Our study applied computational methods to distinguish the representations of objects’ perceived real-world size, retinal size, and inferred real-world depth features in both human brains and ANNs. Consistent with prior studies reporting real-world size representations (Huang et al., 2022; Khaligh-Razavi et al., 2018; Konkle and Caramazza, 2013; Konkle and Oliva, 2012b; Luo et al., 2023; Quek et al., 2023; Wang et al., 2022a), we found that both human brains and ANNs contain significant information about real-world size. Critically, our study goes beyond the contributions of prior studies in several key ways, offering both theoretical and methodological advances: (a) we eliminated the confounding impact of perceived real-world depth (in addition to retinal size) on the real-world size representation; (b) we conducted analyses based on more ecologically valid naturalistic images; (c) we obtained precise feature values for each object in every image, instead of simply dividing objects into two or seven coarse categories; and (d) we utilized a multi-modal, partial correlation RSA that combines EEG, hypothesis-based models, and ANNs. Our approach allowed us to investigate representational time courses and reverse engineering manipulations in unparalleled detail. By integrating EEG data with hypothesis-based models and ANNs, this method offers a powerful tool for dissecting the neural underpinnings of object size and depth perception in more ecological contexts, which enriches our comprehension of the brain’s representational mechanisms.
Using EEG, we uncovered a representational timeline for visual object processing, with object real-world depth information represented first, followed by retinal size, and finally real-world size. While size and depth are highly correlated to each other, our results suggest that the human brain indeed has dissociated time courses and mechanisms to process them. The later representation time window for object real-world size may suggest that the brain requires more sophisticated, higher-level information to form this representation, perhaps incorporating semantic and/or memory information about familiar objects, which was corroborated by our ANN and Word2Vec analyses. These findings also align with a recent fMRI study (Luo et al., 2023) using natural images to explore the neural selectivity for real-world size, finding that low-level visual information could hardly account for neural size preferences, although that study did not consider covariables like retinal size and real-world depth.
In contrast to the later-emerging real-world size representations, it makes sense that retinal size representations could be processed more quickly based on more fundamental, lower-level information such as shape and edge discrimination. Interestingly, real-world depth representations emerged even earlier than retinal size, suggesting that depth feature may precede real-world size processing. This early emergence might reflect the role of depth cues in facilitating object segmentation, as proposed in classical theories of vision (Marr, 1982), where depth helps delineate object boundaries from complex backgrounds. Additionally, there was a secondary, albeit substantially later, significant depth representation time window, which might indicate that our brains also have the ability to integrate object retinal size and higher-level real-size information to form the final representation of real-world depth. Our comparisons between human brains and artificial models and explorations on ANNs and Word2Vec offer further insights and suggest that although real-world object size and depth are closely related, object real-world size appears to be a more stable and higher-level dimension.
The concept of ‘object space’ in cognitive neuroscience research is crucial for understanding how various visual features of objects are represented. Historically, various visual features have been considered important dimensions in constructing object space, including animate-inanimate (Kriegeskorte et al., 2008; Naselaris et al., 2012), spikiness (Bao et al., 2020; Coggan and Tong, 2023), and physical appearance (Edelman et al., 1998). In this study, we focus on one particular dimension, real-world size (Huang et al., 2022; Konkle and Caramazza, 2013; Konkle and Oliva, 2012b). How we generate neural distinctions of different object real-world size and where this ability comes from remain uncertain. Some previous studies found that object shape rather than texture information could trigger neural size representations (Huang et al., 2022; Long et al., 2016; Long et al., 2018; Wang et al., 2022a). Our results attempt to further advance their findings that object real-world size is a stable and higher-level dimension substantially driven by object semantics in object space.
Increasingly, research has begun to use ANNs to study the mechanisms of object recognition (Ayzenberg et al., 2023; Cichy and Kaiser, 2019; Doerig et al., 2023; Kanwisher et al., 2023). We can explore how the human brain processes information at different levels by comparing brain activity with models (Cichy et al., 2016; Khaligh-Razavi and Kriegeskorte, 2014; Kuzovkin et al., 2018; Xie et al., 2020), and we can also analyze the representation patterns of the models with some specific manipulations and infer potential processing mechanisms in the brain (Golan et al., 2020; Huang et al., 2022; Lu and Ku, 2023; Xu et al., 2021). In the current study, our comparisons between EEG signals and different ANNs showed that the visual model’s early layer had a higher similarity to the brain in the early stage, while the visual-semantic model’s late layer had a higher similarity to the brain in the late stage. In addition, to assess whether this EEG-ANN similarity pattern generalizes to more biologically inspired architectures, we conducted the same analyses and found that CORnet (Kubilius et al., 2019) showed similar patterns to those observed for ResNet and CLIP, providing converging evidence across models (Figure 3—figure supplement 5). However, for the representation of objects, partial correlation results for different ANNs didn’t demonstrate the superiority of the multi-modal model at late layers. This might be due to models like CLIP, which contain semantic information, learning more complex image descriptive information (like the relationship between object and the background in the image). Real-world size might be a semantic dimension of the object itself, and its representation does not require overall semantic descriptive information of the image. In contrast, retinal size and real-world depth could rely on image background information for estimation, thus their representations in the CLIP late layer disappeared when input images had only pure object but no background.
Although our study does not directly test specific models of visual object processing, the observed temporal dynamics provide important constraints for theoretical interpretations. In particular, we find that real-world size representations emerge significantly later than low-level visual features such as retinal size and depth. This temporal profile is difficult to reconcile with a purely feedforward account of visual processing (e.g. DiCarlo et al., 2012), which posits that object properties are rapidly computed in a sequential hierarchy of increasingly complex visual features. Instead, our results are more consistent with frameworks that emphasize recurrent or top-down processing, such as the reverse hierarchy theory (Hochstein and Ahissar, 2002), which suggests that high-level conceptual information may emerge later and involve feedback to earlier visual areas. This interpretation is further supported by representational similarities with late-stage artificial neural network layers and with a semantic word embedding model (Word2Vec), both of which reflect learned, abstract knowledge rather than low-level visual features. Taken together, these findings suggest that real-world size is not merely a perceptual attribute, but one that draws on conceptual or semantic-level representations acquired through experience. While our EEG analyses focused on posterior electrodes and thus cannot definitively localize cortical sources, we see this study as a step toward linking low-level visual input with higher-level semantic knowledge. Future work incorporating broader spatial coverage (e.g. anterior sensors), source localization, or complementary modalities such as MEG and fMRI will be critical to adjudicate between alternative models of object representation and to more precisely trace the origin and flow of real-world size information in the brain. Moreover, building on the promising findings of our study, future work may further delve into the detailed processes of object processing and object space. One important problem to solve is how real-world size interacts with other object dimensions in object space. In addition, our approach could be used with future studies investigating other influences on object processing, such as how different task conditions impact and modulate the processing of various visual features.
Moreover, we must also emphasize that in this study, we were concerned with perceived real-world size and depth reflecting a perceptual estimation of our world, which are slightly different from absolute physical size and depth. The differences in brain encoding between perceived and absolute physical size and depth require more comprehensive measurements of an object’s physical attributes for further exploration. Also, we focused on perceiving depth and size from 2D images in this study, which might have some differences in brain mechanism compared to physically exploring the 3D world. Nevertheless, we believe our study offers a valuable contribution to object recognition, especially the encoding process of object real-world size in natural images. Additionally, we acknowledge that our metric for real-world depth was derived indirectly as the ratio of perceived real-world size to retinal size. While this formulation is grounded in geometric principles of perspective projection and served the purpose of analytically dissociating depth from size in our RSA framework, it remains a proxy rather than a direct measure of perceived egocentric distance. Future work incorporating behavioral or psychophysical depth ratings would be valuable for validating and refining this metric.
In conclusion, we used computational methods to distinguish the representations of real-world size, retinal size, and real-world depth features of objects in ecologically natural images in both human brains and ANNs. We found an unconfounded representation of object real-world size, which emerged at later time windows in the human EEG signal and at later layers of artificial neural networks compared to real-world depth, and which also appeared to be preserved as a stable dimension in object space. Thus, although size and depth properties are closely correlated, the processing of perceived object size and depth may arise through dissociated time courses and mechanisms. Our research provides a temporally resolved characterization of how certain key object properties – such as object real-world size, depth, and retinal size – are represented in the brain, which advances our understanding of real-world size encoding as a stable and semantically grounded dimension in object space, and contributes to the development of more brain-like visual models.