The eyes anticipate where objects will move based on their shape.

Imagine staring into a clear river, starving, desperately searching for a fish to spear and cook. You see a dark shape lurking beneath the surface. It doesn't resemble any sort of fish you've encountered before - but you're hungry. To catch it, you need to anticipate which way it will move when you lunge for it, to compensate for your own sensory and motor processing delays1,2,3. Yet you know nothing about the behaviour of this creature, and do not know in which direction it will try to escape. What cues do you then use to drive such anticipatory responses? Fortunately, many species4, including humans, have the remarkable ability to predict the directionality of objects based on their shape - even if they are unfamiliar and so we cannot rely on semantic knowledge about their movements5. While it is known that such directional inferences can guide attention5, we do not yet fully understand how such causal inferences are made, or the extent to which they enable anticipatory behaviours. Does the oculomotor system, which moves our eyes to optimise visual input, use directional inferences from shape to anticipate upcoming motion direction? Such anticipation is necessary to stabilise the moving object on the high-resolution fovea of the retina while tracking the shape, a primary goal of the oculomotor system6, and to guide any future interactions7,8. Here, we leveraged a well-known behaviour of the oculomotor system: anticipatory smooth eye movements (ASEM), where an increase in eye velocity is observed in the direction of a stimulus' expected motion, before the stimulus actually moves3, to show that the oculomotor system extracts directional information from shape, and uses this inference to predict and anticipate upcoming motion.

Mental object rotation based on two-dimensional visual representations.

The discovery of mental rotation was one of the most significant landmarks in experimental psychology, leading to the ongoing assumption that to visually compare objects from different three-dimensional viewpoints, we use explicit internal simulations of object rotations, to 'mentally adjust' one object until it matches the other1. These rotations are thought to be performed on three-dimensional representations of the object, by literal analogy to physical rotations. In particular, it is thought that an imagined object is continuously adjusted at a constant three-dimensional angular rotation rate from its initial orientation to the final orientation through all intervening viewpoints2. While qualitative theories have tried to account for this phenomenon3, to date there has been no explicit, image-computable model of the underlying processes. As a result, there is no quantitative account of why some object viewpoints appear more similar to one another than others when the three-dimensional angular difference between them is the same4,5. We reasoned that the specific pattern of non-uniformities in the perception of viewpoints can reveal the visual computations underlying mental rotation. We therefore compared human viewpoint perception with a model based on the kind of two-dimensional 'optical flow' computations that are thought to underlie motion perception in biological vision6, finding that the model reproduces the specific errors that participants make. This suggests that mental rotation involves simulating the two-dimensional retinal image change that would occur when rotating objects. When we compare objects, we do not do so in a distal three-dimensional representation as previously assumed, but by measuring how much the proximal stimulus would change if we watched the object rotate, capturing perspectival appearance changes7.

Serial dependence for oculomotor control depends on early sensory signals.

To create an accurate percept of the world, the visual system relies on past experience and prior assumptions.1 For example, although the retinal projection of an object moving in depth changes drastically, we still perceive the object at a constant size and velocity.2,3 Consequently, if we see the same object with a constant retinal size at two different depth levels, the perceived size differs (illustrated by the Ponzo illusion). Past experience also directly influences perceptual judgments, an effect known as serial dependence.4,5 Such sequential effects have also been reported for oculomotor behavior, even on the trial-by-trial level.6-10 An integration of past experiences seems like a smart and sophisticated mechanism to reduce uncertainty and improve behavior in a world full of statistical regularities. By leveraging the Ponzo illusion to dissociate perceived size and speed from retinal signals, we show that serial-dependence effects for oculomotor control are mediated by retinal error signals. These sequential effects likely take place in early sensory processing because they transfer to different visual stimuli. In contrast to recently reported history effects for perceptual decisions,11 sequential effects for oculomotor control deviate from perceptual mechanisms by not integrating spatial context and by ignoring size and velocity constancy. Although this dissociation might appear suboptimal, we argue that this effect reveals the different goals of the oculomotor and perceptual systems. The oculomotor system tries to reduce retinal error signals to bring and keep the target close to the fovea, whereas the visual system interprets retinal input to achieve an accurate representation of the world.12.

Humans represent the precision and utility of information acquired across fixations.

Our environment contains an abundance of objects which humans interact with daily, gathering visual information using sequences of eye-movements to choose which object is best-suited for a particular task. This process is not trivial, and requires a complex strategy where task affordance defines the search strategy, and the estimated precision of the visual information gathered from each object may be used to track perceptual confidence for object selection. This study addresses the fundamental problem of how such visual information is metacognitively represented and used for subsequent behaviour, and reveals a complex interplay between task affordance, visual information gathering, and metacogntive decision making. People fixate higher-utility objects, and most importantly retain metaknowledge about how much information they have gathered about these objects, which is used to guide perceptual report choices. These findings suggest that such metacognitive knowledge is important in situations where decisions are based on information acquired in a temporal sequence.

A review of interactions between peripheral and foveal vision.

Visual processing varies dramatically across the visual field. These differences start in the retina and continue all the way to the visual cortex. Despite these differences in processing, the perceptual experience of humans is remarkably stable and continuous across the visual field. Research in the last decade has shown that processing in peripheral and foveal vision is not independent, but is more directly connected than previously thought. We address three core questions on how peripheral and foveal vision interact, and review recent findings on potentially related phenomena that could provide answers to these questions. First, how is the processing of peripheral and foveal signals related during fixation? Peripheral signals seem to be processed in foveal retinotopic areas to facilitate peripheral object recognition, and foveal information seems to be extrapolated toward the periphery to generate a homogeneous representation of the environment. Second, how are peripheral and foveal signals re-calibrated? Transsaccadic changes in object features lead to a reduction in the discrepancy between peripheral and foveal appearance. Third, how is peripheral and foveal information stitched together across saccades? Peripheral and foveal signals are integrated across saccadic eye movements to average percepts and to reduce uncertainty. Together, these findings illustrate that peripheral and foveal processing are closely connected, mastering the compromise between a large peripheral visual field and high resolution at the fovea.

Stronger saccadic suppression of displacement and blanking effect in children.

Humans do not notice small displacements to objects that occur during saccades, termed saccadic suppression of displacement (SSD), and this effect is reduced when a blank is introduced between the pre- and postsaccadic stimulus (Bridgeman, Hendry, & Stark, 1975; Deubel, Schneider, & Bridgeman, 1996). While these effects have been studied extensively in adults, it is unclear how these phenomena are characterized in children. A potentially related mechanism, saccadic suppression of contrast sensitivity-a prerequisite to achieve a stable percept-is stronger for children (Bruno, Brambati, Perani, & Morrone, 2006). However, the evidence for how transsaccadic stimulus displacements may be suppressed or integrated is mixed. While they can integrate basic visual feature information from an early age, they cannot integrate multisensory information (Gori, Viva, Sandini, & Burr, 2008; Nardini, Jones, Bedford, & Braddick, 2008), suggesting a failure in the ability to integrate more complex sensory information. We tested children 7 to 12 years old and adults 19 to 23 years old on their ability to perceive intrasaccadic stimulus displacements, with and without a postsaccadic blank. Results showed that children had stronger SSD than adults and a larger blanking effect. Children also had larger undershoots and more variability in their initial saccade endpoints, indicating greater intrinsic uncertainty, and they were faster in executing corrective saccades to account for these errors. Together, these results suggest that children may have a greater internal expectation or prediction of saccade error than adults; thus, the stronger SSD in children may be due to higher intrinsic uncertainty in target localization or saccade execution.

Transsaccadic integration benefits are not limited to the saccade target.

Across saccades, humans can integrate the low-resolution presaccadic information of an upcoming saccade target with the high-resolution postsaccadic information. There is converging evidence to suggest that transsaccadic integration occurs at the saccade target. However, given divergent evidence on the spatial specificity of related mechanisms such as attention, visual working memory, and remapping, it is unclear whether integration is also possible at locations other than the saccade target. We tested the spatial profile of transsaccadic integration, by testing perceptual performance at six locations around the saccade target and between the saccade target and initial fixation. Results show that integration benefits do not differ between the saccade target and surrounding locations. Transsaccadic integration benefits are not specific to the saccade target and can occur at other locations when they are behaviorally relevant, although there is a trend for worse performance for the location above initial fixation compared with those in the direction of the saccade. This suggests that transsaccadic integration may be a more general mechanism used to reconcile task-relevant pre- and postsaccadic information at attended locations other than the saccade target.NEW & NOTEWORTHY This study shows that integration of pre- and postsaccadic information across saccades is not restricted to the saccade target. We found performance benefits of transsaccadic integration at attended locations other than the saccade target, and these benefits did not differ from those found at the saccade target. This suggests that transsaccadic integration may be a more general mechanism used to reconcile pre- and postsaccadic information at task-relevant locations.

The spatial and temporal properties of attentional selectivity for saccades and reaches.

The preparation and execution of saccades and goal-directed movements elicits an accompanying shift in attention at the locus of the impending movement. However, some key aspects of the spatiotemporal profile of this attentional shift between eye and hand movements are not resolved. While there is evidence that attention is improved at the target location when making a reach, it is not clear how attention shifts over space and time around the movement target as a saccade and a reach are made to that target. Determining this spread of attention is an important aspect in understanding how attentional resources are used in relation to movement planning and guidance in real world tasks. We compared performance on a perceptual discrimination paradigm during a saccade-alone task, reach-alone task, and a saccade-plus-reach task to map the temporal profile of the premotor attentional shift at the goal of the movement and at three surrounding locations. We measured performance relative to a valid baseline level to determine whether motor planning induces additional attentional facilitation compared to mere covert attention. Sensitivity increased relative to movement onset at the target and at the surrounding locations, for both the saccade-alone and saccade-plus-reach conditions. The results suggest that the temporal profile of the attentional shift is similar for the two tasks involving saccades (saccade-alone and saccade-plus-reach tasks), but is very different when the influence of the saccade is removed. In this case, performance in the saccade-plus-reach task reflects the lower sensitivity observed when a reach-alone task is being conducted. In addition, the spatial profile of this spread of attention is not symmetrical around the target. This suggests that when a saccade and reach are being planned together, the saccade drives the attentional shift, and the reach-alone carries little attentional weight.

Transsaccadic integration is dominated by early, independent noise.

Humans are able to integrate pre- and postsaccadic percepts of an object across saccades to maintain perceptual stability. Previous studies have used Maximum Likelihood Estimation (MLE) to determine that integration occurs in a near-optimal manner. Here, we compared three different models to investigate the mechanism of integration in more detail: an early noise model, where noise is added to the pre- and postsaccadic signals before integration occurs; a late-noise model, where noise is added to the integrated signal after integration occurs; and a temporal summation model, where integration benefits arise from the longer transsaccadic presentation duration compared to pre- and postsaccadic presentation only. We also measured spatiotemporal aspects of integration to determine whether integration can occur for very brief stimulus durations, across two hemifields, and in spatiotopic and retinotopic coordinates. Pre-, post-, and transsaccadic performance was measured at different stimulus presentation durations, both at the saccade target and a location where the pre- and postsaccadic stimuli were presented in different hemifields across the saccade. Results showed that for both within- and between-hemifields conditions, integration could occur when pre- and postsaccadic stimuli were presented only briefly, and that the pattern of integration followed an early noise model. Whereas integration occurred when the pre- and post-saccadic stimuli were presented in the same spatiotopic coordinates, there was no integration when they were presented in the same retinotopic coordinates. This contrast suggests that transsaccadic integration is limited by early, independent, sensory noise acting separately on pre- and postsaccadic signals.

Optimal trans-saccadic integration relies on visual working memory.

Saccadic eye movements alter the visual processing of objects of interest by bringing them from the periphery, where there is only low-resolution vision, to the high-resolution fovea. Evidence suggests that people are able to achieve trans-saccadic integration in a near-optimal manner; however the mechanisms underlying integration are still unclear. Visual working memory (VWM) is sustained across a saccade, and it has been suggested that this memory resource is used to store and compare the pre- and post- saccadic percepts. This study directly tested the hypothesis that VWM is necessary for optimal trans-saccadic integration, by introducing memory load during a saccade, and testing subsequent integration performance on feature similar and dissimilar stimuli. Results show that integration performance was impaired when there was an additional memory task. Additionally, performance on the memory task was affected by feature-specific integration stimuli. Our results suggest that VWM supports the integration of pre- and post- saccadic stimuli because integration performance is impaired under VWM load.

Attention modulates trans-saccadic integration.

With every saccade, humans must reconcile the low resolution peripheral information available before a saccade, with the high resolution foveal information acquired after the saccade. While research has shown that we are able to integrate peripheral and foveal vision in a near-optimal manner, it is still unclear which mechanisms may underpin this important perceptual process. One potential mechanism that may moderate this integration process is visual attention. Pre-saccadic attention is a well documented phenomenon, whereby visual attention shifts to the location of an upcoming saccade before the saccade is executed. While it plays an important role in other peri-saccadic processes such as predictive remapping, the role of attention in the integration process is as yet unknown. This study aimed to determine whether the presentation of an attentional distractor during a saccade impaired trans-saccadic integration, and to measure the time-course of this impairment. Results showed that presenting an attentional distractor impaired integration performance both before saccade onset, and during the saccade, in selected subjects who showed integration in the absence of a distractor. This suggests that visual attention may be a mechanism that facilitates trans-saccadic integration.

The profile of attention differs between locations orthogonal to and in line with reach direction.

People make movements in a variety of directions when interacting with the world around them. It has been well documented that attention shifts to the goal of an upcoming movement, whether the movement is a saccade or a reach. However, recent evidence suggests that the direction of a movement may influence the spatial spread of attention (Stewart & Ma-Wyatt, 2015, Journal of Vision, 15(5), 10). We investigated whether the spatiotemporal profile of attention differs depending on where that location is situated relative to the direction of movement, and if this pattern is consistent across different movement effectors. We compared attentional facilitation at locations in line with or orthogonal to the movement, for reach-only, reach-plus-saccade, and saccade-only conditions. Results show that the spatiotemporal profile of attention differs across different movement combinations, and is also different at target locations orthogonal to and in line with the movement direction. Specifically, when a reach alone was made, there was a general decrease in attention at all locations during the movement and a general increase in attention at all locations with a saccade only. However, the concurrent reach and saccade condition showed a premovement attentional facilitation at locations orthogonal to movement direction, but not those in line with the movement direction. These results suggest attentional guidance may be more important at differing time points, depending on the type of movement.

The structure of the Caenorhabditis elegans manganese superoxide dismutase MnSOD-3-azide complex.

C. elegans MnSOD-3 has been implicated in the longevity pathway and its mechanism of catalysis is relevant to the aging process and carcinogenesis. The structures of MnSOD-3 provide unique crystallographic evidence of a dynamic region of the tetrameric interface (residues 41-54). We have determined the structure of the MnSOD-3-azide complex to 1.77-Å resolution. Analysis of this complex shows that the substrate analog, azide, binds end-on to the manganese center as a sixth ligand and that it ligates directly to a third and new solvent molecule also positioned within interacting distance to the His30 and Tyr34 residues of the substrate access funnel. This is the first structure of a eukaryotic MnSOD-azide complex that demonstrates the extended, uninterrupted hydrogen-bonded network that forms a proton relay incorporating three outer sphere solvent molecules, the substrate analog, the gateway residues, Gln142, and the solvent ligand. This configuration supports the formation and release of the hydrogen peroxide product in agreement with the 5-6-5 catalytic mechanism for MnSOD. The high product dissociation constant k4 of MnSOD-3 reflects low product inhibition making this enzyme efficient even at high levels of superoxide.

The spatiotemporal characteristics of the attentional shift relative to a reach.

While the attentional shift preceding a saccadic eye movement has been well documented, the mechanisms surrounding the attentional shift preceding a reach are not well understood. It is unknown whether these mechanisms may be the same as those used in perceptual tasks, or those used in the planning of a saccade. We mapped the spatiotemporal properties of attention relative to a reach to determine the time course of attentional facilitation for hand movements alone. Participants had to reach toward a target and during the reach a perceptual probe could appear at one of six locations around the target, and at nine temporal offsets relative to the cue. Results showed a consistent pattern of facilitation in the planning stages of the reach, with attention increasing and then reaching a plateau during the completion of the movement before dropping off. These results demonstrate that planning a hand movement necessitates a shift in attention across the visual field around 150 ms before the onset of a reach. While these results are broadly consistent with the results of experiments mapping attentional shifts for saccades, the spatiotemporal profile of facilitation found shows that reaching without a concurrent eye movement also causes shifts in attention across the visual field. These results also suggest that the profile of the attentional shift preceding and during a hand movement is different at different locations across the visual field.

The ontogeny of color: developmental origins of divergent pigmentation in Drosophila americana and D. novamexicana.

Pigmentation is a model trait for evolutionary and developmental analysis that is particularly amenable to molecular investigation in the genus Drosophila. To better understand how this phenotype evolves, we examined divergent pigmentation and gene expression over developmental time in the dark-bodied D. americana and its light-bodied sister species D. novamexicana. Prior genetic analysis implicated two enzyme-encoding genes, tan and ebony, in pigmentation divergence between these species, but questions remain about the underlying molecular mechanisms. Here, we describe stages of pupal development in both species and use this staging to determine when pigmentation develops and diverges between D. americana and D. novamexicana. For the developmental stages encompassing pigment divergence, we compare mRNA expression of tan and ebony over time and between species. Finally, we use allele-specific expression assays to determine whether interspecific differences in mRNA abundance have a cis-regulatory basis and find evidence of cis-regulatory divergence for both tan and ebony. cis-regulatory divergence affecting tan had a small effect on mRNA abundance and was limited to a few developmental stages, yet previous data suggests that this divergence is likely to be biologically meaningful. Our study suggests that small and developmentally transient expression changes may contribute to phenotypic diversification more often than commonly appreciated. Recognizing the potential phenotypic impact of such changes is important for a scientific community increasingly focused on dissecting quantitative variation, but detecting these types of changes will be a major challenge to elucidating the molecular basis of complex traits.

Intraspecific polymorphism to interspecific divergence: genetics of pigmentation in Drosophila.

Genetic changes contributing to phenotypic differences within or between species have been identified for a handful of traits, but the relationship between alleles underlying intraspecific polymorphism and interspecific divergence is largely unknown. We found that noncoding changes in the tan gene, as well as changes linked to the ebony gene, contribute to pigmentation divergence between closely related Drosophila species. Moreover, we found that alleles linked to tan and ebony fixed in one Drosophila species also contribute to variation within another species, and that multiple genotypes underlie similar phenotypes even within the same population. These alleles appear to predate speciation, which suggests that standing genetic variation present in the common ancestor gave rise to both intraspecific polymorphism and interspecific divergence.

Purification, crystallization and X-ray structures of the two manganese superoxide dismutases from Caenorhabditis elegans.

Caenorhabditis elegans expresses two manganese superoxide dismutase enzymes (MnSOD-2 and MnSOD-3) that are targeted to the mitochondrion. MnSOD-2 is constitutively expressed, while synthesis of MnSOD-3 is inducible. The structures of these two mononuclear metalloenzymes have been determined to 1.8 and 1.7 A resolution, respectively. Pink crystals formed in space group P4(1)2(1)2 for each, with unit-cell parameters a = b = 81.0, c = 137.4 A for MnSOD-2 and a = b = 81.8, c = 136.0 A for MnSOD-3. The final structure of MnSOD-3 was refined to R = 21.6% and R(free) = 26.2% at 293 K, and R = 18.9% and R(free) = 22.6% at 100 K, while that of MnSOD-2 was refined to R = 16.9% and R(free) = 20.1% at 100 K. The asymmetric unit cell is comprised of two subunits. The resulting structures are very similar to that of human MnSOD and form a tetramer corresponding to a dimer of dimers. The subunit interface between dimers is comprised of two four-helix bundles that stabilize the biologically significant homotetramer.