Nikolaus Kriegeskorte

Tal Golan, Matthew Siegelman, Nikolaus Kriegeskorte et al.

Neural network language models can serve as computational hypotheses about how humans process language. We compared the model-human consistency of diverse language models using a novel experimental approach: controversial sentence pairs. For each controversial sentence pair, two language models disagree about which sentence is more likely to occur in natural text. Considering nine language models (including n-gram, recurrent neural networks, and transformer models), we created hundreds of such controversial sentence pairs by either selecting sentences from a corpus or synthetically optimizing sentence pairs to be highly controversial. Human subjects then provided judgments indicating for each pair which of the two sentences is more likely. Controversial sentence pairs proved highly effective at revealing model failures and identifying models that aligned most closely with human judgments. The most human-consistent model tested was GPT-2, although experiments also revealed significant shortcomings of its alignment with human perception.

Baihan Lin, Nikolaus Kriegeskorte

A central question for neuroscience is how to characterize brain representations of perceptual and cognitive content. An ideal characterization should distinguish different functional regions with robustness to noise and idiosyncrasies of individual brains that do not correspond to computational differences. Previous studies have characterized brain representations by their representational geometry, which is defined by the representational dissimilarity matrix (RDM), a summary statistic that abstracts from the roles of individual neurons (or responses channels) and characterizes the discriminability of stimuli. Here we explore a further step of abstraction: from the geometry to the topology of brain representations. We propose topological representational similarity analysis (tRSA), an extension of representational similarity analysis (RSA) that uses a family of geo-topological summary statistics that generalizes the RDM to characterize the topology while de-emphasizing the geometry. We evaluate this new family of statistics in terms of the sensitivity and specificity for model selection using both simulations and fMRI data. In the simulations, the ground truth is a data-generating layer representation in a neural network model and the models are the same and other layers in different model instances (trained from different random seeds). In fMRI, the ground truth is a visual area and the models are the same and other areas measured in different subjects. Results show that topology-sensitive characterizations of population codes are robust to noise and interindividual variability and maintain excellent sensitivity to the unique representational signatures of different neural network layers and brain regions. These methods enable researchers to calibrate comparisons among representations in brains and models to be sensitive to the geometry, the topology, or a combination of both.

Hossein Adeli, Seoyoung Ahn, Nikolaus Kriegeskorte et al.

The spreading of attention has been proposed as a mechanism for how humans group features to segment objects. However, such a mechanism has not yet been implemented and tested in naturalistic images. Here, we leverage the feature maps from self-supervised vision Transformers and propose a model of human object-based attention spreading and segmentation. Attention spreads within an object through the feature affinity signal between different patches of the image. We also collected behavioral data on people grouping objects in natural images by judging whether two dots are on the same object or on two different objects. We found that our models of affinity spread that were built on feature maps from the self-supervised Transformers showed significant improvement over baseline and CNN based models on predicting reaction time patterns of humans, despite not being trained on the task or with any other object labels. Our work provides new benchmarks for evaluating models of visual representation learning including Transformers.

Tal Golan, Wenxuan Guo, Heiko H. Schütt et al.

Comparing representations of complex stimuli in neural network layers to human brain representations or behavioral judgments can guide model development. However, even qualitatively distinct neural network models often predict similar representational geometries of typical stimulus sets. We propose a Bayesian experimental design approach to synthesizing stimulus sets for adjudicating among representational models efficiently. We apply our method to discriminate among candidate neural network models of behavioral face dissimilarity judgments. Our results indicate that a neural network trained to invert a 3D-face-model graphics renderer is more human-aligned than the same architecture trained on identification, classification, or autoencoding. Our proposed stimulus synthesis objective is generally applicable to designing experiments to be analyzed by representational similarity analysis for model comparison.

Fan L. Cheng, Nikolaus Kriegeskorte

Prompting steers large language models (LLMs) and vision-language models (VLMs) without weight updates, but it remains unclear how instruction changes reshape internal representations to produce behavior. We introduce a nested geometric decomposition framework that treats prompting as a transformation of the representational geometry of the content following the prompt. For each prompt pair, we align representations of the same stimuli under two prompts using increasingly expressive stimulus-invariant maps: translation, rigid transformation with uniform scaling, sequential axis scaling, affine transformation, and nonlinear transformation. We then causally test each map by replacing a single layer's prompt-A hidden state for held-out stimuli with its mapped counterpart and measuring recovery of prompt-B representational geometry and behavior. Across three LLMs, three VLMs, and six text or image datasets spanning style, emotion, scene content, and number, prompts consistently reshape representations toward the instructed task structure. Cross-validated variance decomposition shows that much prompt-induced activation change is captured by shape-preserving maps, especially translation and rigid transformation with uniform scaling, while tier profiles reveal model- and task-specific routing strategies across layers. Crucially, although translation and rigid tiers already improve behavioral agreement, affine transformation is the first tier to nearly recover target-prompt task geometry and yields corresponding behavioral gains. This suggests that cross-dimensional linear mixing is a key mechanism by which prompts reorganize representations toward instructed task structure. Our framework decomposes prompt-induced representational change into interpretable geometric components and reveals how models route task-relevant structure to produce prompt-driven behavior.

Ben Baker, Richard D. Lange, Andrew Richmond et al.

Representations play a central role in the study of both biological and artificial intelligence, as well as philosophy of mind. Across neuroscience, computer science, and philosophy, a recurring theme is that representations not only carry information but should be ``useful'' for or ``usable'' by an agent in some sense. Here, we review how the ``usefulness'' of representations has been conceptualized and how it figures into different conceptions of representation. We identify and explore four aspects of use and usability: representations generally carry \textit{information}; that information may or may not be \textit{useful} and it may or may not be encoded in a usable \textit{format}; and the representations may or may not be \textit{used downstream}. Building on these four aspects of information and use, we then organize existing perspectives on neural representations into three levels: Representations as Information (Level 1); Representations as Usable (Level 2); and Representations as Used (Level 3). Our account is meant to give readers an appreciation for the diversity of notions of ``neural representation,'' help them navigate the vast and multi-disciplinary literature on the topic, and help them clarify the appropriate notion of representation for their own investigations.

Benjamin Peters, James J. DiCarlo, Todd Gureckis et al. · harvard

Vision is widely understood as an inference problem. However, two contrasting conceptions of the inference process have each been influential in research on biological vision as well as the engineering of machine vision. The first emphasizes bottom-up signal flow, describing vision as a largely feedforward, discriminative inference process that filters and transforms the visual information to remove irrelevant variation and represent behaviorally relevant information in a format suitable for downstream functions of cognition and behavioral control. In this conception, vision is driven by the sensory data, and perception is direct because the processing proceeds from the data to the latent variables of interest. The notion of "inference" in this conception is that of the engineering literature on neural networks, where feedforward convolutional neural networks processing images are said to perform inference. The alternative conception is that of vision as an inference process in Helmholtz's sense, where the sensory evidence is evaluated in the context of a generative model of the causal processes giving rise to it. In this conception, vision inverts a generative model through an interrogation of the evidence in a process often thought to involve top-down predictions of sensory data to evaluate the likelihood of alternative hypotheses. The authors include scientists rooted in roughly equal numbers in each of the conceptions and motivated to overcome what might be a false dichotomy between them and engage the other perspective in the realm of theory and experiment. The primate brain employs an unknown algorithm that may combine the advantages of both conceptions. We explain and clarify the terminology, review the key empirical evidence, and propose an empirical research program that transcends the dichotomy and sets the stage for revealing the mysterious hybrid algorithm of primate vision.

Pinyuan Feng, Hossein Adeli, Wenxuan Guo et al.

Recent work has demonstrated that complex visual stimuli can be decoded from human brain activity using deep generative models, helping brain science researchers interpret how the brain represents real-world scenes. However, most current approaches leverage mapping brain signals into intermediate image or text feature spaces before guiding the generative process, masking the effect of contributions from different brain areas on the final reconstruction output. In this work, we propose NeuroAdapter, a visual decoding framework that directly conditions a latent diffusion model on brain representations, bypassing the need for intermediate feature spaces. Our method demonstrates competitive visual reconstruction quality on public fMRI datasets compared to prior work, while providing greater transparency into how brain signals shape the generation process. To this end, we contribute an Image-Brain BI-directional interpretability framework (IBBI) which investigates cross-attention mechanisms across diffusion denoising steps to reveal how different cortical areas influence the unfolding generative trajectory. Our results highlight the potential of end-to-end brain-to-image decoding and establish a path toward interpreting diffusion models through the lens of visual neuroscience.

Benjamin Peters, Nikolaus Kriegeskorte

Human visual perception carves a scene at its physical joints, decomposing the world into objects, which are selectively attended, tracked, and predicted as we engage our surroundings. Object representations emancipate perception from the sensory input, enabling us to keep in mind that which is out of sight and to use perceptual content as a basis for action and symbolic cognition. Human behavioral studies have documented how object representations emerge through grouping, amodal completion, proto-objects, and object files. Deep neural network (DNN) models of visual object recognition, by contrast, remain largely tethered to the sensory input, despite achieving human-level performance at labeling objects. Here, we review related work in both fields and examine how these fields can help each other. The cognitive literature provides a starting point for the development of new experimental tasks that reveal mechanisms of human object perception and serve as benchmarks driving development of deep neural network models that will put the object into object recognition.

Ruben S. van Bergen, Nikolaus Kriegeskorte

Biological visual systems exhibit abundant recurrent connectivity. State-of-the-art neural network models for visual recognition, by contrast, rely heavily or exclusively on feedforward computation. Any finite-time recurrent neural network (RNN) can be unrolled along time to yield an equivalent feedforward neural network (FNN). This important insight suggests that computational neuroscientists may not need to engage recurrent computation, and that computer-vision engineers may be limiting themselves to a special case of FNN if they build recurrent models. Here we argue, to the contrary, that FNNs are a special case of RNNs and that computational neuroscientists and engineers should engage recurrence to understand how brains and machines can (1) achieve greater and more flexible computational depth, (2) compress complex computations into limited hardware, (3) integrate priors and priorities into visual inference through expectation and attention, (4) exploit sequential dependencies in their data for better inference and prediction, and (5) leverage the power of iterative computation.

Tal Golan, Prashant C. Raju, Nikolaus Kriegeskorte

Distinct scientific theories can make similar predictions. To adjudicate between theories, we must design experiments for which the theories make distinct predictions. Here we consider the problem of comparing deep neural networks as models of human visual recognition. To efficiently compare models' ability to predict human responses, we synthesize controversial stimuli: images for which different models produce distinct responses. We applied this approach to two visual recognition tasks, handwritten digits (MNIST) and objects in small natural images (CIFAR-10). For each task, we synthesized controversial stimuli to maximize the disagreement among models which employed different architectures and recognition algorithms. Human subjects viewed hundreds of these stimuli, as well as natural examples, and judged the probability of presence of each digit/object category in each image. We quantified how accurately each model predicted the human judgments. The best performing models were a generative Analysis-by-Synthesis model (based on variational autoencoders) for MNIST and a hybrid discriminative-generative Joint Energy Model for CIFAR-10. These DNNs, which model the distribution of images, performed better than purely discriminative DNNs, which learn only to map images to labels. None of the candidate models fully explained the human responses. Controversial stimuli generalize the concept of adversarial examples, obviating the need to assume a ground-truth model. Unlike natural images, controversial stimuli are not constrained to the stimulus distribution models are trained on, thus providing severe out-of-distribution tests that reveal the models' inductive biases. Controversial stimuli therefore provide powerful probes of discrepancies between models and human perception.

Baihan Lin, Marieke Mur, Tim Kietzmann et al.

Representational similarity analysis (RSA) has been shown to be an effective framework to characterize brain-activity profiles and deep neural network activations as representational geometry by computing the pairwise distances of the response patterns as a representational dissimilarity matrix (RDM). However, how to properly analyze and visualize the representational geometry as dynamics over the time course from stimulus onset to offset is not well understood. In this work, we formulated the pipeline to understand representational dynamics with RDM movies and Procrustes-aligned Multidimensional Scaling (pMDS), and applied it to neural recording of monkey IT cortex. Our results suggest that the the multidimensional scaling alignment can genuinely capture the dynamics of the category-specific representation spaces with multiple visualization possibilities, and that object categorization may be hierarchical, multi-staged, and oscillatory (or recurrent).

Tim C Kietzmann, Courtney J Spoerer, Lynn Sörensen et al.

The human visual system is an intricate network of brain regions that enables us to recognize the world around us. Despite its abundant lateral and feedback connections, object processing is commonly viewed and studied as a feedforward process. Here, we measure and model the rapid representational dynamics across multiple stages of the human ventral stream using time-resolved brain imaging and deep learning. We observe substantial representational transformations during the first 300 ms of processing within and across ventral-stream regions. Categorical divisions emerge in sequence, cascading forward and in reverse across regions, and Granger causality analysis suggests bidirectional information flow between regions. Finally, recurrent deep neural network models clearly outperform parameter-matched feedforward models in terms of their ability to capture the multi-region cortical dynamics. Targeted virtual cooling experiments on the recurrent deep network models further substantiate the importance of their lateral and top-down connections. These results establish that recurrent models are required to understand information processing in the human ventral stream.

Katherine R. Storrs, Nikolaus Kriegeskorte

Neural network models can now recognise images, understand text, translate languages, and play many human games at human or superhuman levels. These systems are highly abstracted, but are inspired by biological brains and use only biologically plausible computations. In the coming years, neural networks are likely to become less reliant on learning from massive labelled datasets, and more robust and generalisable in their task performance. From their successes and failures, we can learn about the computational requirements of the different tasks at which brains excel. Deep learning also provides the tools for testing cognitive theories. In order to test a theory, we need to realise the proposed information-processing system at scale, so as to be able to assess its feasibility and emergent behaviours. Deep learning allows us to scale up from principles and circuit models to end-to-end trainable models capable of performing complex tasks. There are many levels at which cognitive neuroscientists can use deep learning in their work, from inspiring theories to serving as full computational models. Ongoing advances in deep learning bring us closer to understanding how cognition and perception may be implemented in the brain -- the grand challenge at the core of cognitive neuroscience.

Nikolaus Kriegeskorte, Tal Golan

Originally inspired by neurobiology, deep neural network models have become a powerful tool of machine learning and artificial intelligence, where they are used to approximate functions and dynamics by learning from examples. Here we give a brief introduction to neural network models and deep learning for biologists. We introduce feedforward and recurrent networks and explain the expressive power of this modeling framework and the backpropagation algorithm for setting the parameters. Finally, we consider how deep neural networks might help us understand the brain's computations.

Baihan Lin, Nikolaus Kriegeskorte

Testing two potentially multivariate variables for statistical dependence on the basis finite samples is a fundamental statistical challenge. Here we explore a family of tests that adapt to the complexity of the relationship between the variables, promising robust power across scenarios. Building on the distance correlation, we introduce a family of adaptive independence criteria based on nonlinear monotonic transformations of distances. We show that these criteria, like the distance correlation and RKHS-based criteria, provide dependence indicators. We propose a class of adaptive (multi-threshold) test statistics, which form the basis for permutation tests. These tests empirically outperform some of the established tests in average and worst-case statistical sensitivity across a range of univariate and multivariate relationships, offer useful insights to the data and may deserve further exploration.

Nikolaus Kriegeskorte, Robert M. Mok

Building machines that learn and think like humans is essential not only for cognitive science, but also for computational neuroscience, whose ultimate goal is to understand how cognition is implemented in biological brains. A new cognitive computational neuroscience should build cognitive-level and neural- level models, understand their relationships, and test both types of models with both brain and behavioral data.

Patrick McClure, Nikolaus Kriegeskorte

As deep neural networks (DNNs) are applied to increasingly challenging problems, they will need to be able to represent their own uncertainty. Modeling uncertainty is one of the key features of Bayesian methods. Using Bernoulli dropout with sampling at prediction time has recently been proposed as an efficient and well performing variational inference method for DNNs. However, sampling from other multiplicative noise based variational distributions has not been investigated in depth. We evaluated Bayesian DNNs trained with Bernoulli or Gaussian multiplicative masking of either the units (dropout) or the weights (dropconnect). We tested the calibration of the probabilistic predictions of Bayesian convolutional neural networks (CNNs) on MNIST and CIFAR-10. Sampling at prediction time increased the calibration of the DNNs' probabalistic predictions. Sampling weights, whether Gaussian or Bernoulli, led to more robust representation of uncertainty compared to sampling of units. However, using either Gaussian or Bernoulli dropout led to increased test set classification accuracy. Based on these findings we used both Bernoulli dropout and Gaussian dropconnect concurrently, which we show approximates the use of a spike-and-slab variational distribution without increasing the number of learned parameters. We found that spike-and-slab sampling had higher test set performance than Gaussian dropconnect and more robustly represented its uncertainty compared to Bernoulli dropout.

Patrick McClure, Nikolaus Kriegeskorte

Deep neural networks (DNNs) provide useful models of visual representational transformations. We present a method that enables a DNN (student) to learn from the internal representational spaces of a reference model (teacher), which could be another DNN or, in the future, a biological brain. Representational spaces of the student and the teacher are characterized by representational distance matrices (RDMs). We propose representational distance learning (RDL), a stochastic gradient descent method that drives the RDMs of the student to approximate the RDMs of the teacher. We demonstrate that RDL is competitive with other transfer learning techniques for two publicly available benchmark computer vision datasets (MNIST and CIFAR-100), while allowing for architectural differences between student and teacher. By pulling the student's RDMs towards those of the teacher, RDL significantly improved visual classification performance when compared to baseline networks that did not use transfer learning. In the future, RDL may enable combined supervised training of deep neural networks using task constraints (e.g. images and category labels) and constraints from brain-activity measurements, so as to build models that replicate the internal representational spaces of biological brains.