Will Xiao

Spandan Madan, Will Xiao, Mingran Cao et al.

We characterized the generalization capabilities of DNN-based encoding models when predicting neuronal responses from the visual cortex. We collected \textit{MacaqueITBench}, a large-scale dataset of neural population responses from the macaque inferior temporal (IT) cortex to over $300,000$ images, comprising $8,233$ unique natural images presented to seven monkeys over $109$ sessions. Using \textit{MacaqueITBench}, we investigated the impact of distribution shifts on models predicting neural activity by dividing the images into Out-Of-Distribution (OOD) train and test splits. The OOD splits included several different image-computable types including image contrast, hue, intensity, temperature, and saturation. Compared to the performance on in-distribution test images -- the conventional way these models have been evaluated -- models performed worse at predicting neuronal responses to out-of-distribution images, retaining as little as $20\%$ of the performance on in-distribution test images. The generalization performance under OOD shifts can be well accounted by a simple image similarity metric -- the cosine distance between image representations extracted from a pre-trained object recognition model is a strong predictor of neural predictivity under different distribution shifts. The dataset of images, neuronal firing rate recordings, and computational benchmarks are hosted publicly at: https://bit.ly/3zeutVd.

Mengmi Zhang, Marcelo Armendariz, Will Xiao et al.

Primates constantly explore their surroundings via saccadic eye movements that bring different parts of an image into high resolution. In addition to exploring new regions in the visual field, primates also make frequent return fixations, revisiting previously foveated locations. We systematically studied a total of 44,328 return fixations out of 217,440 fixations. Return fixations were ubiquitous across different behavioral tasks, in monkeys and humans, both when subjects viewed static images and when subjects performed natural behaviors. Return fixations locations were consistent across subjects, tended to occur within short temporal offsets, and typically followed a 180-degree turn in saccadic direction. To understand the origin of return fixations, we propose a proof-of-principle, biologically-inspired and image-computable neural network model. The model combines five key modules: an image feature extractor, bottom-up saliency cues, task-relevant visual features, finite inhibition-of-return, and saccade size constraints. Even though there are no free parameters that are fine-tuned for each specific task, species, or condition, the model produces fixation sequences resembling the universal properties of return fixations. These results provide initial steps towards a mechanistic understanding of the trade-off between rapid foveal recognition and the need to scrutinize previous fixation locations.

Li Yuan, Will Xiao, Giorgia Dellaferrera et al.

Artificial neural networks (ANNs) are considered the current best models of biological vision. ANNs are the best predictors of neural activity in the ventral stream; moreover, recent work has demonstrated that ANN models fitted to neuronal activity can guide the synthesis of images that drive pre-specified response patterns in small neuronal populations. Despite the success in predicting and steering firing activity, these results have not been connected with perceptual or behavioral changes. Here we propose an array of methods for creating minimal, targeted image perturbations that lead to changes in both neuronal activity and perception as reflected in behavior. We generated 'deceptive images' of human faces, monkey faces, and noise patterns so that they are perceived as a different, pre-specified target category, and measured both monkey neuronal responses and human behavior to these images. We found several effective methods for changing primate visual categorization that required much smaller image change compared to untargeted noise. Our work shares the same goal with adversarial attack, namely the manipulation of images with minimal, targeted noise that leads ANN models to misclassify the images. Our results represent a valuable step in quantifying and characterizing the differences in perturbation robustness of biological and artificial vision.

Will Xiao, Gabriel Kreiman

A fundamental question for understanding brain function is what types of stimuli drive neurons to fire. In visual neuroscience, this question has also been posted as characterizing the receptive field of a neuron. The search for effective stimuli has traditionally been based on a combination of insights from previous studies, intuition, and luck. Recently, the same question has emerged in the study of units in convolutional neural networks (ConvNets), and together with this question a family of solutions were developed that are generally referred to as "feature visualization by activation maximization." We sought to bring in tools and techniques developed for studying ConvNets to the study of biological neural networks. However, one key difference that impedes direct translation of tools is that gradients can be obtained from ConvNets using backpropagation, but such gradients are not available from the brain. To circumvent this problem, we developed a method for gradient-free activation maximization by combining a generative neural network with a genetic algorithm. We termed this method XDream (EXtending DeepDream with real-time evolution for activation maximization), and we have shown that this method can reliably create strong stimuli for neurons in the macaque visual cortex (Ponce et al., 2019). In this paper, we describe extensive experiments characterizing the XDream method by using ConvNet units as in silico models of neurons. We show that XDream is applicable across network layers, architectures, and training sets; examine design choices in the algorithm; and provide practical guides for choosing hyperparameters in the algorithm. XDream is an efficient algorithm for uncovering neuronal tuning preferences in black-box networks using a vast and diverse stimulus space.

Will Xiao, Honglin Chen, Qianli Liao et al.

The backpropagation (BP) algorithm is often thought to be biologically implausible in the brain. One of the main reasons is that BP requires symmetric weight matrices in the feedforward and feedback pathways. To address this "weight transport problem" (Grossberg, 1987), two more biologically plausible algorithms, proposed by Liao et al. (2016) and Lillicrap et al. (2016), relax BP's weight symmetry requirements and demonstrate comparable learning capabilities to that of BP on small datasets. However, a recent study by Bartunov et al. (2018) evaluate variants of target-propagation (TP) and feedback alignment (FA) on MINIST, CIFAR, and ImageNet datasets, and find that although many of the proposed algorithms perform well on MNIST and CIFAR, they perform significantly worse than BP on ImageNet. Here, we additionally evaluate the sign-symmetry algorithm (Liao et al., 2016), which differs from both BP and FA in that the feedback and feedforward weights share signs but not magnitudes. We examine the performance of sign-symmetry and feedback alignment on ImageNet and MS COCO datasets using different network architectures (ResNet-18 and AlexNet for ImageNet, RetinaNet for MS COCO). Surprisingly, networks trained with sign-symmetry can attain classification performance approaching that of BP-trained networks. These results complement the study by Bartunov et al. (2018), and establish a new benchmark for future biologically plausible learning algorithms on more difficult datasets and more complex architectures.