As the amount of correlation between the signals increases, the benefit derived from integrating these signals should decrease Jones, One possible implication is that the interactions between the signals are quite subtle, so that the benefit gained from integration is still quite high.
Another is that foveal and peripheral information can interact on numerous processing levels, and the interactions outlined in earlier sections occur on a different level to integration: in this case, the representations being integrated may not be subject to the same correlated noise.
Could integration rely on peripheral-foveal connections? Following from the previous point, we can speculate that transsaccadic perception may be facilitated by the peripheral-foveal connections outlined above Foveal-peripheral interactions during fixation. Processing a presaccadic peripheral stimulus in foveal cortex Williams et al. Paeye et al. How are moving objects integrated?
During the saccade, the object changes its position, and it is unclear whether the visual system is able to integrate pre- and postsaccadic information from different locations. What is the relationship between spatiotopic feature integration and retinotopic after effects?
The aforementioned studies showed integration in a strictly spatiotopic reference frame i. Critically, this happens even with very short presentation durations of the stimuli and at short saccade latencies. This is different from the transfer of some aftereffects e. This apparent discrepancy in results could be caused by different factors: adaptation and integration might simply tap into different levels of processing that are organized in different reference frames, and that operate on different time scales.
We have already discussed some open questions at the end of each of the previous sections, but there are, of course, additional overarching questions that we want to briefly sketch out here. As reviewed above Foveal-peripheral interactions during fixation , there is converging evidence that foveal feedback signals facilitate the recognition of objects in peripheral vision during fixation.
At present, it is unclear whether, and how, foveal feedback signals contribute to transsaccadic re-calibration and integration.
Both phenomena involve the joint processing of presaccadic, peripheral, and postsaccadic, foveal information. Irrespective of whether foveal feedback signals rely on the remapping of receptive fields or on another unknown neurophysiological mechanism, there is some evidence for a close link between foveal feedback effects and saccades. First, foveal feedback effects do not occur when a saccade is planned to another location Fan et al.
This suggests that the foveal feedback effect that is typically observed during fixation might be caused by covertly planning a saccade to the peripheral object. Second, the typical time course of foveal feedback signals of about to ms Fan et al. This would suggest that feedback information from presaccadic peripheral vision and feedforward information from postsaccadic foveal vision coincide at the same time in foveal cortex.
However, under cognitively demanding conditions, the time course of foveal feedback can also be much slower at about ms see Figure 2 C; Fan et al. Because it is unlikely that saccade latencies would be delayed to a similar extent, this would mean that feedforward information from postsaccadic foveal vision and feedback information from presaccadic peripheral vision can be temporarily decoupled.
It would be interesting to test if this temporal decoupling also eliminates transsaccadic integration, unlike re-calibration that seems to be rather independent of saccade execution and temporal contiguity Paeye et al. As we saw above Re-calibration of peripheral and foveal vision , the perception of a saccade stimulus feature can be re-calibrated based on transsaccadic changes to that feature. In principle, this might interact with the integration of peripheral and foveal information Transsaccadic integration of peripheral and foveal information.
Is the true physical value of the stimulus integrated, or the re-calibrated percept? The answer to this question depends on the level at which re-calibration occurs. If re-calibration causes a re-tuning of peripheral neurons based on foveal perception, then the answer would be necessarily positive, as the percept would reflect the physiological coding of the stimulus in early visual cortex, so the re-calibrated percept would be integrated.
As discussed earlier Open Questions in Re-calibration of peripheral and foveal vision , this associative learning approach would incorporate additional components that could be integrated: the true value of the stimulus, or the percept formed from a learned association. We do not know which of these elements would be integrated, but we can speculate that the answer may depend on the relative reliability of the true value of the signal versus the strength of the learned association: if the reliability of the true peripheral signal is extremely low, then the system may place more weight on prior learned knowledge.
In this review, we have summarized relevant literature to address three core questions on the interactions between peripheral and foveal vision.
Overall, a clear picture of a highly integrated visual system emerges, in which the differences between peripheral and foveal vision are reconciled by a diverse array of mechanisms: during fixation, peripheral vision is enhanced by using foveal feedback signals for object recognition and by extrapolating information from the fovea toward the periphery; a transsaccadic learning mechanism calibrates peripheral and foveal percepts; information from peripheral and foveal vision can be integrated across saccades to optimize the uptake of information.
This suggests that despite their large differences in processing and representation, peripheral and foveal vision are more intricately related than commonly assumed. The rod-free zone, up to a radius of about 0. The term fovea is used with various definitions, resulting in different estimations of size. Intermediate eccentricities between the fovea and the periphery are often called parafovea.
Because many perceptual measures continuously decline from the center of the visual field, and because most of the studies and effects we want to discuss in this review are agnostic about the exact delineation of the fovea, we use the term fovea colloquially to refer to the center of the visual field, and periphery to refer to the surrounding area.
This is reminiscent of the parafoveal preview benefit in reading for a review see Schotter et al. Although the direction of this effect is opposite to the foveal-feedback effect, the two effects may be related to each other. Although object and scene information cannot be unequivocally mapped to foveal and peripheral vision, there is some evidence that this effect is present specifically for the interaction of periphery and fovea e.
Roux-Sibilon et al. We do not discuss it here because it does not pertain directly to the relationship between peripheral and foveal perception. We point the reader to the review by Zimmermann and Lappe for a discussion of the possible mechanisms subtending this phenomenon. Melcher, and developing more slowly e. National Center for Biotechnology Information , U. J Vis. Published online Nov 3. Emma E. Stewart , 1 , 1 Matteo Valsecchi , 2 , 2 and Alexander C. Alexander C. Author information Article notes Copyright and License information Disclaimer.
Email: moc. Email: ti. Received Jun 3; Accepted Sep Copyright The Authors. This work is licensed under a Creative Commons Attribution 4. This article has been cited by other articles in PMC. Abstract Visual processing varies dramatically across the visual field.
Keywords: Peripheral, foveal, review. Brief overview of differences between peripheral and foveal vision Although the human eye is often compared to a photographic camera, processing across the visual field is not homogeneous like in a camera film or a digital sensor. Open in a separate window. Figure 1. Foveal-peripheral interactions during fixation The aforementioned literature has described clear differences between foveal and peripheral vision on both a physiological level and a perceptual level.
Foveal feedback signals supporting peripheral object recognition A rather recent finding in the history of vision science concerns the processing of peripherally displayed stimuli in foveal retinotopic cortex. Figure 2. Figure 3. Open questions Foveal feedback signals and extrapolation are both rather recent empirical findings and therefore our knowledge about those effects is still scarce, leaving many open questions for further investigation.
Re-calibration of peripheral and foveal vision As we have outlined in the introduction, peripheral and foveal vision differ in many aspects, and, for the most part, peripheral vision achieves the ability to represent a large portion of the visual field by giving up the ability to represent all the elements that it might contain individually, and with high acuity. The idea that transsaccadic learning subtends our ability to match foveal and peripheral appearance is not new, and was already expressed very clearly by Hermann von Helmholtz in his Treatise on Physiological Optics Helmholtz, ; Quote from the English translation, : Now when we perceive any object in indirect vision, and thus have received a limited impression of it on a peripheral part of the retina, and then turn the eye so as to look straight at it, we get afterwards an impression of the same object with the same apparent size on the center of the retina; and thus we can gradually learn by experience when a certain peripheral impression is the same in quality and size as a central impression.
In particular, they clearly predicted that a repeated transsaccadic change would have the effect of unifying the two physically different stimuli, sensed foveally and peripherally, into a common phenomenological experience, for instance, in the case of color appearance: Using a device to measure eye movements connected to a computer, it should be possible to arrange stimulation on a display screen so that whenever an observer looks directly at a patch of colour it appears red, but whenever the observer's eye looks away from the patch, its color changes to green.
Gaze-contingent color perception The first experiments on gaze-contingent color perception were conducted by Kohler Figure 4. Open questions In general, the studies on gaze-contingent color perception and those on transsaccadic re-calibration have shown that visuomotor experience can help us to accommodate some of the sensory differences that exist between peripheral and foveal vision. Transsaccadic integration of peripheral and foveal information The execution of a saccadic eye movement leads to a large discontinuity in visual processing and the visual system must reconcile the presaccadic, peripheral view of an object or location, with its postsaccadic, foveal counterpart.
Figure 5. Integration of feature information across saccades This section covers three broad themes of transsaccadic integration: how peripheral information can directly alter the postsaccadic percept, how the combination of peripheral and foveal information can result in a weighted average, and how the utilisation of both peripheral and foveal information is beneficial to perception. Presaccadic information alters postsaccadic perception We have seen that foveal and peripheral information interact to alter perception, both during fixation and during saccades through re-calibration.
Figure 6. Feature integration: Reliability benefit The concept of reliability is central to the next theme of integration that will be discussed in the following section: optimal feature integration see Figure 5. What is integrated, and how does integration occur?
Where in the visual field does integration occur? The first such limitation is one that was again foreshadowed by Helmholtz , English translation, : The peculiar ultimate basis, which gives convincing power to all our conscious inductions, is the law of causation. Open questions 1. Future directions We have already discussed some open questions at the end of each of the previous sections, but there are, of course, additional overarching questions that we want to briefly sketch out here.
What is the role of foveal feedback signals for transsaccadic perception? Is re-calibrated information integrated?
Conclusions In this review, we have summarized relevant literature to address three core questions on the interactions between peripheral and foveal vision. Commercial relationships: none. Corresponding author: Alexander C.
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Journal of Vision, 6 12 : 4, —, The hypothesis of an optimized gaze behavior was addressed by Vater et al. Based on the findings from 29 studies covering different types of sports and tasks, levels of expertise and methods applied, they suggested three different, task-dependent functionalities of peripheral vision as indicated by different gaze behaviors.
If the task requires and the situation allows one to focus on one specific aspect-as for example the basketball free throw with high precision and low time demands-gaze is stabilized on one specific cue-like the rim of the basket-to get accurate visual information. If, however, the task still requires to process visual information with high spatial acuity but more than one cue prevails crucial information-like in 2 vs.
This gaze behavior was labeled foveal spot Figure 1 , middle. Figure 1. Three types of gaze behavior for the use of foveal and peripheral vision in sports modified from Vater et al.
It should be noted that the small white spot illustrates the current gaze point and the grayish circled areas indicate potential attention locations as a function of respective relevant cues. Finally, in case of a foveal spot the breadth of attention can be modified as a function of the task demands. In a number of studies e. For example, in beach-volleyball defense Hossner et al.
It was assumed, that in these situations high spatial acuity is not required otherwise gaze would be positioned to make use of foveal vision and athletes use a gaze anchor Figure 1 , right which allows one to monitor the movement of objects with peripheral vision see also Vater et al.
In contrast to the foveal spot, covert attention can be distributed to multiple objects. This, however, requires an optimal positioning of the gaze as it was shown by Hausegger et al. Specifically, if attacks could be performed with arms and legs Qwan Ki Do , athletes positioned the gaze anchor higher at the opponent's body compared to situations where mainly the legs are used to attack Tae Kwan Do. Finally, a visual pivot is used in situations that demand the processing of accurate visual information from a number of spatially distributed cues that cannot be covered by para- foveal vision alone and requires to reposition the fovea Figure 1 , left.
As for the gaze anchor, it is hypothesized that the visual pivot is optimally located in-between the relevant information sources to allow for frequent fixation transitions by minimal saccadic costs. The visual pivot can be located in free space but also on information-rich cues, like the opponent's hip in soccer-defense situations e. As emphasized by Vater et al. As the situation evolves, different gaze behavior might interact, as for example in a basketball-offense situation in which the ball carrier, first might anchor his gaze centrally between his teammates to evaluate who might receive a pass.
In the next moment, however, he might decide to shoot and to foveally spot the rim of the basket. Though, whether, indeed, the gaze positioned at the rim of the basket is used as foveal spot or as pivot point to still fixate the teammate which runs into an optimal playing position is difficult to determine by gaze data alone, because the gaze location might not indicate information processing. As emphasized above, due to the complex interactions as they occur in sports with highly dynamic situational conditions and motor tasks with different demands, foveal and peripheral vision are necessary to be successful.
Consequently, the mechanisms introduced are not mutually exclusive but, rather, complementary. We suggest that constraints inherent in the task require to balance an optimal positioning of the gaze with an optimally early onset of this fixation before movement initiation.
The former depends on potential costs that occur when repositioning the fovea using a saccade i. The latter is necessary to facilitate movement parameterization via inhibition processes, thus, is highly dependent on the demands associated with the task.
Consequently, an optimal gaze-anchoring location might not only be useful for processing peripheral information i. Moreover, the positioning and the relative onset of this gaze-anchoring should be highly dependent on the associated cost-benefit equation. To the best of our knowledge, this hypothesis has not been addressed yet and will be pursued in future research projects in our laboratory.
Directly related to these questions, fundamental questions on the role of vision and attention in complex movement behavior should be made clear. As explained earlier, visual capabilities largely influence our gaze behavior and the use of foveal and peripheral vision.
The interaction with attentional capabilities, however, and particularly research directly combining attention and vision measures in sport has been addressed only rudimentarily.
Piras et al. But, this approach challenges eye-tracking technology, particularly when investigating complex movement behavior. Therefore, experimental approaches should be favored as proposed by Vater et al. For example, based on previous research, it can be predicted that the ball carrier in soccer is the most likely gaze location.
If one is able to react to a peripheral player's action without looking away from the ball carrier, this could be seen as an indicator of peripheral vision usage.
Finally, sport scientists should aim to transfer the obtained knowledge back into sports practice to improve the athletes' skills. The so-called perceptual training draws on the expert-performance approach e. Although a fair number of studies examined the effectiveness of these training programs, more often than not, research failed to show significant transfer effects for an overview, e.
In addition to issues with ecological validity e. For example, research suggests that so-called gaze training by means of attentional cueing does not foster the learning of anticipation skills in beach-volleyball when compared to active control groups e.
Therefore, in the next years, increased efforts should be devoted to the theory-practice transfer answering questions like the trainability of expert-like gaze behavior in sports. All authors listed have made a substantial, direct and intellectual contribution to the work, and approved it for publication. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
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Ericsson, K. Ericsson and J. Smith Cambridge: Cambridge University Press , 1— Google Scholar. Because many perceptual measures continuously decline from the center of the visual field, and because most of the studies and effects we want to discuss in this review are agnostic about the exact delineation of the fovea, we use the term fovea colloquially to refer to the center of the visual field, and periphery to refer to the surrounding area. This is reminiscent of the parafoveal preview benefit in reading for a review see Schotter et al.
Although the direction of this effect is opposite to the foveal-feedback effect, the two effects may be related to each other. Although object and scene information cannot be unequivocally mapped to foveal and peripheral vision, there is some evidence that this effect is present specifically for the interaction of periphery and fovea e. Roux-Sibilon et al. We do not discuss it here because it does not pertain directly to the relationship between peripheral and foveal perception. We point the reader to the review by Zimmermann and Lappe for a discussion of the possible mechanisms subtending this phenomenon.
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