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Neuroscience and Biobehavioral Reviews 52 (2015) 105–116

Contents lists available at ScienceDirect

Neuroscience and Biobehavioral Reviews

jou rn al h om epage: www.elsev ier .com/ locate /neubiorev

eview

inking perception and action by structure or process? Toward anntegrative perspective

rvid Herwig ∗

epartment of Psychology and Cluster of Excellence, “Cognitive Interaction Technology”, Bielefeld University, Bielefeld, Germany

r t i c l e i n f o

rticle history:eceived 1 September 2014eceived in revised form 19 February 2015ccepted 22 February 2015vailable online 27 February 2015

eywords:deomotor theory

a b s t r a c t

Over the past decades cognitive neuroscience’s renewed interest in action has intensified the search ofprinciples explaining how the cognitive system links perception to action and vice versa. To date, atleast two seemingly alternative approaches can be distinguished. Perception and action might be linkedeither by common representational structures, as assumed by the ideomotor approach, or by commonattentional processes, as assumed by the attention approach. This article first reviews the evidence fromdifferent paradigms supporting each approach. It becomes clear that most studies selectively focus eitheron actions directed at goals outside the actors’ perceptual range (supporting the ideomotor approach) or

ttentionye movementsredictionnticipationriority map

on actions directed at targets within the actors’ perceptual range (supporting the attention approach).In a second step, I will try to reconcile both approaches by reviewing recent eye movement studies thatabolish the classical combination of approach and goals under study. Demonstrating that both approachescover target- as well as goal-directed actions, it is proposed that operations addressed in both conceptualframeworks interact with each other.

© 2015 Elsevier Ltd. All rights reserved.

ontents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1062. From targets in the world to goals in the mind . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1063. Relationships between perception and action . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107

3.1. Linkage by structure: the ideomotor approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1073.2. Linkage by process: the attention approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1093.3. Toward an integrative perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110

4. A selective review . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1114.1. Target-directed actions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111

4.1.1. The role of prediction during target-directed eye movements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1114.1.2. Transsaccadic learning and object continuity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112

4.2. Goal-directed actions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1124.2.1. Effect-based control of goal-directed eye movements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1134.2.2. Temporal and spatial visual attention for perception and action . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

5. Summary and conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

∗ Correspondence to: Department of Psychology, Bielefeld University, P.O. Box 100131,E-mail address: [email protected]

ttp://dx.doi.org/10.1016/j.neubiorev.2015.02.013149-7634/© 2015 Elsevier Ltd. All rights reserved.

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1 behavi

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tance because it perfectly mirrors the different treatments of theelusive term “goal” in the attention and ideomotor framework.Accordingly, goals might differ in that they are located within or

1 The idea that actions are driven by goals-in-the-mind is not only deeply embed-ded in commonsense psychology but also part of scientific theories of motivationand volition (e.g., Ach, 1910; Heckhausen, 1991; Miller et al., 1960, but see theecological approach of Turvey, 2013, for an opposite standpoint). Interestingly, theshift from goals-in-the-world to goals-in-the-mind can be traced and observed atdifferent levels of analysis. On a macro-level, the shift from behavior controlledby goals-in-the-world to behavior controlled by goals-in-the-mind can be consid-ered as an important evolutionary change from lower (e.g., amphibians) to highervertebrates (e.g., primates) (Neumann, 1990). That is, with the growth of the neo-cortex and the associated ability to represent the environment internally, primatestremendously escalated their room for maneuver. As a consequence, internal rep-resentations allowed decoupling behavior from the immediate stimulus situationleading to a remarkable flexibility of behavior (Goschke, 2013). On a meso-level, asimilar shift from the external to the internal world has taken place in the history ofpsychology. When cognitivism started to reject behaviorism, which solely focused

06 A. Herwig / Neuroscience and Bio

. Introduction

The emergence of nervous systems can be considered a mile-tone of evolution. Thanks to increasing cell differentiation animateeings have continuously extended their capabilities up to theoint that they are even able to systematically gain new knowledgef themselves in general and their nervous systems in particular.owever, despite providing humans with an exceptional machin-ry to think and reason one should not forget that the primaryunction of nervous systems is to control the body and to expressctive movements. This idea, namely that nervous systems are forction, is neatly illustrated by the life cycle of the sea squirt, aarine animal which actively moves around the ocean until it finds

suitable rock and implants itself in place. The first thing the seaquirt does after implanting on the rock, which it never leaves,s to recycle its cerebral ganglion (Dennett, 1991; Glenberg et al.,007). Thus, without the need for moving there seems to be noeed for maintaining a luxury brain. While there is admittedly a

ong way from the sea squirt to human cognition, it is self-evidenthat actions can be considered an essential aspect of our waking lifeecause they are the only way we have to affect our surroundings.

Over the past two decades, increased interest in action andts relationship with cognitive functions like perception, atten-ion, memory and volition has surfaced in parallel in a number ofndependent approaches. Today, a trend toward action as a gen-ine research topic is reflected in many different fields of inquiry,

ncluding motor control, cognitive neuroscience, experimental andognitive psychology, sport science, and developmental psychol-gy (e.g., Engel et al., 2013; Morsella et al., 2009; Prinz et al., 2013).iverse as all these different approaches contributing to action sci-nce may be, they share the basic functional belief that evolutionas optimized cognitive systems to serve the demands of actionLlinás, 2001). The mind and brain is for them much more a deviceor smart action than for true cognition. Accordingly, they claimhat cognition needs to be studied with respect to action and actioneeds to be studied with respect to its cognitive underpinnings.hile most recent approaches consent to such a general claim, the

uestion as to how exactly different functional domains such aserception and action are linked accompanied scientific debatesn action ever since (for a brief historical treatment see Herwigt al., 2013).

In this review article I will resume the question as to how per-eption and action are linked. More precisely, I will reconcile twoeemingly alternative approaches that have been treated more oress separately so far. As will be elaborated below, both approachesriginated from different research traditions and actions undertudy and emphasize different linkage principles of perception andction. For example, the ideomotor framework, which was one ofhe first theoretical approaches to address the cognitive underpin-ings of voluntary action, assumes that perception and action are

inked by common representational structures. That is, actions aressumed to be stored and retrieved by their perceivable reaffer-nces, to the effect that the shared representation (i.e., the commonode) for both perception and action is grounded in the sensoryomain (Hommel et al., 2001; Prinz, 1997). In contrast, the atten-ion framework assumes that perception and action are linked byommon attentional processes. That is, basic issues of attention likeelectivity, competition and priority control are assumed to per-ain to both perception and action, to the effect that targets forerception and targets for action are selected and prioritized byne and the same mechanism (Schneider, 1995; Schneider et al.,013).

However, before exploring different linkage principles of per-eption and action in further detail, in the next section, I will brieflyrovide some working definitions of three basic concepts that aresed throughout this article, namely, actions, targets and goals.

oral Reviews 52 (2015) 105–116

2. From targets in the world to goals in the mind

As has often been pointed out, actions are hard to individu-ate because they are embedded in a continuous stream of ongoingbehavior (e.g., Stränger and Hommel, 1996; Prinz et al., 2009). Inprinciple, this stream can be segmented at various levels and timescales ranging from brief muscle contractions and limb movements,over short-term interactions with external objects to long-termendeavors like taking a trip around the world. To deal with all thesediverse forms, actions are individuated in a continuous stream ofongoing behavior in terms of their underlying goals. That is, anaction starts with the first behavioral activity directed toward aparticular goal and terminates with the achievement of that goal.As a consequence, an action goes beyond a simple body movementbecause it is composed of two elements: the first is ongoing behav-ioral activity; the second is the orientation of these movementstoward a prospective goal state (Herwig et al., 2013).

While this first working definition of action provides a crite-rion for action segmentation and individuation, there is now asecond question that needs to be addressed. What is a goal? Likeactions, the term “goal” is used at various levels (cf. Prinz, 2008).First, we use the term goal at a descriptive level to characterizethe behavior of systems whose activities are directed toward theachievement of certain goal states or goals-in-the-world. For exam-ple, endurance runners, pigeons, and air conditioners all behave ina certain way (e.g., running, flying, regulating) to reach such goalstates (e.g., finish line, dovecote, temperature). On top of such adescriptive characterization of actions as movements directed atgoals-in-the-world, the term goal is also used at an explanatorylevel to define actions as movements driven by goals-in-the-mind.1

While goals-in-the-world are typically described from the thirdperson’s perspective, goals-in-the-mind are somewhat ambiguous.They are used for both, to describe the personal experience asso-ciated with the access to own action goals from a first person’sperspective, and second, to provide a subpersonal account of rep-resentational mechanisms involved in action control from a thirdperson’s perspective (Maasen et al., 2003). To avoid some of thisambiguity, I will thus use the term goal hereafter solely from a thirdperson’s perspective excluding issues of personal experience andintrospection from further analysis.

Besides distinguishing between goals-in-the-world and goals-in-the-mind, goals can be further categorized with respect to therequirements placed on action planning and execution (Prinz,2008). One of such possible categorizations is of particular impor-

on external events, action began to be construed as a function of internal mentalrepresentations (Miller et al., 1960). Finally, on a micro-level, the shift reflects theworkings of the mind and brain in expressing active movements. That is, accordingto the ideomotor framework goals-in-the-world need to be internally representedand anticipated for action selection and control (Herwig, 2014; Hommel et al., 2001).

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A. Herwig / Neuroscience and Bio

eyond the current range of objects or things to which the actoras perceptual access (Herwig et al., 2013). Goals within the per-eptual range are often referred to as targets. Such targets, can bepecified in terms of spatial parameters (e.g., location, shape, size,tc.) and are objects or things which can be looked at, pointed at orhot at and which therefore can be hit (or sometimes missed). Theyre typically investigated in research on attention and motor con-rol using, for example, aiming movements of the eyes (e.g., Deubelnd Schneider, 1996) or hands (e.g., Jeannerod, 1988; Rosenbaum,980). Importantly, target representations can be based on the cur-ent perceptual input in most instances. In contrast, realizing goalseyond the actor’s current perceptual range typically necessitates

situational change. Dressing, for example, first requires open-ng the wardrobe door to see, take out, and finally put on one’savorite shirt. Accordingly, goal representations involved in theseinds of actions have to be based on anticipations or predictions ofuture states that are not yet fully specified in the current perceptualnvironment. As will be elaborated in the next section, such antic-patory goal representations play a crucial role in the ideomotorramework.

To conclude, actions are characterized by their goal-irectedness. That is, they are defined as behavioral activityirected toward a certain goal. Such goals can be categorized atifferent levels including the distinction between goals-in-the-orld and goals-in-the-mind as well as the distinction between

oals within (i.e., targets) or beyond the perceptual range. Withhese preliminaries in mind, let us now turn to different linkagerinciples of perception and action.

. Relationships between perception and action

Linkages between perception and action have been studied in number of cognitive domains. Thus, one should not be surprisedo find different answers to the question as to how the input andhe output side of behavioral and cognitive activity are related toach other. In the following, I will focus on two approaches thatave come up with different linkage principles, namely, linkagey structure (i.e., the ideomotor approach) and linkage by processi.e., the attention approach).2 Finally, the last paragraph of thisection will strive toward an integrative perspective by reconcilinghe ideomotor approach with the attention approach. As such, thisaragraph will also provide the ground for a selective review ofesearch at the interface of both approaches.

.1. Linkage by structure: the ideomotor approach

Contemporary ideomotor approaches assume that linkageetween perception and action originates from shared represen-ational resources or structures (Hommel et al., 2001; Prinz, 1997).deomotor ideas can be traced back to early theoretical work inhe nineteenth century by James (1890/1981), Lotze (1852), andarleß (1861, translated by Pfister and Janczyk, 2012). In a nutshell,

deomotor reasoning assumes that actions are represented throughheir perceivable effects. Thus, although the sensory and the motoromain are still thought to be incommensurate at first, they becomeommensurate through the translation of motor entries into a sen-ory language (Prinz, 1984). Consequently, action and perception
ay induce or interfere with each other by virtue of representa-
ional overlap, or similarity (Kornblum et al., 1990; Prinz, 1990,997).

2 For a discussion of early approaches suggesting a linkage of perception andction by translation of motor codes into sensory codes, see Prinz (1997) and Herwigt al. (2013).

oral Reviews 52 (2015) 105–116 107

In the past two decades, the ideomotor approach has gained alot of empirical support that can be roughly clustered into threemain issues (Hommel and Elsner, 2009). These issues pertain tothree key questions raised by the ideomotor approach, namely: (1)how is action knowledge represented, (2) how is action knowledgeacquired, and (3) how is this knowledge used in action selection andcontrol. Action knowledge in this context incorporates knowledgeabout goals, movements and the relationship between goals andmovements. Moreover, action knowledge can incorporate declar-ative knowledge about action, which is often considered explicitand conscious, as well as procedural knowledge for action, whichis often considered implicit and non-conscious (Prinz, 2014; Schackand Mechsner, 2006).

Concerning the issue of representation, the ideomotor approachassumes that action knowledge and perception are generated andmaintained in the same representational medium (common codingprinciple, Prinz, 1990, 1997). As illustrated in Fig. 1 and speci-fied in detail by the theory of event coding (TEC, Hommel et al.,2001), sensory information from different modalities is thought toconverge on more abstract feature codes referring to distal infor-mation in a common coding system. These abstract feature codesare directly linked to codes belonging to different motor systems.As a consequence, perception and action may induce or interferewith each other by virtue of representational overlap, or similarity.This idea is supported by a number of studies. On the one hand,perception has been shown to affect action. For example, observingactions of others can induce own movements, as it is evident fromresearch on imitation and ideomotor movements (e.g., Knuf et al.,2001; Meltzoff and Prinz, 2002), or interfere with the production ofplanned actions (Brass et al., 2001). On the other hand, action hasbeen shown to affect perception. That is, action planning and pro-duction can either boost (assimilation effect, e.g., Craighero et al.,1999; Symes et al., 2008) or hamper (contrast effect, e.g., Müsselerand Hommel, 1997; Zwickel et al., 2008) the perception of stimu-lus features which are similar to action features (for a recent modeladdressing assimilation and contrast effect, see Thomaschke et al.,2012).3

Concerning the second issue of acquisition, a couple of studiesusing serial- and choice reaction time tasks have shown that theacquisition of action knowledge occurs relatively fast (i.e., afterroughly half an hour) and incidentally (i.e., without an explicitintention to learn). Moreover, knowledge about the relationshipbetween goals and movements is aggregated in the form of bidi-rectional association (see Fig. 2, upper part) of actions and theirensuing perceptual effects (e.g., Elsner and Hommel, 2001; Herwigand Waszak, 2012; Ziessler and Nattkemper, 2001). The acquisitionof such action–effect associations seems to follow similar princi-ples as classical or operant conditioning. For example, action–effectassociations can generalize to other feature-overlapping events(e.g., Beckers et al., 2002; Hommel et al., 2003). Depending onthe learning situation they can however also be acquired context-specifically (Kiesel and Hoffmann, 2004). Moreover, like classicalforms of learning the acquisition of action knowledge is affectedby the temporal contiguity and the probabilistic contingency ofmovement and effect (Elsner and Hommel, 2004). All these find-

3 The planning and control model of Thomaschke et al. (2012) assumes that differ-ent directions of motorvisual priming effects (i.e., assimilation vs. contrast effects)are due to the experimental tasks recruiting different representational structures.More precisely, assimilation effects are due to the activation of spatial represen-tations in movement control, whereas contrast effects are due to the binding ofcategorical representations in action plans.

108 A. Herwig / Neuroscience and Biobehavioral Reviews 52 (2015) 105–116

Fig. 1. Representation of action knowledge and perception in a common coding system. Sensory information is picked up by different sensory systems and converges ona to coa proxi

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e also acquired through observational learning or even solely bynstruction, see Paulus et al., 2011; Pfister et al., 2014).

Finally, the third key question of the ideomotor frameworks how action knowledge is used in action selection and control.ere, two different aspects pertaining to the issue of usage ofction–effect associations can be contrasted (see Fig. 2, lower part).irst, associations allow selection of appropriate movements tochieve an outcome given anticipatory effect representations. Thisrocess is a form of backward computation, because it starts fromesired goal states in the future and acts back to states requiredreceding them. As a consequence, actions can be triggered by acti-ating effect representations either endogenously (e.g., thinking
f the effect) or exogenously (e.g., perceiving the effect). Second,hey allow prediction of the outcome of given movements. Thisrocess is a form of forward computation, because it starts from
given state and acts forward to states expected to happen in the

ig. 2. Key issues of the ideomotor approach pertain to the acquisition and usage of actffect (A) results in the formation of bidirectional action–effect associations (B). Such asrediction of the outcome of given movements (D).

des belonging to different motor systems. Feature codes refer to distal informationmal information from the sensory surface or muscle innervation patterns.

future. Consequently, behavior can be optimized by comparing thepredicted action effects to the actual sensory consequences of theaction. Both aspects have been investigated in a number of studies.Concerning backward computation, it has been shown that actionsare indeed selected through the anticipation of the effects theseactions produce. For example, studies using the response–effectcompatibility paradigm indicate that actions are initiated fasterif actions and effects share certain features on a physical dimen-sion (Kunde, 2001, 2003; Kunde et al., 2011). Using eye trackingmethodology, we recently showed that effect anticipations notonly affect when an action is executed but also how it is executed(Herwig and Horstmann, 2011). Because in all these studies action
effects only appear after action execution any observed influenceson action can only be due to the anticipation of perceptual changesassociated with action execution. Moreover, recent studies usingfunctional brain imaging indicate that effect anticipations exist in
ion knowledge. A movement that is consistently followed by a particular sensorysociations can be used for action selection through effect anticipation (C) and for

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concrete, perceptual-like format as assumed by the ideomotorpproach (Kühn and Brass, 2010; Kühn et al., 2010). Concerningorward computation, it has been repeatedly shown that predictedction effects are processed differently than unpredicted actionffects. For example, predicted effects are perceived to occur earlyintentional binding, see Haggard et al., 2002) and less intensivesensory attenuation, see Blakemore et al., 1999; Weiss et al., 2011)han stimuli that are not resulting from own actions. Moreover,nanticipated effects have been shown to elicit error-related pro-esses and orienting responses making the given event available toehavioral control (Band et al., 2009; Waszak and Herwig, 2007).

To sum up, the ideomotor framework assumes that percep-ion and action are linked by common representational structureshich are grounded in the sensory domain. To date, the majority

f ideomotor studies addressing issues of representation, acqui-ition, and usage of action knowledge focus on relatively simplend discrete actions like pressing a key in a reaction time exper-ment (for few exceptions see, Mechsner et al., 2001; Schack and

echsner, 2006). Moreover, there is also a strong tradition in usingistal action effects that fall beyond the current range to which thector has perceptual access (for few exceptions see, Huestegge andreutzfeldt, 2012; Walter and Rieger, 2012). While selective focus

s always justified in science – perhaps even a prerequisite to suc-essful scientific inquiry – such a strategy may also carry the risk toisregard a whole category of important actions (e.g., oculomotorctions) and to lose sight of other prospering research fields (e.g.,ttention research).

.2. Linkage by process: the attention approach

Somewhat complementary to the linkage-by-structurepproach, the attention approach assumes that perception andction are linked by common attentional processes (Gottlieb, 2007;eubel and Schneider, 2005; Schneider, 1995, 2013; Schneider andeubel, 2002; Schneider et al., 2013). Evidently, both, the numberf objects that can be perceived as well as the number of actionshat can be performed at any given moment in time are limited.hus, selection processes are not only required for perceptionut also for action (Allport, 1987; Neumann, 1987; Schneider,995).4 The biased competition approach of attention (Desimonend Duncan, 1995) provides a framework for investigating suchelection processes. Accordingly, events compete for capacityimited neural representation and control over behavior. Thisompetition is thought to be biased by task-driven top-down fac-ors (e.g., relevance) and stimulus-driven bottom-up factors (e.g.,alience). Concerning space-based features, it has been suggestedhat top-down and bottom-up factors are combined on a commonriority map coding attentional priorities in a location specificanner (Bisley and Goldberg, 2010; Fecteau and Munoz, 2006).In recent years, attentional processes have turned out to

e a promising candidate to link different functional domainsSchneider et al., 2013). For example, there is now a substantialnd increasing body of experimental work investigating atten-ional processes across functional domains like perception andisual working memory (VWM) or perception and action. More
recisely, it has been suggested that VWM may simply be visualttention oriented to internal representations or visual attentionustained internally over time (Chun, 2011; Gazzaley and Nobre,
4 Historically, the question, what the functions of visual attention are, has ofteneen neglected in theories of visual attention. Among the first advocates of the ideahat attention mainly serves action (i.e., selection-for-action) were Allport (1987)nd Neumann (1987). Other work mainly focused on the function of visual attentionor perception (Goebel, 1993; LaBerge and Brown, 1989; Olshausen et al., 1993).chneider (1995), however, was the first to integrate both functions (i.e., selection-or-action and selection-for-perception) in his visual attention model (VAM).

oral Reviews 52 (2015) 105–116 109

2012; Kiyonaga and Egner, 2013). In fact, there is now a bunchof evidence indicating that VWM and visual attention are closelyinterrelated. Attention serves as a kind of gatekeeper by biasingthe encoding into VWM (Bundesen, 1990; Bundesen et al., 2005;Schneider, 2013). Conversely, memorized items in VWM can influ-ence the deployment of attention toward matching items in thevisual field (Awh et al., 2006; Olivers et al., 2006; Soto et al., 2005).Moreover, it has been repeatedly proposed that visual attention canplay a functional role in the active maintenance of information inVWM (Awh et al., 1998; Awh and Jonides, 2001; Postle et al., 2004).

Concerning the functional domains of perception and action,the linkage-by-process approach is further supported by stud-ies investigating the close coupling between selection processesfor perception and target-directed actions, i.e., actions which aredirected at spatial targets within the perceptual range of the actor(Deubel and Schneider, 1996, 2005; Schneider and Deubel, 2002).These studies were mainly inspired by the visual attention model(VAM) proposed by Schneider (1995) which postulates that selec-tion for perception and selection for target-directed actions areperformed by one common visual attention mechanism. In a seriesof experiments Deubel and Schneider demonstrated that saccadiceye movements, manual pointing as well as grasping movementshave to be preceded by the allocation of covert visual atten-tion to the target location. More precisely, perceptual processingwas selectively enhanced at the target location prior to move-ment execution. This finding indicates that biasing competitionfor target-directed actions and biasing competition for perceptionare strongly coupled to one common target (for an extension ofthis finding to multi-step actions, see Baldauf and Deubel, 2010;Deubel and Schneider, 2005). The linkage of attention and actionhas been further addressed within the premotor theory of atten-tion advocated by Rizzolatti et al. (1987). According to this theory,planning of a target-directed action is necessary and sufficient forshifting visual attention. That is, the premotor theory assumes thatvisual attention follows from the programming of target-directedactions (for diverging evidence, however, see Smith and Schenk,2012; Smith et al., 2014), whereas the VAM assumes that visualattention is a prerequisite for target-directed actions.

Fig. 3 depicts the linkage-by-process approach in an admit-tedly simplifying sketch that may however be suited to capturethe main ideas relevant in the present context. Accordingly, visualobject information is weighted by the bottom-up distinctivenessof an object as well as by the top-down relevance of an objecton a topographical map of space. This map is termed attentionalpriority map and provides for each object location attentional pri-orities that are used to bias competition for perception as well as fortarget-directed actions. In recent years, the firing patterns of singleneurons recorded in non-human primates suggest that brain areasof the oculomotor and attention system (e.g., frontal eye field [FEF],lateral intraparietal area [LIP], superior culliculus [SC] and pulv-inar) actually represent attentional priorities, that is the combinedrepresentation of an object’s bottom-up distinctiveness and its rel-evance (Bisley and Goldberg, 2010; Krauzlis et al., 2004; Robinsonand Petersen, 1992; Schall and Thompson, 1999). However, to dateit is not entirely clear how these various areas are concerted to forma common functional priority map.

In summary, the attention approach assumes that perceptionand action are linked by common attentional processes. By mainlyfocusing on actions like grasping, pointing or saccadic eye move-ments it deliberately restricts itself to a subset of actions that aredirected toward space-based targets within the actor’s perceptualrange. In comparison to discrete actions which are directed at goals
beyond the actor’s perceptual range (as mainly investigated by theideomotor approach) this selective focus on space allows a muchmore fine grained analysis by using, for example, continuous inputand output variables. Moreover, the restriction to target-directed

110 A. Herwig / Neuroscience and Biobehavioral Reviews 52 (2015) 105–116

Fig. 3. A simplified sketch of the linkage-by-process approach of perception and action. Task-driven top-down factors and stimulus-driven bottom-up factors are assumed tobe integrated (solid arrows) on a common priority map specialized for attentional control. Attentional control signals on the basis of these priorities are used (dotted arrows)t samet

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o bias competition in visual feature maps thus affecting perception. Moreover, theoward the prioritized location.

ctions provides the opportunity to link current models to findingsathered in single unit recordings in non-human primates. How-ver, at the same time selectively focusing on one specific subset ofctions may once again carry the risk to disregard other impor-ant actions (e.g., actions to produce effects outside the currenterceptual range). Likewise, focusing solely on actions directed atargets within the perceptual range may conceal the importance oflooking into the future”, namely the notion of prediction and antic-pation which forms an integral part of the linkage-by-structurepproach.

.3. Toward an integrative perspective

As should have become clear in this brief sketch of possible link-ge principles of perception and action, the question as to how thenput and the output side of behavioral and cognitive activity areelated to each other is treated differently by diverse approacheshat exist more or less independently of each other. As always incience, there are many ways to deal with such diversity. On thene hand, one may single out one particular approach withoutiving much credit to other ones. Such views are often justifiedy the claim that the approach chosen is the proper one and thatther approaches address different and independent levels of anal-sis that bear no causal significance for understanding the issuender study. While ideomotor and attention approaches certainlyddress different levels of analysis, it is hard to see why one par-icular approach should play no functional role at all. On the otherand, one may consider two or more approaches simultaneouslyy explicitly addressing the issue of how these approaches mighte related to each other. This second suggestion has been partly fol-

owed up previously. For example, Hommel (2011) briefly toucheshe relationship between ideomotor approaches and the premo-or theory of attention in the context of the Simon effect (Simonnd Rudell, 1967). Moreover, Hommel and colleagues (2001) havepeculated that common representations of perception and actionay be mediated through some kind of attentional mechanismhich they termed “intentional-weighting” (see also Memelink

nd Hommel, 2013). Finally, based on findings that action plan-
ing can prime action relevant perceptual dimensions (e.g., Fagiolit al., 2007) and also bias selection in visual search (e.g., Wykowskat al., 2009), Hommel (2010) recently suggested that attention cane considered as a direct derivative of action control. Together these
attentional control signals are used to bias competition for target-directed actions

previous attempts from the ideomotor perspective are importantfirst steps to decrease the gap between both approaches.

However, to promote a thorough elaboration of how bothapproaches are related, two further ideas are worth following. First,the relationship of the ideomotor and the attention approach needsto be studied from both directions, i.e., not only from the ideomo-tor but also from the attentional perspective. Second, the classicalrestriction of research approach and goals under study (i.e., targetswithin vs. goals beyond the perceptual range) needs to be over-come. Therefore, the first direction needs to study target-directedactions, aiming at demonstrating the workings of representationaloverlap of perception and action. Accordingly, central ideomotorconcepts like anticipation, prediction, acquisition and represen-tation will be applied to actions that are directed to space-basedtargets within the actor’s perceptual range. Conversely, the seconddirection will focus on goal-directed actions, aiming at demon-strating the workings of common attentional processes linkingperception and action. Accordingly, central attentional conceptslike selectivity, competition and priority control will be applied toactions which are directed at goals beyond the actor’s perceptualrange.

As a first step to move toward an integrative perspective, thepresent article will by and large focus special emphasis on eyemovements. These movements are, with an average of three tofour fast ballistic steps (i.e., saccades) per second, one of the mostfrequent primate movements under voluntary control (Land andTatler, 2009; for a recent review see Schütz et al., 2011). Impor-tantly, eye movements can be considered a fundamental action aswell as perceptual device (Schneider et al., 2013) and are thus aprime candidate to approach the linkage of perception and action(Huestegge and Koch, 2010; Wolff, 1984). Given their commonmeasurement as proxy of covert visual attention they are also aprime candidate to bridge the gap between the ideomotor and theattention approach.

As a second step to move toward an integrative perspective,the present article will investigate the relationship of the ideomo-tor and the attention approach from both directions as suggestedabove (for an overview see Fig. 4). First, central concepts of the ideo-
motor approach will be applied to target-directed eye movements.As will be elaborated in the next section, the gathered findingspertaining to this line of research strongly suggest that the rep-resentation, acquisition and usage of action–knowledge follow the

A. Herwig / Neuroscience and Biobehavioral Reviews 52 (2015) 105–116 111

Fig. 4. Classification scheme for the present review. Publications are arrangedaccording to their treatment within the ideomotor (linkage-by-structure) or atten-tion (linkage-by-process) approach as well as according to their treatment of goalswithin or beyond the actor’s perceptual range. Gray quadrants depict classical com-b

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Fig. 5. (A) Due to the non-homogeneity of the visual system, pre- and postsaccadicinformation of the saccade target object differ in acuity. (B) The two-phase model oftranssaccadic feature prediction. At Phase 1, pre- and postsaccadic information of thesaccade target object is associated. At Phase 2, these associations are used to predictthe foveal appearance of peripheral objects (peripheral object recognition) and theperipheral appearance of foveal search templates (visual search) from Herwig and

inations of research approach and goals under study.

ame principles for target-directed and goal-directed actions. Thus,lso actions directed at space-based targets within the actor’s per-eptual range – which constitute an essential part of our wakingife – can be construed in the ideomotor framework as goal-directedctions selected and guided by a prediction of their perceptual con-equences. Second, basic concepts of the attention approach wille applied to actions that are directed at goals outside the actor’serceptual range including eye movements leading to distal per-eptual effects and eye movements to memorized object locations.he main findings pertaining to this second line of research indi-ate that selecting information for goal-directed actions as wells for perception are performed by one common visual attentionechanism.

. A selective review

.1. Target-directed actions

Most of the studies conducted in the ideomotor framework soar focus special emphasis on actions directed at goals which falleyond the current range to which the actor has perceptual access.or example, there is a strong tradition in using simple key pressctions which produce auditory or visual action effects in the dis-al environment (for recent reviews see Herwig, 2014; Hommel,009; Nattkemper et al., 2010; Shin et al., 2010). While such aocus was particularly successful in demonstrating the workingf internally anticipated events in action control, it should be notorgotten that a serious part of our daily actions is directed at tar-ets within the perceptual range to the effect that these actionsan be partly based on the current perceptual input. Recently,he question as to whether the ideomotor framework can be alsopplied to target-directed actions has gained increased interest forxample in the field of tool-use actions (e.g., Herwig and Massen,009; Kunde et al., 2012; Massen and Prinz, 2007; Müsseler et al.,008), temporal synchronization (Walter and Rieger, 2012) andye movement control (Herwig and Schneider, 2014; Herwig et al.,n press; Weiß et al., 2014; Huestegge and Kreutzfeldt, 2012).n line with this renewed interest, in the following I will thus
ddress the question how the ideomotor approach can be applied toarget-directed actions by focusing special emphasis on eye move-
ents.

Schneider (2014).

Reprinted with permission from APA.

4.1.1. The role of prediction during target-directed eyemovements

Due to the visual systems non-homogeneity, objects can beprocessed in detail only within an astonishingly small region ofabout 2◦ around the center of gaze corresponding to foveal vision.As a consequence, we constantly move our eyes to assure that thehigh resolution foveal region is brought to interesting spots of thevisual field for detailed visual sampling (see Fig. 5a). Thus, seenfrom an ideomotor perspective, each target-directed eye move-ment is accompanied by a direct perceptual effect. That is, priorto a saccade, peripheral objects are only coarsely represented,whereas the same objects are represented with high acuity whenfoveated. We recently addressed exactly this direct perceptualeffect of saccadic eye movements and proposed a feature predic-tion mechanism based on past experience to deal with changesaccompanying saccadic eye movements (Herwig and Schneider,2014). More precisely, three questions pertaining to the ideomotorframework were investigated (see Fig. 5b). First, we tested whethernew action knowledge is acquired during saccadic eye movementsin the form of associations of pre- and postsaccadic information
of the saccade target object. Second, we asked whether predic-tions based on such transsaccadic associations affect perception,i.e., peripheral object recognition. Third, we investigated whether

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In three experiments, participants were first exposed to anltered visual stimulation where, unnoticed by most participants,ne object systematically changed spatial frequency during theaccade. This first acquisition phase was conducted to establishew and unfamiliar associations of peripheral and foveal object

nformation. To test whether acquired transsaccadic associationsffect perception, participants in Experiments 1 and 2 were thenequired to judge the spatial frequency of a peripheral saccadearget object in a second test phase. The results of these two exper-ments demonstrated that peripheral perception is biased towardreviously associated postsaccadic foveal input and that this effect

s particularly associated with making saccades. In particular, if thebject during acquisition intrasaccadically changed its frequencyrom low in the periphery to high in the fovea, this object’s fre-uency was later on judged to be higher in the periphery comparedo a baseline object that did not change frequency during acquisi-ion. Conversely, objects that previously changed frequency fromigh in the periphery to low in the fovea were later on judgedo be lower. This finding was recently replicated and extended tonother feature dimension, namely shape (Herwig et al., in press).hat is, targets were perceived as less curved for objects whichreviously changed from more circular in the periphery to moreriangular in the fovea. Likewise, shapes were perceived as moreurved for objects which previously changed from triangular to cir-ular. To test whether acquired transsaccadic associations affectction, Herwig and Schneider (2014) further presented partici-ants in Experiment 3 a foveal search template on each trial of theest phase and asked them to search and saccade to this target inhe periphery. Importantly, the spatial frequency of the peripheralarget object could physically either match or mismatch the fre-uency of the foveal search template. On the basis of the learningistory, frequency matches and mismatches could be both, congru-nt and incongruent with the experience during acquisition. Theesults of Experiment 3 revealed that search performance was bet-er for acquisition congruent combinations of peripheral and fovealbjects which demonstrate that saccades were biased toward pre-iously associated presaccadic peripheral input.

Together the findings are well in line with ideomotor theoryy highlighting the importance of past experience for an effect-ased control of eye movements. That is, during target-directed eyeovements new action knowledge linking pre- and postsaccadic

nformation is easily acquired. This knowledge can be used for twourposes: to predict how peripheral objects will look in the fovea,nd what foveal search templates should look like in the periphery.hus, action knowledge affects both, the perception of, as well ashe action toward peripheral objects which indicates a tight linkagef perception and action. Additionally, we suggested that apply-ng the ideomotor framework to target-directed eye movements

ight also contribute to the impression to see the visual field uni-ormly detailed (Dennett, 1991). In this sense, the prediction ofction effects (i.e., precise postsaccadic foveal information) prioro movement execution probably conceals acuity limitations in theeriphery.

.1.2. Transsaccadic learning and object continuityWhile Herwig and Schneider (2014) provided first evidence for

n effect based-control of target-directed eye movements, little isnown about the basic conditions driving the acquisition of actionnowledge for this particular class of actions. To shed light onhis question, Weiß et al. (2014) investigated the role of object
ontinuity in the acquisition of transsaccadic associations. Objectontinuity – the spatiotemporal stability of an object – has proveno be an important factor in establishing object persistence acrossrequently occurring interruptions of visual input in particularly
oral Reviews 52 (2015) 105–116

saccadic eye movements. For example, breaking temporal stabil-ity by inserting a temporal blank after the saccadic eye movementstrongly improves participants’ ability to detect displacements ofan object across a saccade (Deubel et al., 1996). Likewise, displace-ments are better detected following a spatial disruption of objectcontinuity (e.g., by a transsaccadic shape change, Demeyer et al.,2010).

Given this strong influence of spatiotemporal factors on theperceived object persistency across saccades, Weiß et al. (2014)addressed the question whether object continuity is also a nec-essary precondition to learn transsaccadic associations for laterfeature prediction. To test this question, the paradigm to assessfeature prediction in visual search (introduced in Herwig andSchneider, 2014) was adopted and a condition with object con-tinuity (Experiment 1a) was compared against conditions wherewe separately disrupted temporal (Experiment 1b) and spatialobject continuity (Experiment 1c) during learning. More precisely,temporal object continuity was disrupted by inserting a post-saccadic blank and spatial object continuity was disrupted by atask-irrelevant shape change. Although there were overt and covertindicators for better change detection under disruption, interest-ingly, visual search performance revealed that neither disruption oftemporal object continuity nor disruption of spatial object continu-ity impaired transsaccadic learning. Thus, the basic finding reportedin Herwig and Schneider (2014, Experiment 3) was replicated irre-spective of disrupting object continuity across saccades.

This main finding shows that associating peripheral input andfoveal input across saccades is a robust, default mode of thehuman visual system. Moreover, the independence of transsac-cadic learning of object-continuity testing implies a close linkbetween transsaccadic learning and the more general concept ofaction–effect learning (e.g., Elsner and Hommel, 2001; Herwiget al., 2007; Herwig and Waszak, 2009, 2012; Waszak and Herwig,2007). As elaborated in Section 3.2, studies on action–effect learn-ing are deeply rooted in the ideomotor framework and typicallyfocus on manual actions like key presses and auditory effects liketones. They address the more general question as to how par-ticipants learn that a particular action can produce a particularaction–effect. Importantly, comparable to the present finding per-taining to transsaccadic learning, studies on action–effect learningalso demonstrated that the disruption of temporal contiguity (upto a delay of 1000 ms between action and effect) does not pre-vent learning (Elsner and Hommel, 2004). Thus, the finding ofWeiß et al. (2014) provides first evidence that the acquisition ofaction-knowledge follows similar principles for target-directed andgoal-directed actions.

4.2. Goal-directed actions

To date, most of the studies conducted in the attention approachfocus on actions like grasping, pointing or saccadic eye movements– actions which are directed toward space-based targets within theactor’s perceptual range (e.g., Deubel and Schneider, 1996, 2005;Schneider and Deubel, 2002). While such a focus allows a muchmore fine grained analysis and the opportunity to link findingsacross species, it may carry the risk to disregard another impor-tant class of actions, namely actions which are executed to produceeffects outside the current perceptual range. As elaborated above,such goal-directed actions have been so far mainly investigated inthe ideomotor approach. Recently, however, there is also increas-ing interest in the attention field to incorporate goals outside theperceptual range. For example, studies addressing the relation-
ship between attention and VWM (e.g., Awh et al., 1998, 2006;Olivers et al., 2006; Soto et al., 2005) as well as between attentionand motor prediction (e.g., Hughes et al., 2013; Jones et al., 2013)have alluded to the intricate interplay of goal representations and

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ttention allocation. In accordance with this renewed interest inoals which cannot be solely based on the current perceptual input,he second set of publications reviewed in Section 4.2 will focus onoal-directed eye movements.

.2.1. Effect-based control of goal-directed eye movementsAlthough it is widely accepted that eye movements are not only

uided by the current visual input but also by the current task atand (Buswell, 1935; Land and Tatler, 2009; Yarbus, 1967), only

ew studies addressed the question as to how exactly goals outsidehe actor’s perceptual range control eye movements. In line withdeomotor reasoning, such distal goals might affect eye movementsn the same way as they affect other action modalities – that ishrough an anticipative mechanism. Admittedly, the idea that therere distal environmental effects of eye movements is not particu-arly intrusive at first glance. In fact, plenty of our eye movementsead to changes in sensation (see Herwig and Schneider, 2014;erwig et al., in press; Weiß et al., 2014) but are not accompaniedy changes in the environment. However, it has to be noted that we,s social beings, move our eyes not only to get information but alsoo supply information to our fellows. Thus, in social interaction gazeas a unique characteristic because it can cause changes in the envi-onment or to be more precise in our counterparts (e.g., empathicacial expressions, eyebrow flash, etc.). Accordingly, we recentlyypothesized that eye movements in social context should be con-idered as goal-directed actions (Herwig and Horstmann, 2011).

To investigate the effect-based control of goal-directed eyeovements, Herwig and Horstmann (2011) conducted two experi-ents in which participants in an acquisition phase had to saccade

o neutral faces which changed their facial expression (to happy orngry, depending on the position of the face) briefly after the gazeell upon them. Thus, the environmental effect of the eye movementas outside the participants’ perceptual range prior to movement

xecution. Nevertheless, participants’ saccades were directed dif-erently to the same neutral faces depending on the predictednvironmental effect. That is, the first saccade was directed moreften to the mouth region of a neutral face about to change into aappy one and to the eyebrows region of a neutral face about tohange into an angry expression (see Fig. 6). Measuring eye move-ents provided a proxy of covert visual attention because there is a

trong coupling of covert and overt forms of orienting briefly before saccade is initiated (Deubel and Schneider, 1996; Schneider andeubel, 2002). Thus, the present finding can be interpreted within

he attention approach as evidence for an influence of predictedction effects on attentional prioritization. Converging evidenceomes from a recent study by Khan et al. (2014). Here, stimuli thatonsistently appeared in the vicinity of the saccade target following

saccadic eye movement attracted the saccadic landing position inhe course of the experiment.

The study by Herwig and Horstmann (2011) revealed two fur-her relevant findings. First, in a following test phase, saccades in

Fig. 6. Differences in saccades’ vertical landing distribution on the neutral face

oral Reviews 52 (2015) 105–116 113

response to facial expressions were initiated more quickly to theposition where the expression was previously triggered. Thus, assuggested by the ideomotor approach, actions (here, goal-directedeye movements) can be triggered by activating effect represen-tations through the perception of previously experienced actioneffects (Elsner and Hommel, 2001). Second, the reported effectson attentional prioritization as well as on action priming criticallydepended on the action mode. That is, saccades’ landing positionsand saccadic latencies were only affected if participants duringacquisition freely selected where to look next (Experiment 1), butnot if saccades where triggered by external stimuli (Experiment2). This latter finding replicates previous results obtained with keypress actions and auditory effects (Herwig et al., 2007; Herwig andWaszak, 2009, 2012) which strongly indicates that the acquisitionand usage of action–effect associations follows similar principlesfor oculomotor and manual actions (see also Weiß et al., 2014, forconverging evidence).

4.2.2. Temporal and spatial visual attention for perception andaction

While most of the studies supporting the attention approachas a linkage principle of perception and action focused on target-directed actions, two recent studies also addressed the attentionapproach in the context of goal-directed eye movements (Herwiget al., 2010; Griffiths et al., 2013). These two studies focused on aparticular class of saccadic eye movements, namely memory-guidedsaccades, which are saccades directed toward memorized locations.As such, these eye movements are directed at goals outside theactor’s current perceptual range.

In two experiments, Griffiths et al. (2013) investigated whetherprocessing location information for a memory-guided saccade tem-porarily interferes with perception, i.e., pattern recognition. Moreprecisely, Griffiths and colleagues varied the temporal intervalbetween the peripheral presentation of action relevant locationinformation (T1) and a central presented letter (T2) that had tobe reported following the memory-guided saccade at the endof the trial. This experimental paradigm bears similarities toestablished paradigms that are frequently used to measure tem-poral visual attention like the attentional blink (Raymond et al.,1992) or dwell time paradigm (Duncan et al., 1994). Impor-tantly, however, Griffiths et al. (2013) focused on the processingof spatial information for goal-directed actions while in clas-sical attentional blink and dwell time paradigms T1 typicallyalso requires pattern recognition, i.e., a perceptual judgment.It was hypothesized that, if processing information for goal-directed actions and processing information for perception aresubject to a common temporal attention mechanism, as suggested
by the attention approach, processing of T1 should temporarilyinterfere with processing of T2. Moreover, comparable to the clas-sical attentional blink following pattern recognition we expectedthis interference to be pronounced for conditions in which T1
for the different effect conditions. From Herwig and Horstmann (2011).

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Both experiments revealed that T2 performance increased withnset asynchrony of both targets. In addition, Experiment 2,here T1 was followed by location distractors, showed a task-ependent interference effect. That is, interference was strongernd more durable under dual-task conditions when T1 needed toe processed for a memory-guided saccade. This finding demon-trates that limitations in temporal visual attention do also emergen an experimental paradigm, in which T1 requires the processingf action relevant location information, while T2 requires thetandard pattern recognition task. As such, the data indicatehat processing information for goal-directed eye movements androcessing information for perception seem to underlie a commonttentional mechanism. Thus, the attentional approach also coversnd accounts for linkages between perception and goal-directedctions in the domain of temporal visual attention.

Finally, the study by Herwig et al. (2010) addressed spatialisual attention as a candidate to link perception and action. Morepecifically, we conducted two experiments to investigate whetherhe perception of an onset distractor interferes with the mainte-ance of spatial information for a memory-guided saccade. Thisesearch question was motivated by recent studies suggesting

tight interdependence of visual attention and VWM. Accord-ng to the attention-based rehearsal hypothesis put forward bydward Awh, covert shifts of attention are thought to play a func-ional role in the active maintenance of information in VWMAwh et al., 1998). Given that onset stimuli are known to capturepatial attention, we hypothesized that maintaining spatial infor-ation for goal-directed actions should be subject to interference

y onset distractors presented in the vicinity of the memorizedocation.

We tested this hypothesis by adopting previous research on thelobal effect, a well-known spatial effect of distractor stimuli onhe oculomotor system, showing that visually guided saccades areypically directed to intermediate locations between a target and aimultaneously presented distractor stimulus (Coren and Hoenig,972; Findlay, 1982). Because the global effect is known to occurnly for distractors appearing at less than 20◦ of angular distancerom the target axis (Walker et al., 1997), Experiment 2 systemati-ally varied the distance between the memorized location and theistractor. The results of both experiments showed that a distractorhich was briefly flashed in the vicinity of the memorized locationuring the retention interval attracted the saccade’s landing posi-ion. Comparable to the classical global effect, this attraction wasnly found, if the distractor was presented within a sector of 20egrees around the target axis.

In accordance with the attention approach, the results of Herwigt al. (2010) indicate that perception and action recruit the samepatial selection mechanisms that can be probably ascribed tottentional priority maps (see Fig. 3). Moreover, the finding thathe “memory-based” global effect is subject to the same spatialonstrains as the classical global effect once again support the ideahat actions directed at targets within the actor’s perceptual rangend actions directed at goals outside the actor’s perceptual rangeollow similar principles of attentional prioritization. Accordingly,ttentional priority maps may be best characterized as the “neuro-omputational space” where memory, perception and action meet.

. Summary and conclusions

The present article aimed at reconciling different linkage prin-iples of perception and action as advanced by the ideomotornd attention approach. While ideomotor approaches assume thaterception and action are linked by common representational

oral Reviews 52 (2015) 105–116

structures, attentional approaches focus special emphasis on com-mon attentional processes as a linkage principle of perception andaction. To strive toward an integrative perspective, the presentarticle followed two basic strategies. First, eye movements, as aprime candidate to bridge the gap between the ideomotor and theattention approach, were prominently featured. Second, possiblerelationships of the ideomotor and attention approach were inves-tigated from two directions: by applying central concepts of theideomotor approach to target-directed actions typically addressedwithin the attention framework (Section 4.1), and by applying cen-tral concepts of the attention approach to goal-directed actionstypically addressed within the ideomotor framework (Section 4.2).

In a nutshell, the reviewed studies revealed two main out-comes. First, central concepts of the ideomotor approach includingthe representation, acquisition and usage of action-knowledgealso govern target-directed eye movements. Consequently, it isproposed that also target-directed actions are selected and guidedby a prediction of their perceptual consequences. Second, centralconcepts of the attention approach including selectivity, competi-tion and priority control also apply to goal-directed actions, i.e., eyemovements directed at goals outside the actor’s perceptual range.Consequently, it is proposed that selecting information for goal-directed actions as well as for perception is performed by commonvisual attention mechanisms.

Together, both main outcomes point to the conclusion that oper-ations addressed in both conceptual frameworks interact with eachother. This claim poses a challenge to the functional divide thatoften seems to exist between ideomotor and attention approaches.Pertaining to ideomotor approaches, it is thus important to developmore detailed views on the question how attentional processesaffect the way common representational structures are used forperception and action. Recent work suggests that progress alongthis line will critically depend on ways to connect the concept ofprediction and attention (see also Summerfield and Egner, 2009;Roussel et al., 2013). Likewise, for attention approaches, it is impor-tant to elaborate how representational structures affect the waycommon attentional processes are used for perception and action.As suggested by the results reviewed in Section 4.2, progress alongthis line will probably depend on ways to extend the currentlyspace-based concept of an attentional priority map (see Fecteauand Munoz, 2006) to visual features beyond space.

Acknowledgements

I am grateful to the research group ‘Competition and prior-ity control in mind and brain: new perspectives from task-drivenvision’ at the ‘Center for Interdisciplinary Research’ (ZiF) inBielefeld. The idea to reconcile the attention and ideomotorapproach originated during the great research year of the groupin 2012–2013 (see http://www.uni-bielefeld.de/%28en%29/ZIF/FG/2012Priority/). This work was supported by a grant from the Ger-man Research Council (Deutsche Forschungsgemeinschaft; DFG) toArvid Herwig and Werner Schneider (He6388/1-1).

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