Controlled shifts of attention between competing stimuli are crucial for effective everyday visual behaviour. While these typically involve overt shifts of fixation, many past studies used covert attention shifts in which fixation is unchanged, meaning that some response components may result from the inhibition of eye movements. In this study, the neural events in the human brain when making overt shifts of attention are studied through the combination of event-related potential recording with simultaneous eye tracking. Fixation shifts under competition (central target remains visible when a peripheral target appears) were compared with noncompetition (central target disappears). A longer latency for competition compared with noncompetition, which is found in the saccadic response, is already present in the early occipital positivity when a single target is presented for the fixation shift. These results indicate that the requirement to disengage from a current target affects the time course of neural processing at an early level. However, the relation is more complex when the participant is required to choose which of two targets to fixate.
AmanoK.GodaN.NishidaS. Y.EjimaY.TakedaT.OhtaniY. (2006).
Estimation of the timing of human visual perception from magnetoencephalography. Journal of Neuroscience,
26, 3981–3991.
2.
Anllo-VentoL.HillyardS. A. (1996).
Selective attention to the color and direction of moving stimuli: Electrophysiological correlates of hierarchical feature selection. Perception & Psychophysics,
58, 191–206.
3.
AtkinsonJ. (1984).
Human visual development over the first 6 months of life. A review and a hypothesis. Human Neurobiology,
3, 61–74.
4.
AtkinsonJ. (2000). Newborn vision. In AtkinsonJ. (Ed.), The developing visual brain.
Oxford University Press.
5.
AtkinsonJ.BraddickO. (2012).
Visual attention in the first years: Typical development and developmental disorders. Developmental Medicine & Child Neurology,
54, 589–595.
6.
AtkinsonJ.HoodB.BraddickO.Wattam-BellJ. (1988).
Infants' control of fixation shifts with single and competing targets-mechanisms of shifting attention.Perception,
17, 367–368.
7.
AtkinsonJ.HoodB.Wattam-BellJ.BraddickO. (1992).
Changes in infants’ ability to switch visual attention in the first three months of life. Perception,
21, 643–643.
8.
BartolomeoP.De SchottenM. T.DoricchiF. (2007).
Left unilateral neglect as a disconnection syndrome. Cerebral Cortex,
17, 2479–2490.
9.
ButcherP. R.KalverboerA. F.GeuzeR. H. (2000).
Infants’ shifts of gaze from a central to a peripheral stimulus: A longitudinal study of development between 6 and 26 weeks. Infant Behavior and Development,
23, 3–21.
10.
CollinsC. E.LyonD. C.KaasJ. H. (2005).
Distribution across cortical areas of neurons projecting to the superior colliculus in new world monkeys. The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology,
285, 619–627.
11.
CorbettaM.AkbudakE.ConturoT. E.SnyderA. Z.OllingerJ. M.DruryH. A.LinenweberM. R.PetersenS. E.RaihleM. E.ShulmanG. L.Van EssenD. C. (1998).
A common network of functional areas for attention and eye movements. Neuron,
21, 761–773.
12.
CordonesI.GómezC. M.EscuderoM. (2013).
Cortical dynamics during the preparation of antisaccadic and prosaccadic eye movements in humans in a gap paradigm. PLoS One,
8, e63751.
13.
CsibraG.JohnsonM. H.TuckerL. A. (1997).
Attention and oculomotor control: A high-density ERP study of the gap effect. Neuropsychologia,
35, 855–865.
14.
De SchottenM. T.UrbanskiM.DuffauH.VolleE.LévyR.DuboisB.BartolomeoP. (2005).
Direct evidence for a parietal-frontal pathway subserving spatial awareness in humans. Science,
309, 2226–2228.
15.
DelormeA.WesterfieldM.MakeigS. (2007).
Medial prefrontal theta bursts precede rapid motor responses during visual selective attention. Journal of Neuroscience,
27, 11949–11959.
16.
Di RussoF.MartínezA.HillyardS. A. (2003).
Source analysis of event-related cortical activity during visuo-spatial attention. Cerebral Cortex,
13, 486–499.
17.
DorrisM. C.MunozD. P. (1995).
A neural correlate for the gap effect on saccadic reaction times in monkey. Journal of Neurophysiology,
73, 2558–2562.
18.
EimerM.ForsterB.VelzenJ. V.PrabhuG. (2005).
Covert manual response preparation triggers attentional shifts: ERP evidence for the premotor theory of attention. Neuropsychologia,
43, 957–966.
19.
EimerM.Van VelzenJ.DriverJ. (2002).
Cross-modal interactions between audition, touch, and vision in endogenous spatial attention: ERP evidence on preparatory states and sensory modulations. Journal of Cognitive Neuroscience,
14, 254–271.
20.
ElisonJ. T.PatersonS. J.WolffJ. J.ReznickJ. S.SassonN. J.GuH.BotteronK. N.DagerS. R.EstesA. M.GerigC.HazlettH. C.SchultzR. T.StynerM.ZwaigenbaumL.PivenJ.EvansA. C., & IBIS Network. (2013).
White matter microstructure and atypical visual orienting in 7-month-olds at risk for autism. The American Journal of Psychiatry,
170, 899–908.
21.
FarroniT.SimionF.UmiltàC.BarbaB. D. (1999).
The gap effect in newborns. Developmental Science,
2, 174–186.
22.
FerreeT. C.LuuP.RussellG. S.TuckerD. M. (2001).
Scalp electrode impedance, infection risk, and EEG data quality. Clinical Neurophysiology,
112, 536–544.
23.
FischerB. (1986).
The role of attention in the preparation of visually guided eye movements in monkey and man. Psychological Research,
48, 251–257.
24.
GoldbergM. E.WurtzR. H. (1972).
Activity of superior colliculus in behaving monkey. I. Visual receptive fields of single neurons. Journal of Neurophysiology,
35, 542–559.
25.
GomezC.AtienzaM.GomezG.VazquezM. (1996).
Response latencies and event-related potentials during the gap paradigm using saccadic responses in human subjects. International Journal of Psychophysiology,
23, 91–99.
26.
HampelF. R. (1974).
The influence curve and its role in robust estimation. Journal of the American Statistical Association,
69, 383–393.
27.
HolmS. (1979).
A simple sequentially rejective multiple test procedure. Scandinavian Journal of Statistics,
6, 65–70.
28.
HoodB.AtkinsonJ. (1993).
Disengaging visual attention in the infant and adult. Infant Behavior and Development,
16, 405–422.
29.
Huber-HuberC.DityeT.FernándezM. M.AnsorgeU. (2016).
Using temporally aligned event-related potentials for the investigation of attention shifts prior to and during saccades. Neuropsychologia,
92, 129–141.
30.
JohnsonM. H. (1990).
Cortical maturation and the development of visual attention in early infancy. Journal of Cognitive Neuroscience,
2, 81–95.
31.
JohnsonM. H. (2002). The development of visual attention: A cognitive neuroscience perspective. In JohnsonM. H.MunakataY.GilmoreR. O. (Eds.), Brain development and cognition: A reader (pp. 134–150).
John Wiley & Sons.
32.
KalesnykasR.HallettP. (1987).
The differentiation of visually guided and anticipatory saccades in gap and overlap paradigms. Experimental Brain Research,
68, 115–121.
33.
KatsumiO.HiroseT.TsukadaT. (1988).
Effect of number of elements and size of stimulus field on recordability of pattern reversal visual evoked response. Investigative Ophthalmology & Visual Science,
29, 922–927.
34.
KawakuboY.KasaiK.OkazakiS.Hosokawa-KakuraiM.WatanabeK.-I.KuwabaraH.IshijimaM.YamasueH.IwanamiA.MaekawaH.KatoN. (2007).
Electrophysiological abnormalities of spatial attention in adults with autism during the gap overlap task. Clinical Neurophysiology,
118, 1464–1471.
35.
KulkeL. (2015). Cortical mechanisms of visual attention in typically developing infants and adults (PhD thesis).
University College London.
36.
KulkeL. (2017).
The effect of stimulus size and eccentricity on attention shift latencies. Vision,
1, E25.
37.
KulkeL. (2019).
Neural mechanisms of overt attention shifts to emotional faces. Neuroscience,
418, 59–68.
38.
KulkeL.AtkinsonJ.BraddickO. (2015).
Automatic detection of attention shifts in infancy: Eye tracking in the fixation shift paradigm. PLoS One,
10, e0142505.
39.
KulkeL.AtkinsonJ.BraddickO. (2016a).
Neural differences between covert and overt attention studied using EEG with simultaneous remote eye tracking. Frontiers in Human Neuroscience,
10, 592.
40.
KulkeL.AtkinsonJ.BraddickO. (2016b).
Neural mechanisms of attention become more specialised during infancy: Insights from combined eye tracking and EEG. Developmental Psychobiology,
59, 250–260.
41.
LuckS. J. (2005). An introduction to the event-related potential technique.
MIT Press.
42.
MartinezA.Anllo-VentoL.SerenoM. I.FrankL. R.BuxtonR. B.DubowitzD. J.WongE. C.HinrichsH.HeinzeH. J.HillyardS. A. (1999).
Involvement of striate and extrastriate visual cortical areas in spatial attention. Nature Neuroscience,
2, 364–369.
43.
MatsuzawaM.ShimojoS. (1997).
Infants’ fast saccades in the gap paradigm and development of visual attention. Infant Behavior and Development,
20, 449–455.
44.
McAdamsC. J.ReidR. C. (2005).
Attention modulates the responses of simple cells in monkey primary visual cortex. Journal of Neuroscience,
25, 11023–11033.
45.
MercuriE.AtkinsonJ.BraddickO.AnkerS.CowanF.RutherfordM.PennockJ.DubowitzL. (1997a).
Basal ganglia damage and impaired visual function in the newborn infant. Archives of Disease in Childhood-Fetal and Neonatal Edition,
77, F111–F114.
46.
MercuriE.AtkinsonJ.BraddickO.AnkerS.CowanF.RutherfordM.PennockJ.DubowitzL. (1997b).
Visual function in full-term infants with hypoxic-ischaemic encephalopathy. Neuropediatrics,
28, 155–161.
47.
NeggersS.RaemaekersM.LampmannE.PostmaA.RamseyN. (2005).
Cortical and subcortical contributions to saccade latency in the human brain. European Journal of Neuroscience,
21, 2853–2863.
48.
PerryR.ZekiS. (2000).
The neurology of saccades and covert shifts in spatial attention: An event-related fMRI study. Brain,
123, 2273–2288.
49.
PraamstraP.OostenveldR. (2003).
Attention and movement-related motor cortex activation: A high-density EEG study of spatial stimulus–response compatibility. Cognitive Brain Research,
16, 309–322.
50.
ReganD. (1989). Human brain electrophysiology: Evoked potentials and evoked magnetic fields in science and medicine.
Elsevier.
51.
ReulenJ. P. (1984).
Latency of visually evoked saccadic eye movements. Biological Cybernetics,
50, 251–262.
52.
RicciD.CesariniL.GalliniF.SerraoF.LeoneD.BaranelloG.VascoG.AlfieriP.StaccioliS.RomeoD. M.TinelliF.MolleF.LeporeD.BaldascinoA.RamenghiL. A.TorrioliM. G.RomagnoliC.CowanF.AtkinsonJ., . . . De RoseP. (2010).
Cortical visual function in preterm infants in the first year. The Journal of Pediatrics,
156, 550–555.
53.
RolfsM.VituF. (2007).
On the limited role of target onset in the gap task: Support for the motor-preparation hypothesis. Journal of Vision,
7, 7.
54.
RuggM.MilnerA.LinesC.PhalpR. (1987).
Modulation of visual event-related potentials by spatial and non-spatial visual selective attention. Neuropsychologia,
25, 85–96.
55.
SaslowM. (1967).
Effects of components of displacement-step stimuli upon latency for saccadic eye movement. Journal of the Optical Society of America,
57, 1024–1029.
56.
SchillerP. H. (1985). A model for the generation of visually guided saccadic eye movements. In RoseD.DobsonV. G. (Eds.), Models of the visual cortex.
John Wiley & Sons Ltd.
57.
SchillerP. H.StrykerM. (1972).
Single-unit recording and stimulation in superior colliculus of the alert rhesus monkey. Journal of Neurophysiology,
35, 915–924.
58.
SchillerP. H.TehovnikE. J. (2005).
Neural mechanisms underlying target selection with saccadic eye movements. Progress in Brain Research,
149, 157–171.
59.
SheligaB. M.RiggioL.RizzolattiG. (1994).
Orienting of attention and eye movements. Experimental Brain Research,
98, 507–522.
60.
ShomsteinS.KravitzD. J.BehrmannM. (2012).
Attentional control: Temporal relationships within the fronto-parietal network. Neuropsychologia,
50, 1202–1210.
61.
TuckerD. M. (1993).
Spatial sampling of head electrical fields: The geodesic sensor net. Electroencephalography and Clinical Neurophysiology,
87, 154–163.
62.
VassilevA.ManahilovV.MitovD. (1983).
Spatial frequency and the pattern onset-offset response. Vision Research,
23, 1417–1422.
63.
WangF.ChenM.YanY.ZhaopingL.LiW. (2015).
Modulation of neuronal responses by exogenous attention in macaque primary visual cortex. Journal of Neuroscience,
35, 13419–13429.
64.
WeaverM. D.van ZoestW.HickeyC. (2017).
A temporal dependency account of attentional inhibition in oculomotor control. NeuroImage,
147, 880–894.
65.
YamaguchiS.TsuchiyaH.KobayashiS. (1994).
Electroencephalographic activity associated with shifts of visuospatial attention. Brain,
117, 553–562.
66.
YamaguchiS.TsuchiyaH.KobayashiS. (1995).
Electrophysiologic correlates of visuo-spatial attention shift. Electroencephalography and Clinical Neurophysiology,
94, 450–461.
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