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So far, the focus has been on how ANNs can help to explain the tuning of neurons at various stages of the visual cortical hierarchy. However, the role of modulatory feedback connections, which play a role in attention and perceptual organization, has not yet been resolved. The present study presents a biologically plausible neural network that performs scene segmentation and can shift attention using modulatory feedback connections from higher to lower cortical brain areas. The model replicates several neurophysiological signatures of recurrent processing. Specifically, figural regions elicit more activity in model units than background regions. The modulation of activity by figure and ground occurs at a delay after the first feedforward response, because it depends on a loop through the higher model areas. Importantly, the figural response enhancement is amplified by object-based attention, which stays focused on the figural regions and does not spill over to the adjacent background, just as is observed in the visual cortex. Our results indicate how progress in artificial intelligence can be used to garner insight into the recurrent cortical processing for scene segmentation and object-based attention.<\/jats:p>","DOI":"10.1371\/journal.pone.0337087","type":"journal-article","created":{"date-parts":[[2025,12,10]],"date-time":"2025-12-10T18:35:54Z","timestamp":1765391754000},"page":"e0337087","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":1,"title":["Modulatory feedback determines attentional object segmentation in a model of the ventral stream"],"prefix":"10.1371","volume":"20","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6249-841X","authenticated-orcid":true,"given":"Paolo","family":"Papale","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jonathan R.","family":"Williford","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stijn","family":"Balk","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pieter R.","family":"Roelfsema","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2025,12,10]]},"reference":[{"issue":"6753","key":"pone.0337087.ref001","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1038\/44134","article-title":"fMRI evidence for objects as the units of attentional selection","volume":"401","author":"KM O\u2019Craven","year":"1999","journal-title":"Nature"},{"key":"pone.0337087.ref002","article-title":"Selective attention and the organization of visual information","author":"J Duncan","year":"1984","journal-title":"J Exp Psychol Gen"},{"issue":"1","key":"pone.0337087.ref003","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1037\/0096-3445.126.1.3","article-title":"Object-based attentional selection--grouped arrays or spatially invariant representations?: comment on vecera and Farah (1994)","volume":"126","author":"AF Kramer","year":"1997","journal-title":"J Exp Psychol Gen"},{"issue":"4","key":"pone.0337087.ref004","doi-asserted-by":"crossref","first-page":"1011","DOI":"10.1037\/0096-1523.24.4.1011","article-title":"Object-based attention and occlusion: evidence from normal participants and a computational model","volume":"24","author":"M Behrmann","year":"1998","journal-title":"J Exp Psychol Hum Percept Perform"},{"issue":"7","key":"pone.0337087.ref005","doi-asserted-by":"crossref","first-page":"1003","DOI":"10.1016\/j.neuron.2023.03.016","article-title":"Solving the binding problem: Assemblies form when neurons enhance their firing rate-they don\u2019t need to oscillate or synchronize","volume":"111","author":"PR Roelfsema","year":"2023","journal-title":"Neuron"},{"issue":"8","key":"pone.0337087.ref006","doi-asserted-by":"crossref","first-page":"2542","DOI":"10.3758\/s13414-011-0200-0","article-title":"Incremental grouping of image elements in vision","volume":"73","author":"PR Roelfsema","year":"2011","journal-title":"Atten Percept Psychophys"},{"key":"pone.0337087.ref007","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1146\/annurev.neuro.29.051605.112939","article-title":"Cortical algorithms for perceptual grouping","volume":"29","author":"PR Roelfsema","year":"2006","journal-title":"Annu Rev Neurosci"},{"issue":"5043","key":"pone.0337087.ref008","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1126\/science.1734518","article-title":"Information processing in the primate visual system: an integrated systems perspective","volume":"255","author":"DC Van Essen","year":"1992","journal-title":"Science"},{"issue":"6","key":"pone.0337087.ref009","doi-asserted-by":"crossref","first-page":"974","DOI":"10.1038\/s41593-019-0392-5","article-title":"Evidence that recurrent circuits are critical to the ventral stream\u2019s execution of core object recognition behavior","volume":"22","author":"K Kar","year":"2019","journal-title":"Nat Neurosci"},{"key":"pone.0337087.ref010","doi-asserted-by":"crossref","unstructured":"Thorat S, van Gerven M, Peelen M. 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