have revealed that temporal dynamics also plays a crucial function in visual attention. This paper reviews the representation, function and neural mechanism of temporal dynamics in visual attention from the following four aspects:(1) Tracking dynamic structure of external stimulus by attention;(2) Intrinsic dynamic characteristics of attention;(3) Time-based multiple object representation;(4) Relationship between visual dynamics and classical attentional phenomena. We propose that the dynamic structure and temporal organization are fundamental to visual attention, and the research on it might provide new solutions to many unresolved issues in visual attention research.
引文
1 Ester EF,Fukuda K,May LM,Vogel EK,Awh E.Evidence for a fixed capacity limit in attending multiple locations.Cogn Affect Behav Neurosci 2014;14(1):62-77.
2 Pylyshyn ZW,Storm RW.Tracking multiple independent targets:Evidence for a parallel tracking mechanism.Spat Vis1988;3(3):179-197.
3 Carrasco M.Visual attention:the past 25 years.Vision Res2011;51(13):1484-1525.
4 Posner MI,Snyder CR,Davidson BJ.Attention and the detection of signals.J Exp Psychol Gen 1980;109(2):160-174.
5 Posner MI.Orienting of attention.Q J Exp Psychol 1980;32(1):3-25.
6 Eriksen CW,Yeh Y.Allocation of attention in the visual field.J Exp Psychol Hum Percept Perform 1985;11(5):583-597.
7 Müller MM,Malinowski P,Gruber T,Hillyard SA.Sustained division of the attentional spotlight.Nature 2003;424(6946):309-312.
8 McMains SA,Somers DC.Multiple spotlights of attentional selection in human visual cortex.Neuron 2004;42(4):677-686.
9 Jans B,Peters JC,De Weerd P.Visual spatial attention to multiple locations at once:The jury is still out.Psychol Rev2010;117(2):637-682.
10 Desimone R,Duncan J.Neural mechanisms of selective visual attention.Annu Rev Neurosci 1995;18:193-222.
11 Reynolds JH,Heeger DJ.The normalization model of attention.Neuron 2009;61:168-185.
12 Large EW,Jones MR.The dynamics of attending:How people track time-varying events.Psychol Rev 1999;106(1):119-159.
13 VanRullen R.Perceptual cycles.Trends Cogn Sci 2016;20(10):723-735.
14 Schroeder CE,Lakatos P.Low-frequency neuronal oscillations as instruments of sensory selection.Trends Neurosci2009;32(1):9-18.
15 Jones MR,Boltz M.Dynamic attending and responses to time.Psychol Rev 1989;96(3):459-491.
16 Hickok G,Farahbod H,Saberi K.The rhythm of perception:entrainment to acoustic rhythms induces subsequent perceptual oscillation.Psychol Sci 2015;26(7):1006-1013.
17 Patel AD,Iversen JR.The evolutionary neuroscience of musical beat perception:the Action Simulation for Auditory Prediction(ASAP)hypothesis.Front Syst Neurosci 2014;8:57.
18 Spaak E,de Lange FP,Jensen O.Local entrainment of alpha oscillations by visual stimuli causes cyclic modulation of perception.J Neurosci 2014;34(10):3536-3544.
19 Fries P,Reynolds JH,Rorie AE,Desimone R.Modulation of oscillatory neuronal synchronization by selective visual attention.Science 2001;291(5508):1560-1563.
20 Womelsdorf T,Fries P,Mitra PP,Desimone R.Gamma-band synchronization in visual cortex predicts speed of change detection.Nature 2006;439(7077):733-736.
21 Gregoriou GG,Gotts SJ,Zhou H,Desimone R.High-frequency,long-range coupling between prefrontal and visual cortex during attention.Science 2009;324(5931):1207-1210.
22 Lakatos P,Karmos G,Mehta AD,Ulbert I,Schroeder CE.Entrainment of neuronal oscillations as a mechanism of attentional selection.Science 2008;320(5872):110-113.
23 Lakatos P,Barczak A,Neymotin SA,McGinnis T,Ross D,Javitt DC,O’Connell MN.Global dynamics of selective attention and its lapses in primary auditory cortex.Nat Neurosci 2016;19(12):1707-1717.
24 VanRullen R,Koch C.Is perception discrete or continuous?Trends Cogn Sci 2003;7(5):207-213.
25 Busch NA,Dubois J,VanRullen R.The phase of ongoing EEG oscillations predicts visual perception.J Neurosci2009;29(24):7869-7876.
26 Busch NA,VanRullen R.Spontaneous EEG oscillations reveal periodic sampling of visual attention.Proc Natl Acad Sci U S A 2010;107(37):16048-16053.
27 Myers NE,Stokes MG,Walther L,Nobre AC.Oscillatory brain state predicts variability in working memory.J Neurosci2014;34(23):7735-7743.
28 DuguéL,Marque P,VanRullen R.The phase of ongoing oscillations mediates the causal relation between brain excitation and visual perception.J Neurosci 2011;31(33):11889-11893.
29 Drewes J,VanRullen R.This is the rhythm of your eyes:The phase of ongoing electroencephalogram oscillations modulates saccadic reaction time.J Neurosci 2011;31(12):4698-4708.
30 Mathewson KE,Gratton G,Fabiani M,Beck DM,Ro T.To see or not to see:Prestimulus phase predicts visual awareness.J Neurosci 2009;29(9):2725-2732.
31 Samaha J,Postle BR.The speed of alpha-band oscillations predicts the temporal resolution of visual perception.Curr Biol 2015;25(22):2985-2990.
32 Wutz A,Melcher D,Samaha J.Frequency modulation of neural oscillations according to visual task demands.Proc Natl Acad Sci U S A 2018;115(6):1346-1351.
33 Buschman TJ,Kastner S.From behavior to neural dynamics:An integrated theory of attention.Neuron 2015;88(1):127-144.
34 Landau AN,Fries P.Attention samples stimuli rhythmically.Curr Biol 2012;22(11):1000-1004.
35 Fiebelkorn IC,Saalmann YB,Kastner S.Rhythmic sampling within and between objects despite sustained attention at a cued location.Curr Biol 2013;23(24):2553-2558.
36 Lamy D,Egeth H.Object-based selection:the role of attentional shifts.Percept Psychophys 2002;64:52-66.
37 Song K,Meng M,Chen L,Zhou K,Luo H.Behavioral oscillations in attention:Rhythmicαpulses mediated throughθband.J Neurosci 2014;34(14):4837-4844.
38 Klimesch W,Sauseng P,Hanslmayr S.EEG alpha oscillations:The inhibition-timing hypothesis.Brain Res Rev 2007;53(1):63-88.
39 Huang Y,Chen L,Luo H.Behavioral oscillation in priming:Competing perceptual predictions conveyed in alternating theta-band rhythms.J Neurosci 2015;35(6):2830-2837.
40 Davidson MJ,Alais D,van Boxtel JJA,Tsuchiya N.Attention periodically samples competing stimuli during binocular rivalry.Elife 2017.pii:e40868.
41 Tomassini A,Spinelli D,Jacono M,Sandini G,Morrone MC.Rhythmic oscillations of visual contrast sensitivity synchronized with action.J Neurosci 2015;35(18):7019-7029.
42 Ho HT,Leung J,Burr DC,Alais D,Morrone MC.Auditory sensitivity and decision criteria oscillate at different frequencies separately for the two ears.Curr Biol 2017;27(23):3643-3649.e3.
43 Wang Y,Luo H.Behavioral oscillation in face priming:Prediction about face identity is updated at a theta-band rhythm.Prog Brain Res 2017;236:211-224.
44 Irwin D,Gordon R.Eye movements,attention and trans-saccadic memory.Vis Cogn 1998;5(1-2):127-155.
45 VanRullen R.Visual attention:A rhythmic process?Curr Biol 2013;23(24):R1110-R1112.
46 VanRullen R,Carlson T,Cavanagh P.The blinking spotlight of attention.Proc Natl Acad Sci U S A 2007;104(49):19204-19209.
47 Landau AN,Schreyer HM,van Pelt S,Fries P.Distributed attention is implemented through theta-rhythmic gamma modulation.Curr Biol 2015;25(17):2332-2337.
48 DuguéL,Roberts M,Carrasco M.Attention reorients periodically.Curr Biol 2016;26(12):1595-1601.
49 Jia J,Liu L,Fang F,Luo H.Sequential sampling of visual objects during sustained attention.PLoS Biol 2017;15(6):e2001903.
50 Buschman TJ,Miller EK.Serial,covert shifts of attention during visual search are reflected by the frontal eye fields and correlated with population oscillations.Neuron 2009;63(3):386-396.
51 Treisman AM,Gelade G.A feature-integration theory of attention.Cognit Psychol 1980;12:97-136.
52 DuguéL,Marque P,VanRullen R.Theta oscillations modulate attentional search performance periodically.J Cogn Neurosci 2015;27(5):945-958.
53 DuguéL,McLelland D,Lajous M,VanRullen R.Attention searches nonuniformly in space and in time.Proc Natl Acad Sci U S A 2015;112(49):15214-15219.
54 Koch C,Ullman S.Shifts in selective visual attention:towards the underlying neural circuitry.Hum Neurobiol1985;4(4):219-227.
55 Foulsham T,Underwood G.What can saliency models predict about eye movements?Spatial and sequential aspects of fixations during encoding and recognition.J Vis 2008;8(2):6.1-17.
56 Wang W,Chen C,Wang Y,Jiang T,Fang F,Yao Y.Simulating human saccadic scanpaths on natural images.IEEE Conference on Computer Vision and Pattern Recognition(CVPR);Colorado,USA,2011.
57 Klein RM.Inhibition of return.Trends Cogn Sci 2000;4(4):138-147.
58 Rollenhagen JE,Olson CR.Low-frequency oscillations arising from competitive interactions between visual stimuli in macaque inferotemporal cortex.J Neurophysiol 2005;94(5):3368-3387.
59 Kienitz R,Schmiedt JT,Shapcott KA,Kouroupaki K,Saunders RC,Schmid MC.Theta rhythmic neuronal activity and reaction times arising from cortical receptive field interactions during distributed attention.Curr Biol 2018;28(15):2377-2387.
2 Pylyshyn ZW,Storm RW.Tracking multiple independent targets:Evidence for a parallel tracking mechanism.Spat Vis1988;3(3):179-197.
3 Carrasco M.Visual attention:the past 25 years.Vision Res2011;51(13):1484-1525.
4 Posner MI,Snyder CR,Davidson BJ.Attention and the detection of signals.J Exp Psychol Gen 1980;109(2):160-174.
5 Posner MI.Orienting of attention.Q J Exp Psychol 1980;32(1):3-25.
6 Eriksen CW,Yeh Y.Allocation of attention in the visual field.J Exp Psychol Hum Percept Perform 1985;11(5):583-597.
7 Müller MM,Malinowski P,Gruber T,Hillyard SA.Sustained division of the attentional spotlight.Nature 2003;424(6946):309-312.
8 McMains SA,Somers DC.Multiple spotlights of attentional selection in human visual cortex.Neuron 2004;42(4):677-686.
9 Jans B,Peters JC,De Weerd P.Visual spatial attention to multiple locations at once:The jury is still out.Psychol Rev2010;117(2):637-682.
10 Desimone R,Duncan J.Neural mechanisms of selective visual attention.Annu Rev Neurosci 1995;18:193-222.
11 Reynolds JH,Heeger DJ.The normalization model of attention.Neuron 2009;61:168-185.
12 Large EW,Jones MR.The dynamics of attending:How people track time-varying events.Psychol Rev 1999;106(1):119-159.
13 VanRullen R.Perceptual cycles.Trends Cogn Sci 2016;20(10):723-735.
14 Schroeder CE,Lakatos P.Low-frequency neuronal oscillations as instruments of sensory selection.Trends Neurosci2009;32(1):9-18.
15 Jones MR,Boltz M.Dynamic attending and responses to time.Psychol Rev 1989;96(3):459-491.
16 Hickok G,Farahbod H,Saberi K.The rhythm of perception:entrainment to acoustic rhythms induces subsequent perceptual oscillation.Psychol Sci 2015;26(7):1006-1013.
17 Patel AD,Iversen JR.The evolutionary neuroscience of musical beat perception:the Action Simulation for Auditory Prediction(ASAP)hypothesis.Front Syst Neurosci 2014;8:57.
18 Spaak E,de Lange FP,Jensen O.Local entrainment of alpha oscillations by visual stimuli causes cyclic modulation of perception.J Neurosci 2014;34(10):3536-3544.
19 Fries P,Reynolds JH,Rorie AE,Desimone R.Modulation of oscillatory neuronal synchronization by selective visual attention.Science 2001;291(5508):1560-1563.
20 Womelsdorf T,Fries P,Mitra PP,Desimone R.Gamma-band synchronization in visual cortex predicts speed of change detection.Nature 2006;439(7077):733-736.
21 Gregoriou GG,Gotts SJ,Zhou H,Desimone R.High-frequency,long-range coupling between prefrontal and visual cortex during attention.Science 2009;324(5931):1207-1210.
22 Lakatos P,Karmos G,Mehta AD,Ulbert I,Schroeder CE.Entrainment of neuronal oscillations as a mechanism of attentional selection.Science 2008;320(5872):110-113.
23 Lakatos P,Barczak A,Neymotin SA,McGinnis T,Ross D,Javitt DC,O’Connell MN.Global dynamics of selective attention and its lapses in primary auditory cortex.Nat Neurosci 2016;19(12):1707-1717.
24 VanRullen R,Koch C.Is perception discrete or continuous?Trends Cogn Sci 2003;7(5):207-213.
25 Busch NA,Dubois J,VanRullen R.The phase of ongoing EEG oscillations predicts visual perception.J Neurosci2009;29(24):7869-7876.
26 Busch NA,VanRullen R.Spontaneous EEG oscillations reveal periodic sampling of visual attention.Proc Natl Acad Sci U S A 2010;107(37):16048-16053.
27 Myers NE,Stokes MG,Walther L,Nobre AC.Oscillatory brain state predicts variability in working memory.J Neurosci2014;34(23):7735-7743.
28 DuguéL,Marque P,VanRullen R.The phase of ongoing oscillations mediates the causal relation between brain excitation and visual perception.J Neurosci 2011;31(33):11889-11893.
29 Drewes J,VanRullen R.This is the rhythm of your eyes:The phase of ongoing electroencephalogram oscillations modulates saccadic reaction time.J Neurosci 2011;31(12):4698-4708.
30 Mathewson KE,Gratton G,Fabiani M,Beck DM,Ro T.To see or not to see:Prestimulus phase predicts visual awareness.J Neurosci 2009;29(9):2725-2732.
31 Samaha J,Postle BR.The speed of alpha-band oscillations predicts the temporal resolution of visual perception.Curr Biol 2015;25(22):2985-2990.
32 Wutz A,Melcher D,Samaha J.Frequency modulation of neural oscillations according to visual task demands.Proc Natl Acad Sci U S A 2018;115(6):1346-1351.
33 Buschman TJ,Kastner S.From behavior to neural dynamics:An integrated theory of attention.Neuron 2015;88(1):127-144.
34 Landau AN,Fries P.Attention samples stimuli rhythmically.Curr Biol 2012;22(11):1000-1004.
35 Fiebelkorn IC,Saalmann YB,Kastner S.Rhythmic sampling within and between objects despite sustained attention at a cued location.Curr Biol 2013;23(24):2553-2558.
36 Lamy D,Egeth H.Object-based selection:the role of attentional shifts.Percept Psychophys 2002;64:52-66.
37 Song K,Meng M,Chen L,Zhou K,Luo H.Behavioral oscillations in attention:Rhythmicαpulses mediated throughθband.J Neurosci 2014;34(14):4837-4844.
38 Klimesch W,Sauseng P,Hanslmayr S.EEG alpha oscillations:The inhibition-timing hypothesis.Brain Res Rev 2007;53(1):63-88.
39 Huang Y,Chen L,Luo H.Behavioral oscillation in priming:Competing perceptual predictions conveyed in alternating theta-band rhythms.J Neurosci 2015;35(6):2830-2837.
40 Davidson MJ,Alais D,van Boxtel JJA,Tsuchiya N.Attention periodically samples competing stimuli during binocular rivalry.Elife 2017.pii:e40868.
41 Tomassini A,Spinelli D,Jacono M,Sandini G,Morrone MC.Rhythmic oscillations of visual contrast sensitivity synchronized with action.J Neurosci 2015;35(18):7019-7029.
42 Ho HT,Leung J,Burr DC,Alais D,Morrone MC.Auditory sensitivity and decision criteria oscillate at different frequencies separately for the two ears.Curr Biol 2017;27(23):3643-3649.e3.
43 Wang Y,Luo H.Behavioral oscillation in face priming:Prediction about face identity is updated at a theta-band rhythm.Prog Brain Res 2017;236:211-224.
44 Irwin D,Gordon R.Eye movements,attention and trans-saccadic memory.Vis Cogn 1998;5(1-2):127-155.
45 VanRullen R.Visual attention:A rhythmic process?Curr Biol 2013;23(24):R1110-R1112.
46 VanRullen R,Carlson T,Cavanagh P.The blinking spotlight of attention.Proc Natl Acad Sci U S A 2007;104(49):19204-19209.
47 Landau AN,Schreyer HM,van Pelt S,Fries P.Distributed attention is implemented through theta-rhythmic gamma modulation.Curr Biol 2015;25(17):2332-2337.
48 DuguéL,Roberts M,Carrasco M.Attention reorients periodically.Curr Biol 2016;26(12):1595-1601.
49 Jia J,Liu L,Fang F,Luo H.Sequential sampling of visual objects during sustained attention.PLoS Biol 2017;15(6):e2001903.
50 Buschman TJ,Miller EK.Serial,covert shifts of attention during visual search are reflected by the frontal eye fields and correlated with population oscillations.Neuron 2009;63(3):386-396.
51 Treisman AM,Gelade G.A feature-integration theory of attention.Cognit Psychol 1980;12:97-136.
52 DuguéL,Marque P,VanRullen R.Theta oscillations modulate attentional search performance periodically.J Cogn Neurosci 2015;27(5):945-958.
53 DuguéL,McLelland D,Lajous M,VanRullen R.Attention searches nonuniformly in space and in time.Proc Natl Acad Sci U S A 2015;112(49):15214-15219.
54 Koch C,Ullman S.Shifts in selective visual attention:towards the underlying neural circuitry.Hum Neurobiol1985;4(4):219-227.
55 Foulsham T,Underwood G.What can saliency models predict about eye movements?Spatial and sequential aspects of fixations during encoding and recognition.J Vis 2008;8(2):6.1-17.
56 Wang W,Chen C,Wang Y,Jiang T,Fang F,Yao Y.Simulating human saccadic scanpaths on natural images.IEEE Conference on Computer Vision and Pattern Recognition(CVPR);Colorado,USA,2011.
57 Klein RM.Inhibition of return.Trends Cogn Sci 2000;4(4):138-147.
58 Rollenhagen JE,Olson CR.Low-frequency oscillations arising from competitive interactions between visual stimuli in macaque inferotemporal cortex.J Neurophysiol 2005;94(5):3368-3387.
59 Kienitz R,Schmiedt JT,Shapcott KA,Kouroupaki K,Saunders RC,Schmid MC.Theta rhythmic neuronal activity and reaction times arising from cortical receptive field interactions during distributed attention.Curr Biol 2018;28(15):2377-2387.