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No one knows what attention is

No one knows what attention is In this article, we challenge the usefulness of “attention” as a unitary construct and/or neural system. We point out that the concept has too many meanings to justify a single term, and that “attention” is used to refer to both the explanandum (the set of phenomena in need of explanation) and the explanans (the set of processes doing the explaining). To illustrate these points, we focus our discussion on visual selective attention. It is argued that selectivity in processing has emerged through evolution as a design feature of a complex multi-channel sensorimotor system, which generates selective phenomena of “attention” as one of many by-products. Instead of the traditional analytic approach to attention, we suggest a synthetic approach that starts with well-understood mecha- nisms that do not need to be dedicated to attention, and yet account for the selectivity phenomena under investigation. We conclude that what would serve scientific progress best would be to drop the term “attention” as a label for a specific functional or neural system and instead focus on behaviorally relevant selection processes and the many systems that implement them. . . . . . . . . Keywords Attention Motor control Selection Sensorimotor Decision making Phylogenetic Intention Evolution Parietal cortex Superior colliculus Introduction however, that the overuse and popularity of this statement in cognitive research has been detrimental to progress – that in “Everyone knows what attention is” (James, 1890)isone of fact, no one knows what attention is. More specifically, we the most popular quotes from William James and certainly the argue that the concept of “attention” is one of the most mis- most famous statement about human attention. We argue, leading and misused terms in the cognitive sciences. In the present paper, we stake the position that the term “attention” Note that William James’ own approach to attention was mainly concerned should be abandoned and the nature of the research in this area with phenomenology, an aspect of attention that we do not further consider in be re-conceptualized to focus on the subsets of processes and this article given the modern functional and neural approaches we address mechanisms that lead to task-specific performance. Similar instead. positions have been proposed and discussed previously (see Anderson, 2011;Di Lollo, 2018; Hommel & Colzato, 2015; * Timothy N. Welsh t.welsh@utoronto.ca Krauzlis, Bollimunta, Arcizet, & Wang, 2014;Mole, 2011). The present paper reaffirms and expands this position by plac- Institute of Psychology, Cognitive Psychology Unit and Leiden ing particular and new emphasis on the interconnected and Institute for Brain and Cognition, Leiden University, Leiden, the integrative nature of the human sensorimotor information pro- Netherlands cessing systems. This emphasis on integrated sensori- Faculty of Kinesiology, Sport, and Recreation, University of Alberta, cognitive-motor processes takes inspiration from the synthetic Edmonton, Alberta, Canada approach to understanding “cognition” (Hommel & Colzato, Department of Neuroscience, University of Montreal, 2015) and a proposed phylogenetic refinement of the scientific Montreal, Quebec, Canada approach to understanding behavior (Cisek, 2019 [this issue]). School of Health and Human Performance, Dalhousie University, In the present paper, we start by discussing and outlining Halifax, Nova Scotia, Canada the central problem with the way “attention” has been concep- Department of Cognitive, Linguistic and Psychological Sciences, tualized and studied thus far. We make the case for adopting a Brown University, Providence, RI, USA synthetic approach to studying cognitive phenomena wherein Centre for Motor Control, Faculty of Kinesiology and Physical the focus is on the subset of processes and mechanisms that Education, University of Toronto, 55 Harbord Street, have been attributed to and investigated under the umbrella of Toronto, ON M5S 2W6, Canada Atten Percept Psychophys (2019) 81:2288–2303 2289 “attention,” rather than on “attention” as one overarching con- Before we explain our main position and arguments, cept. To bolster our analysis of the state of affairs, we present we would like to emphasize that the theoretical problems two test cases. In the first, we examine the debate about the that we highlight are particularly visible with respect to conceptual distinction between attention and intention, and the concept of attention, but by no means restricted to show that this debate fails to adequately account for the avail- that concept (Hommel, 2019a). For example, very similar able data. In the second, we review research on selection and arguments to those that we present in the following have reward history to show how conventional analytic approaches been put forward to question the concept of memory. to solving this problem are ineffective. As an alternative to the Decades of research on human memory have seen an analytic approach, we provide a brief review of the phyloge- ever-increasing number of memory systems that were netic evolution of the human brain (for an expanded account, thought to represent separable aspects of memory perfor- see Cisek, 2019 [this issue]) and show how selective attention mance, which then were thought to be explained by the emerged as just one necessary consequence of the challenges existence of corresponding memory systems, with rather facing animals behaving in the natural world. In the end, we limited contributions to a mechanistic understanding of conclude that the traditional analytic attempt to lump many the underlying processes (Bechtel, 2008). As recent con- diverse empirical observations under one common umbrella siderations suggest, however, the various types of mem- called “attention” and to try to explain all of them by referring ory may not at all reflect the operations of separable to one coherent attentional system has actually failed, and dedicated systems, but rather stand for different should be replaced by a more synthetic approach. This syn- byproducts of normally functioning cognitive systems thetic approach focuses on, and starts with, ecologically rele- (Buckner & Schacter, 2004), and emerged at different vant mechanisms and processes and then tries to account for times during the evolution of our species (Murray, as many phenomena (“attentional” or not) as possible. Wise, & Graham, 2017). Similar arguments have been put forward for the concept of emotion (Barrett, 2017; Hommel, 2019b) and may be developed for other con- cepts as well, including “cognition” itself (Cisek, 2019). The concept of attention We focus here on attention because we believe that at least some of the related phenomena are best understood We are not the first to raise concerns about problems with in terms of the kinds of interactions between sensory, the term “attention.” Multiple authors have highlighted motor, and cognitive phenomena that are the focus of the tendency to reify attention, creating circular explana- this special issue. tions for empirical results (Anderson, 2011;DiLollo, Theorizing about human attention suffers from at least 2018). Another common criticism is that multiple process- three main problems. First, the concept of attention invites es underlie what is typically labeled as “attention” (Di misconceptions of one coherent set of cognitive or neural Lollo, 2018; Hommel & Colzato, 2015). Mole (2011) operations, depending on one’s level of analysis, that all highlights that James’ statement came at the time where contribute to what we call “attention” (e.g., Kahneman, there was debate among theorists as to whether the main 1973). Second, the concept of “attention” can also easily role of attention was in thinking, perceiving, or acting. be misunderstood as both an important explanandum that James’ contemporary, F.H. Bradley, produced one of the psychology is rightly expected to explain and the earliest criticisms of the concept of attention, titling his explanans that is supposed to form the explanation – thus essay “Is There Any Special Activity of Attention.” In rendering the latter a pseudo-explanation. And, third, the brief, his position was that there were too many examples concept is thought to distinguish a particular set of cog- of phenomena labeled “attention”, with little concern nitive or neural operations from other, seemingly different about the processes underlying such phenomena sets of operations, such as those related to decisions, in- (Bradley, 1886). Now, over 130 years later, there is a tentions, motivations, emotions, and, of particular rele- continued and heightened need to question the role of vance to the present special issue, action planning and “attention”, the use of the term “attention,” and a search execution. As we show in the expanded discussion of for what “attention” is (e.g., Busemeyer, Gluth, these three points in the following paragraphs, all these Rieskamp, & Turner, 2019; Gottlieb, 2012). We reaffirm assumptions are incorrect. these positions and further suggest that compartmentaliz- First, let us start by considering which phenomena re- ing “attention” and then searching for the “attentional searchers have, historically speaking, been trying to explain system” hinders the development of a comprehensive un- when using the term “attention.” According to some tradition- derstanding of human behavior because it ignores inte- al and conventional views, “attention” is the set of cognitive/ grated, parallel, and reciprocal relationships among senso- neural mechanisms responsible for maximizing the efficient ry, cognitive, and action processes. utilization of our limited capacities to process, store, and 2290 Atten Percept Psychophys (2019) 81:2288–2303 retrieve information. However, consultancy of introductory mechanistic understanding of how the observed phenomena textbooks (e.g., Eysenck & Keane, 2000) and the internet are causally produced (Krauzlis et al., 2014). reveals a dramatic variety of abilities attributed to attention: Third, research on attention has followed, and suffered the ability to select external events for further internal process- from, the common analytical approach to psychological func- ing (focused attention); ignore misleading information and/or tioning (see Hommel & Colzato, 2015,for amore detailed an irrelevant location (selective attention); process irrelevant discussion of this issue). The analytic approach comprises a information (involuntary attention); selectively integrate in- search for an exhaustive definition (which, given the diversity formation belonging to one event within and across sensory of the subfunctions of attention, is impossible), the identifica- modalities (feature integration); prioritize processing of events tion of assumed subfunctions (e.g., overt vs. covert, early vs. from a particular location (spatial attention); systematically late, focused vs. divided, voluntary vs. automatic attention, search for a target event (visual search); perform multiple etc.) with separable functional and neural processes, and the tasks at the same time (divided attention); control the spatial concentration of research on tasks and subfunctions rather parameters of eye movements (selective attention for action); than actual processes. The problem of this analytic approach prioritize one goal over others (goal-centered attention); pri- is that it underestimates and overlooks commonalities be- oritize one object, memory item, or conscious representation tween subfunctions and, in a wider perspective, commonali- over others (object-centered attention); and consolidate infor- ties with other concepts. For instance, the very fact that we use mation for later use and concentrate in anticipation of a pos- concepts like attention, decision making, intention, emotion, sible event over some time (sustained attention). At face value, and motivation in different situations and theoretical contexts it seems highly unlikely that the same set of functional/neural by no means implies that the underlying functional and neural mechanisms are involved in, and responsible for, this broad processes are different and separable. Indeed, attempts to sys- variety of phenomena (Allport, 1993), and a bulk of behav- tematically distinguish the processes “underlying” attention ioral and neural evidence confirms that most subfunctions can from the processes that do not, often fail to produce any co- be dissociated from each other (e.g., Fan, McCandliss, herent consensus. In the following section, we outline one Sommer, Raz, & Posner, 2002). Accordingly, it is unsurpris- exemplar case of the failure of the analytic approach – the ing that no theory has been suggested so far that comes even attempt to separate “attention” from “intention.” close to providing a coherent account of all phenomena sailing under the label of “attention.” Second, the term “attention” is often used to capture both Attention versus intention: A failed the problem and the solution of cognitive processing; i.e., to dichotomy describe both the phenomenon one aims to explain and the mechanism proposed to provide the explanation. For instance, Most researchers agree that the posterior parietal cortex repre- the term attention is used to refer to the consequences of both sents the core of the neural substrate of selective attention, and “voluntary” and “involuntary” factors in favoring the repre- is a key node of an “attentional network” (Corbetta & sentation of one event having a stronger impact on decision Shulman, 2011; Posner & Dehaene, 1994; Ptak, 2012). In making and action than representations of other events (e.g., particular, individual neurons in the posterior parietal cortex Yantis, 1998). But the concept of attention is also used to refer appear to reflect the locus of attention (Bisley & Goldberg, to the system, mechanism, or ability to deal with (or avoid) the 2010; Robinson, Goldberg, & Stanton, 1978) and parietal consequences of such unequal potencies of representations to damage often leads to phenomena of spatial neglect drive behavior (e.g., Broadbent, 1958). Along the same lines, (Bartolomeo, 2007; Corbetta & Shulman, 2011). However, a attention is considered by some to represent the critical capac- separate line of research implicates these same regions of the ity limitation, the cognitive bottleneck that needs to be brain in processes related to movement control (Mountcastle, accounted for and explained (e.g., Pashler & Johnston, Lynch, Georgopoulos, Sakata, & Acuna, 1975; Snyder, 1998), while still others consider attention to be the cognitive Batista, & Andersen, 1997). In particular, regions of the pos- means to deal with such bottlenecks (e.g., Bundesen, 1990). terior parietal cortex are strongly and reciprocally intercon- These conceptual confusions have created a situation in which nected with parts of the frontal lobe that are involved in the it is no longer clear what the to-be-explained problem actually planning and guidance of movement (Johnson, Ferraina, is (do we have a cognitive bottleneck that we need to make the Bianchi, & Caminiti, 1996;Markov etal., 2014), individual best of, or do we have too much information we need to neurons are strongly modulated by the type of action per- choose from?), and whether attention is a concept that refers formed with respect to identical stimuli (Cui & Andersen, to the problem or to the solution. This runs into the danger that 2007; Snyder et al., 1997), and inactivation of the posterior research and theorizing on attention is based on circular rea- parietal cortex causes biases in free-choice tasks soning (attentional phenomena are explained by assuming and (Christopoulos, Kagan, & Andersen, 2018), but not decisions pointing to attentional systems) rather than on a deeper based on visual evidence (Katz, Yates, Pillow, & Huk, 2016). Atten Percept Psychophys (2019) 81:2288–2303 2291 These apparently contradictory findings have fueled a heat- 1996; Gersch, Kowler, & Dosher, 2004;Hoffman & ed and persistent debate, now in its fifth decade, on whether Subramaniam, 1995; Peterson, Kramer, & Irwin, 2004). the posterior parietal cortex is involved in guiding “attention” These studies demonstrate there is preferential processing of or whether it reflects the individual’s “intention.” As is typical stimuli at the goal of a saccade just before the onset of the eye for the dominant analytical approach to psychological science, movement (presumably because “attention” has been shifted researchers have tried to resolve this debate by defining the to the goal location). concepts of attention and intention in ways that make them Other research has extended the study of these action- appear mutually exclusive: “Attention” is what restricts the attention interactions to manual actions, showing that plan- inflow of sensory information to cognition, what enters con- ning and performing reaching and grasping movements prior- scious thought for further processing, whereas “intention” is itizes the processing of the target objects of these movements the output of cognition, the will (free or otherwise) to perform (e.g., Pratt & Abrams, 1994; Rizzolati, Riggio, & Sheliga, a specific action. Defined in this way, the two appear like 1994; Tipper, Lortie, & Baylis, 1992; see also Wu, 2014). distinct concepts that must be dissociable through careful ex- Even when the eyes remain fixated, perceptual discrimination perimental design. And yet, after decades of work by some of is better at the to-be-reached goal than non-goal locations the world’s most accomplished neuroscientists, a clear disso- (Baldauf & Deubel, 2008; Baldauf, Wolf, & Deubel, 2006; ciation of the function of posterior parietal cortex remains Deubel & Schneider, 2003; Deubel, Schneider, & Paprotta, elusive. A prominent review expressed this frustration many 1998; Khan, Song, & McPeek, 2011). The “attentional im- years ago, suggesting that “current hypotheses concerning pa- pact” or prioritized processing associated with intended future rietal function may not be the actual dimensions along which movements goes beyond mere spatial prioritization because the parietal lobes are functionally organized; on this view, other studies have shown that moving or planning to move what we are lacking is a conceptual advance that leads us to also facilitates the detection of action-related features of the test better hypotheses” (Culham & Kanwisher, 2001). object targeted by the movement. For example, preparing for a To escape this rather uncomfortable state of affairs, some grasping movement facilitates the detection of size oddballs, researchers have argued for a more integrative view on atten- while preparing for a pointing movement facilitates the detec- tion and intention. A particularly promising approach is the tion of location oddballs (Fagioli, Hommel, & Schubotz, 2007; pre-motor theory, which argues that shifts of attention are see also Craighero, Fadiga, Rizzolatti, & Umiltà, 1999). Other triggered by sub-threshold saccadic commands in oculomotor studies (Bekkering & Neggers, 2002; Moher, Anderson, & areas and, conversely, shifts of attention in space lead to action Song, 2015; Tipper, Meegan, & Howard, 2002; Weir et al., planning (e.g., Rizzolatti, Riggio, Dascola, & Umilta, 1987). 2003; Welsh & Pratt, 2008; Welsh & Zbinden, 2009;see also Support for the former idea has come from a large number of Gallivan, Barton, Chapman, Wolpert, & Flanagan, 2015; observations including: (a) behavioral studies showing that Glazebrook, Welsh, & Tremblay, 2016; Yoxon, Constable, & attention and eye movements are strongly linked behaviorally Welsh, 2019 [this issue]) show that the processing of specific (e.g., Deubel & Schneider, 1996; Kowler, Anderson, Dosher, object features can be prioritized depending on the relative (i.e., &Blaser, 1995; Sheliga, Riggio, & Rizzolatti, 1995); (b) task/action-specific) salience of those features for the to-be- fMRI studies of visual attention showing activation in eye- performed action. That is, the same feature (e.g., orientation) movement areas for attention tasks (e.g., Beauchamp, Petit, can be prioritized in one action context (e.g., grasping), but not Ellmore, Ingeholm, & Haxby, 2001; Corbetta et al., 1998; another action context (e.g., pointing). Hence, it is neither phys- Nobre, Sebestyen, & Miniussi, 2000)and for movement acti- ical stimulus properties nor action goals alone that generate vation (decoding) in retinotopically defined visual cortex dur- selectivity, but rather selectivity is shaped by the reciprocal ing movement tasks (Gallivan, Chapman, Gale, Flanagan, & and iterative interactions between these factors. These findings Culham, 2019); (c) stimulation studies showing that activation thus suggest that multiple functional and neural systems are of neurons in the superior colliculus (SC), frontal eye field involved in selective attention. (FEF), and lateral intraparietal area (LIP) can change the focus In addition to the interactive nature of stimulus properties of attention (Cavanaugh & Wurtz, 2004;Cutrell&Marrocco, and action goals in determining selection and prioritization of 2002; Moore & Fallah, 2001; Muller, Philiastides, & locations and features, it does not seem that selection stops Newsome, 2005); and (d) neurological studies of patients with solely within any putative attentional system. Indeed, neural attentional disorders following damage to the frontal cortex, activity related to multiple simultaneously active intentions to parietal cortex, or midbrain (e.g., Husain & Kennard, 1996; act at potential target locations, as well as the selection of the Posner, Cohen, & Rafal, 1982; Posner, Rafal, Choate, & final target, has been identified in various structures more Vaughan, 1985; Sapir, Soroker, Berger, & Henik, 1999). Of commonly associated with the planning and execution of ac- particular relevance to the present purpose are behavioral stud- tions, such as the dorsal premotor area, the parietal reach re- ies revealing that perceptual discrimination at the goal location gion, and the motor cortex (Cisek & Kalaska, 2005; Klaes, of an upcoming saccade is improved (Deubel & Schneider, Westendorff, Chakrabarti, & Gail, 2011; Pesaran, Nelson, & 2292 Atten Percept Psychophys (2019) 81:2288–2303 Andersen, 2008; Scherberger & Andersen, 2007;Song& selection. Given that this would make it no longer apparent McPeek, 2010; Thura & Cisek, 2014). Behaviorally, the pres- that such a reduced version of “attention” has anything to do ence of multiple co-existing response representations and the with other “attentional” functions like integration, orientation, dynamic selection of target from non-target stimuli and ac- or vigilance, this would eventually call for dropping the con- tions is also expressed through the spatiotemporal character- cept – and the same argument holds for “intention.” On the istics of reaching and grasping movements. Specifically, in- positive side, this would prevent researchers from trying to stead of the efficient straight and direct movements that one find commonalities in processes and substrates that are unlike- might anticipate if attentional selection had been completed ly to be found. On the negative side, however, there is no prior to the intention to act, the trajectories of hand and eye theoretical justification to pick just these functional aspects movements veer towards or away from non-target stimuli de- but not others. What looks like a definitional issue thus be- pending on the timing and salience of the non-target stimuli comes a theoretical bias that is lacking justification. (e.g., Chapman et al., 2010; Gallivan & Chapman, 2014; Alternatively, one might search for hybrid approaches that Howard & Tipper, 1997;Moheretal., 2015; Neyedli & allow for additional components and factors. A typical ap- Welsh, 2012; Song & Nakayama, 2006, 2008; Welsh, 2011; proach of this sort was the resource theory of attention, which Welsh & Elliott, 2004; Wispinski, Gallivan, & Chapman, in triggered heated debates in the 1970s and 1980s (Kahneman, press). Thus, the characteristics of the physically executed 1973; Navon, 1984). While the first approaches were simple action actually reflect the “attentional” state of the target and and elegant by assuming one kind of resource that needs to be non-target stimuli. Collectively, these data indicate that atten- distributed over all mental work, the attempt to integrate an tion, selection, and intention are not readily separated in a set increasing number of unpredicted findings led to the invention of discrete serial processes, but are more dynamic and contin- of increasing numbers and types of separate resources. In the uous in nature and embedded within a densely interconnected, end, this made systematic predictions impossible (Navon, parallel processing system. 1984), which is the main reason why this approach no longer While more work needs to be done to synthesize these plays an important role – except in the field of ego-depletion, neural and behavioral observations into a coherent framework, where history seems to repeat itself (Friese, Loschelder, it seems clear (to us) that the conceptual distinction between Gieseler, Frankenbach, & Inzlicht, in press). The main reason attention and intention is not sufficient to account for the va- why hybrid approaches that simply lump together different riety of findings discussed here. The distinction fails to pro- factors are not overly successful rests in the fact that the re- vide a meaningful contribution or framework for sorting the spective factors are not truly integrated into a coherent available findings into useful categories to stimulate further framework. theorizing, and it also clashes with the demonstration of so A similar tendency can be seen with respect to selective attention, where Awh, Belopolsky, and Theeuwes (2012)have many interactions between input processing and output gen- eration. But what is the solution to this and the many other tried to integrate findings that are no longer consistent with the conceptual problems we are encountering in thinking about historical distinction between endogenous attention, which human attention (e.g., controlled vs. automatic processing; represents the prioritized processing of stimuli to which the facilitation vs. inhibition, etc.)? agent “wants” to attend, and exogenous attention, which rep- resents the prioritized processing of stimuli that are unrelated to the present action and goals. The history of distinguishing A failed analytic solution: Selection between endogenous and exogenous attention is very similar and reward history to the distinction between attention and intention. Each started out by trying to improve definitions about what the concept As noted above, the dominant analytical approach to psycho- referred to, only to be faced later with the inability to system- logical functioning begins with an exhaustive search for a atically sort the available findings into two distinct categories. definition of a concept, including the borders of where it dif- In a nutshell, endogenous attention is sometimes too automat- fers from other concepts. So, to understand “attention,” one ic and exogenous attention is sometimes too dependent on the would tend to first define how it differs from “intention,” current prioritized stimulus feature or action goal to make this “decision-making,”“motivation,” etc. In view of a failure of dichotomy fruitful and tenable (Awh et al., 2012;Folk, this approach, as is obvious for the case of attention versus Remington, & Johnston, 1992;Hommel & Wiers, 2017). intention, two reactions are to be expected. First, one might Awh et al. suggest solving this problem by adding a third consider the previous attempts to define attention and inten- variable – selection history – to the list of factors. In particular, tion as flawed and try to improve the definitions by further the idea is that goals (the factor responsible for endogenous reducing the conceptual overlap between the two concepts. attention), salience (the factor responsible for exogenous at- For instance, one may further reduce the concept of attention tention), and selection history (a factor that does not seem to fit to mere input selection and the concept of intention to output the previous dichotomy and is associated with previous Atten Percept Psychophys (2019) 81:2288–2303 2293 selections and rewards generated by the selections) all con- Indeed, prioritizing shape and orientation when grasping ob- tribute to selectivity by sending their output to an integrative jects makes more sense than prioritizing color because those priority map. Although this approach may account for many features are more likely to determine a successful or an unsuc- of the available findings, we are not convinced that it really cessful grasp. Hence, establishing a bias for shape and orien- solves the problem, but rather provides a patch that holds tation over color when grasping would be a functional adap- concepts together and, in the end, prevents or misdirects the tation. However, this influence implies that selection history search for suitable solutions. Instead, we suggest that a com- affects how goals impact (endogenous) attention. Along the plete dismantling of the concept of attention is required. same lines, the relative salience of the visual dimension At first there does not seem to be anything wrong with the changes substantially during the first years of life (e.g., idea that structures, or a singular structure, in the human cen- Suchman & Trabasso, 1966), which at least opens the possi- tral nervous system are devoted to collecting and integrating bility that selection history impacts salience. information that affects prioritized processing. One candidate While these arguments are fully consistent with Awh structure is the superior colliculus, which is thought to reflect a et al.’s suggestion to consider selection history as a third factor priority map of stimuli in the visual field (Fecteau & Munoz, involved in attentional control, they also imply that the 2006). Importantly, and as required from the view that selec- resulting three factors are not independent but strongly over- tivity for behavioral relevance is the purview of the entire lapping and intertwined – both empirically and conceptually. moving body, the superior colliculus is involved not just in As we have tried to explain, these conceptual-overlap prob- eye movements, but in orienting movements of the eye, head, lems are unlikely to be resolved by more definitions. Rather, body, and hand (Gandhi & Katnani, 2011; Stuphorn, what is needed is a theory that not only assumes that integra- Hoffmann, & Miller, 1999). Emphasizing this point, Song tion takes place but that explains how that integration works. and colleagues (Song, Rafal, & McPeek, 2011; Song & Another reason why just adding selection history as an McPeek, 2015) found that the superior colliculus plays a caus- additional factor raises more questions than answers is that al role in target selection during manual reaching tasks, the concept itself is unclear, particularly in its overlap with supporting the idea that the superior colliculus is part of a other related factors beyond exogenous and endogenous con- general-purpose target selection/orientation system trol. One such factor that is intertwined with selection history (Nummela & Krauzlis, 2010; Song et al., 2011). On the other is reward history. It is uncontroversial that previously hand, there is no need to assume that the superior colliculus is rewarded stimuli receive preferential processing (Anderson, the only map that integrates relevant information to steer at- Laurent, & Yantis, 2011; Anderson & Yantis, 2012), suggest- tention, nor is it necessary to assume that all available infor- ing that reward history is important in determining salience. mation is integrated into that one map. As we argue below, the Conventionally, stimuli must have been selected in order for human brain can be considered to have many sources of se- the organism to have received a reward, thus conflating the lectivity and, in the end, it is the brain as a whole that does the two concepts. Awh et al. (2012) appear to acknowledge this integration. Given that this integration is the explanandum tension, considering both previous reward and previous stim- (the to-be-explained phenomenon), postulating the existence ulus selections to be exemplars of “selection history” while of one map that has no other function than achieving this facing the fact that selection history and reward history cannot integration seems to be one more attempt to “explain” apsy- be identical (given that previous selections might not have chological phenomenon by positing the existence of a dedi- received reward). However, the fit of reward history into this cated system whose only purpose is to somehow create that joint category is much less obvious than this theoretical inte- phenomenon. gration suggests since reward history is itself likely composed Apart from this more general meta-theoretical problem, of many differentiable factors. adding one more factor to a model that just assumes that inte- For instance, the preference for rewarding stimuli is stron- gration takes place without explaining how that can be done is ger when the larger expected gain of the stimulus is due to an unlikely to guide further research. In the case of Awh et al. increased probability of receiving a reward than when there is (2012), one reason is that selection history overlaps consider- a lower cost of failure (Neyedli & Welsh, 2015b). This finding ably with goal-induced endogenous selectivity and salience- suggests that what participants take to be rewarding is itself induced exogenous selectivity. For instance, the fact that plan- multifaceted – not just the magnitude of the reward, but also ning and carrying out particular kinds of actions systematical- how likely it is that they will receive the reward. Furthermore, ly facilitates the processing of particular object features (e.g., across a series of reach-decision experiments, a multitude of of size and orientation for grasping, location for reaching: biasing factors have been observed including: reward value Bekkering & Neggers, 2002; Craighero et al., 1999; Fagioli and probability (Chapman, Gallivan, & Enns, 2015a), the best et al., 2007) is unlikely to be genetically determined, but rather option in a decision set (Wispinski, Truong, Handy, & the consequence of learning and experience of selecting dif- Chapman, 2017), current level of accumulated wealth ferent features for grasping over the lifespan (Hommel, 2010). (Neyedli & Welsh, 2015a), the number of targets and not the 2294 Atten Percept Psychophys (2019) 81:2288–2303 perception of them (Milne et al., 2013), and how the number A synthetic approach of choice-options is represented (Chapman et al., 2014). The problem here is similar to the one Awh et al. (2012)tried to A synthetic approach is valuable only insofar as it synthesizes solve: the definition of the concept under investigation is too elements that actually correspond to real biological processes restricted. In their case, endogenous and exogenous control at both neural and functional levels, and it is a significant were insufficient to account for the variety of pheomenena challenge to figure out what those processes are. One power- being ascribed to the concept of attention, so they added se- ful strategy for doing so – for keeping our synthesis close to lection history. Selection history is itself decomposable into biological reality – is to use evolution as a guide. This guid- (at least) selection and reward history, and reward history is ance can be done through a procedure of “phylogenetic refine- itself decomposable even further. Thus, the nature and the ment,” whereby one progressively elaborates a theory about influence of reward is itself dependent on numerous contex- how neural and behavioral processes evolved along a given tual factors and the expression of the confluence of these fac- lineage, always respecting constraints about the neural modi- tors is not easily captured in a unitary construct. fications and behavioral adaptations that appeared at each The picture gets even more complicated when one con- stage (see Cisek, 2019 [this issue]). For this reason, here we siders studies comparing the impact of rewards of equal step away from the concept of “attention” and take a brief magnitude but in opposite directions (e.g., positive/gain detour into the history of how the relevant neural circuits vs. negative/loss). For example, loss aversion, made fa- evolved in the lineage that leads to homo sapiens (humans). mous by the work of Kahneman and Tversky (1979), While it is often very difficult to know why a given modifica- shows that people treat potential losses as being more aver- tion took place in evolution, establishing what was the se- sive than equivalent gains are rewarding. Interestingly, quence of modifications can be constrained by a wealth of when decisions between positively and negatively reward- comparative and developmental data, leading to strong and ing stimuli are made rapidly (Chapman et al., 2015b), testable hypotheses about how neural circuits and behavioral asymmetries in choice behavior are also observed, but go abilities evolved together. opposite to loss aversion. That is, in these situations, par- The evolutionary history of spatial interaction along the ticipants appear to be disproportionately drawn toward op- primate lineage is a long and complex tale (Fig. 1). A major tions giving gains, while the aversive impact of loss-related advance occurred during the Cambrian epoch, over 500 mil- choices is attenuated. These findings highlight an addition- lion years ago (Mya), with the elaboration of visually guided al complication – that different biases are likely to operate orientation behaviors. Our simple chordate ancestors pos- on different timelines. For instance, in the study of the sessed a visual escape circuit that involved projections from asymmetryindecisionstogo for agoodoptionversus a single photosensitive patch in the rostral neural tube to a avoid a bad one, participants were biased toward positively midbrain structure called the tectum, which projected to the valanced targets 100 ms earlier than they were biased to spinal cord to generate locomotion (Lacalli, 1996, 2018). In move away from negative ones (Chapman et al., 2015b). the lineage leading to vertebrates, the photosensitive patch This finding echoes related work showing that more time is split into two lateral eye patches on both sides of the head required to select optimally between visuomotor choices (Butler, 2000). Because these eye patches projected contra- when they differ based on negative value information com- laterally to the tectum, which projected ipsilaterally to the pared to when they differ based on the probability of re- spinal cord, the circuit caused our ancestors to turn away from ward (Neyedli & Welsh, 2015b). salient visual stimuli such as the shadow from an approaching Taken together, these and other findings strongly suggest predator (Fig. 2a). As the eye patches expanded, they folded that selection history is tightly interconnected with reward into cups and formed a lens (Lamb, 2013), resulting in a two- history and that neither of these concepts are particularly well dimensional retina that provided a topographic mapping of understood. Hence, adding selection history to the two other external stimuli. The tectum expanded in parallel, with a not well understood concepts of exogenous and endogenous matched topographic map of space in its superficial layers attention is unlikely to help much in understanding the mech- and gradients of downstream projections in its deep layers. anisms underlying human selective attention. To be clear, we The result was an “action map” of oriented escape responses are not advocating an alternative theory at this point, but rather to threatening stimuli at specific locations in the external we suggest an alternative theoretical perspective: Let us re- world. place the analytical approach, which seeks to explain complex Microstimulation studies reveal the presence of an orga- phenomena by first carefully defining them and then nized map of oriented escape responses in the tectum of lam- subdividing them into simpler elements, with a synthetic ap- prey (Saitoh, Menard, & Grillner, 2007), a jawless fish whose proach that considers how simple mechanisms and functional ancestors diverged from ours about 550 Mya. These studies processes, each of which is itself behaviorally relevant, can also reveal the presence of another action map, which lies together give rise to complex phenomena. within the rostral region of the tectum. This map is sensitive Atten Percept Psychophys (2019) 81:2288–2303 2295 Fig. 1 A reduced phylogenetic tree of bilaterally symmetric animals, (paleobiodb.org). Small rectangles indicate the estimated latest timing exclusively emphasizing the lineage that leads to humans. Branch of innovations described in the boxes. Note that many branch points points represent some of the divergences between different lineages, and lineages are omitted for clarity. Silhouettes along the right are from with timing estimated on the basis of molecular clock analyses (Erwin phylopic.org et al., 2011). Thick lines indicate the presence of relevant fossil data to space in front of the animal, and projects mostly contra- accomplished through lateral inhibition that produces “win- laterally to the spinal cord, thereby producing orientation and ner-take-all” dynamics (Grossberg, 1973; Mysore & approach actions (Jones, Grillner, & Robertson, 2009; Knudsen, 2011; Wang, 2002). Kardamakis, Saitoh, & Grillner, 2015). It is this latter tectal How is this related to attention? A few sentences after that sub-circuit that is most relevant to attention and selection. In famous phrase we quoted above, James wrote that attention the avoidance circuit, multiple stimuli can engage multiple “implies a withdrawal from some things in order to deal ef- escape actions that can simply be averaged downstream to fectively with others.” That withdrawal from some stimuli to produce adaptive avoidance behavior (Fig. 2b). In contrast, interact with another stimulus is indeed accomplished, quite averaging cannot work in an approach circuit, because the literally, within the approach circuit of the rostral tectum. And average response to two stimuli will cause the animal to miss while these simple circuits for governing interactive behavior both of them (Fig. 2c). Consequently, the approach circuit may seem far removed from the higher cognition of humans, must select between actions, such that one completely sup- they are indeed the precursors to the mechanisms that control presses the other. This kind of selection could be what has been called “selective attention.” The tectum is 2296 Atten Percept Psychophys (2019) 81:2288–2303 Fig. 2 Circuits for avoidance and approach in a hypothetical early stimulus. (b) Spatial averaging of escape directions (numbered arrows) vertebrate. (a) In the avoidance circuit, visual information from the away from two threatening stimuli (black stars) is an effective response. lateral eyes arrives in the contralateral tectum, which projects (c) For approach actions, spatial averaging is maladaptive, making ipsilaterally to the midbrain locomotor regions. Thus, if a stimulus falls winner-take-all dynamics necessary. B and C reused with permission on the left eye, the locomotion will tend to turn to the right until from Cisek (2019) stimulation is balanced and the body is oriented away from the homologous to the human superior colliculus, which, as and each projected to a specific set of relevant effectors. In a discussed earlier, is strongly implicated in both orienting gaze sense, each circuit was an “action map” analogous to the much through eye and head movements and in controlling covert older tectal systems for approach and avoidance, but guiding attention when gaze is stationary (Basso & May, 2017). the much wider repertoire of task-specific interactions avail- Though much has developed in the central nervous system able in the mammalian niche. and the world since our lineage diverged from lamprey in As the behavioral repertoire of mammals continued to ex- the early Cambrian, both the approach and avoidance circuits pand, so did the dorsomedial neocortex, and there was a dif- of the tectum are still present in fish (Herrero, Rodriguez, ferentiation and specialization of action-specific maps of sen- Salas, & Torres, 1998) and in mammals (Comoli et al., 2012). sory space. In primates, expansion of the parietal cortex was Eventually, our ancestors left the seas and some of them, particularly dramatic, yielding a variety of idiosyncratic rep- the amniotes, adapted to a fully terrestrial lifestyle. This adap- resentations of space particular to the needs of different action tation was accompanied by an expansion and lamination of types (Andersen, Snyder, Bradley, & Xing, 1997; Stein, 1992) the telencephalic pallium, an integrative olfactory, visual, and (see Fig. 3). For example, visually guided reaching actions somatosensory region that would eventually give rise to the involve the medial intraparietal cortex (Cui & Andersen, cerebral cortex (Aboitiz & Montiel, 2015;Striedter, 2005). In 2007;Kalaska &Crammond, 1995), which represents targets all mammals, the neocortex consists of two sheets (Finlay & within reach with respect to the direction of gaze and the Uchiyama, 2015), a dorsomedial sector that is spatially topo- position of the hand (Buneo, Jarvis, Batista, & Andersen, graphic and a ventrolateral sector that is non-topographic. In 2002; Gallivan, Cavina-Pratesi, & Culham, 2009), and is in- primates, the former includes a medial and dorsolateral pre- terconnected with frontal regions controlling reaching, such as frontal cortex, cingulate regions, all of the premotor, motor, the dorsal premotor cortex (Johnson et al., 1996;Wise, sensorimotor, and parietal cortex, as well as the retrosplenial Boussaoud, Johnson, & Caminiti, 1997). Grasp control in- cortex. The latter includes parts of the lateral prefrontal cortex, volves the anterior intraparietal area (Baumann, Fluet, & orbitofrontal cortex, and all of the limbic cortex and the tem- Scherberger, 2009), which is sensitive to object shape and is poral lobe. Most relevant to the issue of attention is the interconnected with grasp-related frontal regions such as the dorsomedial sector of the neocortex, which is organized into ventral premotor cortex (Nakamura et al., 2001;Rizzolatti & a set of fronto-parietal circuits dedicated to different classes of Luppino, 2001). The control of gaze involves the lateral species-typical actions (Graziano, 2016; Kaas & Stepniewska, intraparietal area (Snyder, Batista, & Andersen, 2000), which 2016). In early mammals (300 Mya), this system was proba- represents space in a retinotopic frame (Colby & Duhamel, bly quite limited and consisted simply of medial circuits con- 1996; Snyder, Grieve, Brotchie, & Andersen, 1998), and is cerned with locomotion and lateral circuits concerned with interconnected with frontal regions controlling gaze, such as head and mouth movements (Kaas, 2017). Each of these cir- the frontal eye fields and the superior colliculus (Paré & cuits processed sensory information in an idiosyncratic man- Wurtz, 2001) – taking advantage of the tectal orientation/ ner specialized for its specific type of action (e.g., space near approach system that has been steering animals since the the legs for locomotion, space near the snout for ingestion) Cambrian epoch. Atten Percept Psychophys (2019) 81:2288–2303 2297 Fig. 3 The primate cerebral cortex contains a set of parallel sensorimotor relevant given the current behavioral context. AIP anterior intraparietal streams in the dorsomedial regions (blue arrows), each involved in a area, FEF frontal eye fields, IT inferotemporal cortex, LIP lateral specific type of action using specific representations of space. All of intraparietal area, LPFC lateral prefrontal cortex, MIP medial these use information on object identity and outcome value, computed intraparietal area, OFC orbitofrontal cortex, PMd dorsal premotor in the ventrolateral regions (red arrows), to select the actions most cortex, PMv ventral premotor cortex, V1 primary visual cortex In many situations, different fronto-parietal action streams goals and avoid negative outcomes. The phylogenetic per- will compete against each other. For example, one must make spective suggests that the primate posterior parietal cortex is an all-or-none decision as to whether to burrow at the roots of part of a topographically organized dorsomedial neocortical one tree or instead walk over to another tree. In other situa- system for visually guided interactions oriented with respect tions, however, different fronto-parietal streams will be coor- to objects in the world (Cisek, 2007). This system is organized dinated. For example, when a head/snout orientation system as parallel sensorimotor streams, each contributing to a spe- points at a target, that target is then made available to other cific type of action within the animal’s behavioral repertoire, behaviors, such as burrowing or biting. This availability be- whose activity is orchestrated through selective invigoration, comes particularly important in primates, which evolved from energization, or drive from the basal ganglia and other struc- tree-climbing insect eaters and developed large eyes with a tures (Cisek & Thura, 2018; Grillner, Robertson, & central, high-resolution fovea, and acquired a taste for fruit. Stephenson-Jones, 2013). Within each of these fronto- In such animals, the system for controlling the orientation of parietal action streams, target selection occurs through gaze takes on an executive role for many other visually guided winner-take-all dynamics taking place in an idiosyncratic spa- behaviors. Selecting a target for gaze becomes part of tial reference frame specific to each given type of action (e.g., selecting what to reach for or which branch to grasp to climb. retinotopic for eye movements). One of those streams is con- It comes to serve much of the role traditionally ascribed to cerned with orienting gaze through eye and head movements, “selective attention.” Indeed, it has long been proposed that and appears to have an executive role simply because so many selective attention, both overt and covert, is closely related to of the other streams rely on high-resolution visual information the gaze orientation system and involves the same neural that is derived from the fovea. When an animal (including structures (Corbetta et al., 1998; Rizzolatti et al., 1987), in- humans) is placed in a laboratory situation and trained/ cluding the posterior parietal cortex, the frontal eye fields, and instructed to perform just one isolated aspect of complex nat- the superior colliculus. ural behavior, what the researcher will observe in this region is And so, this brief foray into the long journey of primate activity that appears to be related to what the researcher has evolution has brought us back to selective attention and to the defined as “attention,”“intention,” or “decision-making,” de- posterior parietal cortex, but with a different perspective on pending on the particular task variables that are being exper- both. The question is not whether the posterior parietal cortex imentally manipulated. But it does not follow from these cor- plays a role in something called “attention” or something else relations that there exists anything in the brain that can be called “intention,” but how the posterior parietal cortex fits meaningfully delineated as an “attentional system” (or, for that within a broader system that enables animals to select and matter, an “intention system” or “decision system”). The key control interactions with their environment to achieve their insight is that the posterior parietal cortex is not part of an 2298 Atten Percept Psychophys (2019) 81:2288–2303 “attention system” but, rather, that selective attention phenom- In an analytic approach to science, one runs the risk of ena are part of what the posterior parietal cortex produces as it becoming a slave to the concepts that have been generated. goes about its business of controlling goal-directed action. Many researchers have taken terms like “attention,”“inten- tion,” and “decision making” from everyday language and expect this linguistic categorization to somehow map to iden- Summary and conclusions tifiable mechanisms in the brain or functions. Of course, when one starts to peer into actual neural functions, there is no clear We thus conclude that selectivity emerged through evolution delineation, only a set of processes that interact to create se- as a design feature to enable efficient goal-directed action. lectivity in the end. These processes interact not because they Such selectivity became necessary as the action repertoire of belong to a dedicated system, but because the human brain the given line of organisms that led to humans increased. This and body evolved this way and selectivity was a necessary means that selectivity is an emerging property arising from a feature to achieve efficient behavior. Further, everything an myriad underlying processes, and the simple fact that humans individual does throughout their life (distant and recent past) (and other species showing selective attention) evolved the creates, reinforces, and shapes selection: Turning to the left way they did, with selective attention being one of many makes us ignore stimuli on the right, picking one apple makes byproducts, next to “selective intention” and “selective deci- us overlook the others, saying one word prevents us from sion making.” Here, we have primarily emphasized selection uttering any other. And each of the different selections results mechanisms in the superior colliculus and parietal cortex, but in all ranges of rewards, from positive gains to negative losses. similar arguments can be made for other selection Selection and reward are thus inherent ingredients of all our mechanisms in other brain regions. For example, Krauzlis lives and the way we lead them (Allport, 1987). et al. (2014) suggest how some types of “attentional” phenom- To produce selective behavior, multiple, inter-related pro- ena could be products of value-based selection mechanisms of cesses integrate numerous sources of information. One of the the basal ganglia. If selectivity is a design feature that emerged challenges is that these processes unfold over different as the repertoire of behaviors increased in number and sophis- timeframes (e.g., Chapman et al., 2015b; Welsh, Neyedli, & tication (avoidance, approach, saccade, eat, reach, grasp, use Tremblay, 2013). Therefore, in a laboratory setting, if these tools…), it would seem futile to search for a single dedicated processes are only observed during one point or snapshot functional or neural subsystem generating selection. We feel during the selection process, the observation could appear to that this futility is the reason that attention research has so reflect “attention” or “intention” or “decision making and re- many longstanding and rather fruitless debates about the true ward.” The synthetic approach proposed here also rectifies origins and processes of selective attention. These debates are and makes explicit that reward and selection history are commonly binary in nature because the debates start with the intertwined subjects, but likely reflect multiple processes that assumption of one cause or singular central core system. As contribute to goal-oriented behavior. For example, the syn- this one cause is then increasingly challenged by additional thetic approach can account for harm avoidance. research findings, another, commonly opposite cause is Specifically, harmful stimuli should receive priority process- established… and the process and debate continues. We sus- ing for detection, yet the organism should move away from pect that none of these debates will come to an end, simply these stimuli. The primitive neural circuits for reward/ because the proponents of all camps are “correct” in some way approach and harm/avoid processes diverge early in evolu- and in some cases given that selectivity is a feature of the tionary history, providing a process-based account for diver- system that has emerged from the interaction of many factors gent findings regarding positive and negative value-based across evolution. stimuli. Likewise, the synthetic approach explains why sensi- And yet, we strongly feel that these debates do not move tivity to different features of objects depends on the action our field forward; that they do not really increase our under- context (Bekkering & Neggers, 2002; Craighero et al., 1999; standing of how “attentional phenomena” are generated. Fagioli et al., 2007; Welsh & Pratt, 2008) – because the con- Pursuing the analytic approach and trying to use one concept text determines which action-centered parietal stream, with its like “attention” to explain all of these results (that is, as a idiosyncratic representation of the external world, is being singular explanans) is problematic – the term invariably gets selectively invigorated at a given time. spread so thin, across so many different findings, that it ends One of the great conundrums in experimental psychology up being too vague to have any empirical punch. Researchers and neuroscience is exactly how all of these streams of infor- are right to pursue these as multiple explananda, but would be mation diverge from initial sensory areas and then converge to wrong to seek or be forced to rule out only one explanans. produce action. Working backwards from what researchers Rather, inasmuch as it is possible, one should seek to identify observe in behavior, it is known that generally only one the key mechanisms and processes at work and explain each goal-directed movement is performed at a time, though more in turn. than one might be simultaneously represented (e.g., Cisek & Atten Percept Psychophys (2019) 81:2288–2303 2299 Kalaska, 2005). As discussed in the section on evolutionary delineate the phenomena we aim to explain were misleading, adaptations, we advocate for a parallel competitive structure and we feel that this is in particular true for the concept of with winner-take-all dynamics resolving to produce a single attention. Hence, we argue that, in contrast to James’s(1890) action for each action system (e.g., hand and eye). Of course, assertion, no one knows, or can ever know, exactly what at- much of the detail about how this occurs is an open question tention is. and beyond the scope of this article. What we hope to empha- Acknowledgements The authors would like to recognize the financial size here is the synthetic approach to understanding how com- support of the Psychonomic Society. The meeting that brought the au- plex sensory information is transformed into action. The cor- thors of this paper together was funded by the Leading Edge Workshop ollary argument is that progress is hindered when we appeal to Award from the Psychonomic Society (awarded to JHS and TNW). The or attempt to apply catch-all terms like “attention.” Thus, rath- research presented in this paper was supported by numerous grants in- cluding: Advanced Grant of the European Research Council (ERC-2015- er than saying that an individual “pays more attention to a AdG-694722) to BH; Operating Grants from the Canadian Institute of physically salient stimulus,” one should make an attempt to Health Research to PC; National Science Foundation grant BCS-1555006 understand the mechanism by which physical salience trans- to JHS; and Discovery Grants from the Natural Sciences and Engineering lates to more efficient processing and behavior. Instead of Research Council to CSC, PC, HFN and TNW. The authors would also like to thank the Brouwerij 't IJ pub for providing the atmosphere and arguing that rewarding stimuli “demand more attention,” pro- nourishment that stimulated and facilitated the discussions that led to this vide a description of how a particular reward is associated paper. with a particular target, and how, perhaps even more astound- ingly, the cognitive system/brain then recalls this association Open practices statement None of the data or materials for the exper- iments reported here are available, and none of the experiments was in a fraction of a second to guide behavior on a subsequent preregistered. trial. Experiment to figure out how and why visual informa- tion presented at a location selected for action is amplified, Open Access This article is distributed under the terms of the Creative rather than passing the finding off as “just attention.” Hence, Commons Attribution 4.0 International License (http:// turn to the mechanisms that we understand and try to re-create creativecommons.org/licenses/by/4.0/), which permits unrestricted use, the behavior that cognitive and neural scientists are interested distribution, and reproduction in any medium, provided you give appro- in. If that approach turns out to be successful, there will be no priate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. need for undefinable concepts like attention, either in describ- ing the explanandum or in describing the explanans. The synthetic approach we suggest here might appear re- ductionist. 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Springer Journals
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Copyright © 2019 by The Author(s)
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Psychology; Cognitive Psychology
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1943-3921
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1943-393X
DOI
10.3758/s13414-019-01846-w
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Abstract

In this article, we challenge the usefulness of “attention” as a unitary construct and/or neural system. We point out that the concept has too many meanings to justify a single term, and that “attention” is used to refer to both the explanandum (the set of phenomena in need of explanation) and the explanans (the set of processes doing the explaining). To illustrate these points, we focus our discussion on visual selective attention. It is argued that selectivity in processing has emerged through evolution as a design feature of a complex multi-channel sensorimotor system, which generates selective phenomena of “attention” as one of many by-products. Instead of the traditional analytic approach to attention, we suggest a synthetic approach that starts with well-understood mecha- nisms that do not need to be dedicated to attention, and yet account for the selectivity phenomena under investigation. We conclude that what would serve scientific progress best would be to drop the term “attention” as a label for a specific functional or neural system and instead focus on behaviorally relevant selection processes and the many systems that implement them. . . . . . . . . Keywords Attention Motor control Selection Sensorimotor Decision making Phylogenetic Intention Evolution Parietal cortex Superior colliculus Introduction however, that the overuse and popularity of this statement in cognitive research has been detrimental to progress – that in “Everyone knows what attention is” (James, 1890)isone of fact, no one knows what attention is. More specifically, we the most popular quotes from William James and certainly the argue that the concept of “attention” is one of the most mis- most famous statement about human attention. We argue, leading and misused terms in the cognitive sciences. In the present paper, we stake the position that the term “attention” Note that William James’ own approach to attention was mainly concerned should be abandoned and the nature of the research in this area with phenomenology, an aspect of attention that we do not further consider in be re-conceptualized to focus on the subsets of processes and this article given the modern functional and neural approaches we address mechanisms that lead to task-specific performance. Similar instead. positions have been proposed and discussed previously (see Anderson, 2011;Di Lollo, 2018; Hommel & Colzato, 2015; * Timothy N. Welsh t.welsh@utoronto.ca Krauzlis, Bollimunta, Arcizet, & Wang, 2014;Mole, 2011). The present paper reaffirms and expands this position by plac- Institute of Psychology, Cognitive Psychology Unit and Leiden ing particular and new emphasis on the interconnected and Institute for Brain and Cognition, Leiden University, Leiden, the integrative nature of the human sensorimotor information pro- Netherlands cessing systems. This emphasis on integrated sensori- Faculty of Kinesiology, Sport, and Recreation, University of Alberta, cognitive-motor processes takes inspiration from the synthetic Edmonton, Alberta, Canada approach to understanding “cognition” (Hommel & Colzato, Department of Neuroscience, University of Montreal, 2015) and a proposed phylogenetic refinement of the scientific Montreal, Quebec, Canada approach to understanding behavior (Cisek, 2019 [this issue]). School of Health and Human Performance, Dalhousie University, In the present paper, we start by discussing and outlining Halifax, Nova Scotia, Canada the central problem with the way “attention” has been concep- Department of Cognitive, Linguistic and Psychological Sciences, tualized and studied thus far. We make the case for adopting a Brown University, Providence, RI, USA synthetic approach to studying cognitive phenomena wherein Centre for Motor Control, Faculty of Kinesiology and Physical the focus is on the subset of processes and mechanisms that Education, University of Toronto, 55 Harbord Street, have been attributed to and investigated under the umbrella of Toronto, ON M5S 2W6, Canada Atten Percept Psychophys (2019) 81:2288–2303 2289 “attention,” rather than on “attention” as one overarching con- Before we explain our main position and arguments, cept. To bolster our analysis of the state of affairs, we present we would like to emphasize that the theoretical problems two test cases. In the first, we examine the debate about the that we highlight are particularly visible with respect to conceptual distinction between attention and intention, and the concept of attention, but by no means restricted to show that this debate fails to adequately account for the avail- that concept (Hommel, 2019a). For example, very similar able data. In the second, we review research on selection and arguments to those that we present in the following have reward history to show how conventional analytic approaches been put forward to question the concept of memory. to solving this problem are ineffective. As an alternative to the Decades of research on human memory have seen an analytic approach, we provide a brief review of the phyloge- ever-increasing number of memory systems that were netic evolution of the human brain (for an expanded account, thought to represent separable aspects of memory perfor- see Cisek, 2019 [this issue]) and show how selective attention mance, which then were thought to be explained by the emerged as just one necessary consequence of the challenges existence of corresponding memory systems, with rather facing animals behaving in the natural world. In the end, we limited contributions to a mechanistic understanding of conclude that the traditional analytic attempt to lump many the underlying processes (Bechtel, 2008). As recent con- diverse empirical observations under one common umbrella siderations suggest, however, the various types of mem- called “attention” and to try to explain all of them by referring ory may not at all reflect the operations of separable to one coherent attentional system has actually failed, and dedicated systems, but rather stand for different should be replaced by a more synthetic approach. This syn- byproducts of normally functioning cognitive systems thetic approach focuses on, and starts with, ecologically rele- (Buckner & Schacter, 2004), and emerged at different vant mechanisms and processes and then tries to account for times during the evolution of our species (Murray, as many phenomena (“attentional” or not) as possible. Wise, & Graham, 2017). Similar arguments have been put forward for the concept of emotion (Barrett, 2017; Hommel, 2019b) and may be developed for other con- cepts as well, including “cognition” itself (Cisek, 2019). The concept of attention We focus here on attention because we believe that at least some of the related phenomena are best understood We are not the first to raise concerns about problems with in terms of the kinds of interactions between sensory, the term “attention.” Multiple authors have highlighted motor, and cognitive phenomena that are the focus of the tendency to reify attention, creating circular explana- this special issue. tions for empirical results (Anderson, 2011;DiLollo, Theorizing about human attention suffers from at least 2018). Another common criticism is that multiple process- three main problems. First, the concept of attention invites es underlie what is typically labeled as “attention” (Di misconceptions of one coherent set of cognitive or neural Lollo, 2018; Hommel & Colzato, 2015). Mole (2011) operations, depending on one’s level of analysis, that all highlights that James’ statement came at the time where contribute to what we call “attention” (e.g., Kahneman, there was debate among theorists as to whether the main 1973). Second, the concept of “attention” can also easily role of attention was in thinking, perceiving, or acting. be misunderstood as both an important explanandum that James’ contemporary, F.H. Bradley, produced one of the psychology is rightly expected to explain and the earliest criticisms of the concept of attention, titling his explanans that is supposed to form the explanation – thus essay “Is There Any Special Activity of Attention.” In rendering the latter a pseudo-explanation. And, third, the brief, his position was that there were too many examples concept is thought to distinguish a particular set of cog- of phenomena labeled “attention”, with little concern nitive or neural operations from other, seemingly different about the processes underlying such phenomena sets of operations, such as those related to decisions, in- (Bradley, 1886). Now, over 130 years later, there is a tentions, motivations, emotions, and, of particular rele- continued and heightened need to question the role of vance to the present special issue, action planning and “attention”, the use of the term “attention,” and a search execution. As we show in the expanded discussion of for what “attention” is (e.g., Busemeyer, Gluth, these three points in the following paragraphs, all these Rieskamp, & Turner, 2019; Gottlieb, 2012). We reaffirm assumptions are incorrect. these positions and further suggest that compartmentaliz- First, let us start by considering which phenomena re- ing “attention” and then searching for the “attentional searchers have, historically speaking, been trying to explain system” hinders the development of a comprehensive un- when using the term “attention.” According to some tradition- derstanding of human behavior because it ignores inte- al and conventional views, “attention” is the set of cognitive/ grated, parallel, and reciprocal relationships among senso- neural mechanisms responsible for maximizing the efficient ry, cognitive, and action processes. utilization of our limited capacities to process, store, and 2290 Atten Percept Psychophys (2019) 81:2288–2303 retrieve information. However, consultancy of introductory mechanistic understanding of how the observed phenomena textbooks (e.g., Eysenck & Keane, 2000) and the internet are causally produced (Krauzlis et al., 2014). reveals a dramatic variety of abilities attributed to attention: Third, research on attention has followed, and suffered the ability to select external events for further internal process- from, the common analytical approach to psychological func- ing (focused attention); ignore misleading information and/or tioning (see Hommel & Colzato, 2015,for amore detailed an irrelevant location (selective attention); process irrelevant discussion of this issue). The analytic approach comprises a information (involuntary attention); selectively integrate in- search for an exhaustive definition (which, given the diversity formation belonging to one event within and across sensory of the subfunctions of attention, is impossible), the identifica- modalities (feature integration); prioritize processing of events tion of assumed subfunctions (e.g., overt vs. covert, early vs. from a particular location (spatial attention); systematically late, focused vs. divided, voluntary vs. automatic attention, search for a target event (visual search); perform multiple etc.) with separable functional and neural processes, and the tasks at the same time (divided attention); control the spatial concentration of research on tasks and subfunctions rather parameters of eye movements (selective attention for action); than actual processes. The problem of this analytic approach prioritize one goal over others (goal-centered attention); pri- is that it underestimates and overlooks commonalities be- oritize one object, memory item, or conscious representation tween subfunctions and, in a wider perspective, commonali- over others (object-centered attention); and consolidate infor- ties with other concepts. For instance, the very fact that we use mation for later use and concentrate in anticipation of a pos- concepts like attention, decision making, intention, emotion, sible event over some time (sustained attention). At face value, and motivation in different situations and theoretical contexts it seems highly unlikely that the same set of functional/neural by no means implies that the underlying functional and neural mechanisms are involved in, and responsible for, this broad processes are different and separable. Indeed, attempts to sys- variety of phenomena (Allport, 1993), and a bulk of behav- tematically distinguish the processes “underlying” attention ioral and neural evidence confirms that most subfunctions can from the processes that do not, often fail to produce any co- be dissociated from each other (e.g., Fan, McCandliss, herent consensus. In the following section, we outline one Sommer, Raz, & Posner, 2002). Accordingly, it is unsurpris- exemplar case of the failure of the analytic approach – the ing that no theory has been suggested so far that comes even attempt to separate “attention” from “intention.” close to providing a coherent account of all phenomena sailing under the label of “attention.” Second, the term “attention” is often used to capture both Attention versus intention: A failed the problem and the solution of cognitive processing; i.e., to dichotomy describe both the phenomenon one aims to explain and the mechanism proposed to provide the explanation. For instance, Most researchers agree that the posterior parietal cortex repre- the term attention is used to refer to the consequences of both sents the core of the neural substrate of selective attention, and “voluntary” and “involuntary” factors in favoring the repre- is a key node of an “attentional network” (Corbetta & sentation of one event having a stronger impact on decision Shulman, 2011; Posner & Dehaene, 1994; Ptak, 2012). In making and action than representations of other events (e.g., particular, individual neurons in the posterior parietal cortex Yantis, 1998). But the concept of attention is also used to refer appear to reflect the locus of attention (Bisley & Goldberg, to the system, mechanism, or ability to deal with (or avoid) the 2010; Robinson, Goldberg, & Stanton, 1978) and parietal consequences of such unequal potencies of representations to damage often leads to phenomena of spatial neglect drive behavior (e.g., Broadbent, 1958). Along the same lines, (Bartolomeo, 2007; Corbetta & Shulman, 2011). However, a attention is considered by some to represent the critical capac- separate line of research implicates these same regions of the ity limitation, the cognitive bottleneck that needs to be brain in processes related to movement control (Mountcastle, accounted for and explained (e.g., Pashler & Johnston, Lynch, Georgopoulos, Sakata, & Acuna, 1975; Snyder, 1998), while still others consider attention to be the cognitive Batista, & Andersen, 1997). In particular, regions of the pos- means to deal with such bottlenecks (e.g., Bundesen, 1990). terior parietal cortex are strongly and reciprocally intercon- These conceptual confusions have created a situation in which nected with parts of the frontal lobe that are involved in the it is no longer clear what the to-be-explained problem actually planning and guidance of movement (Johnson, Ferraina, is (do we have a cognitive bottleneck that we need to make the Bianchi, & Caminiti, 1996;Markov etal., 2014), individual best of, or do we have too much information we need to neurons are strongly modulated by the type of action per- choose from?), and whether attention is a concept that refers formed with respect to identical stimuli (Cui & Andersen, to the problem or to the solution. This runs into the danger that 2007; Snyder et al., 1997), and inactivation of the posterior research and theorizing on attention is based on circular rea- parietal cortex causes biases in free-choice tasks soning (attentional phenomena are explained by assuming and (Christopoulos, Kagan, & Andersen, 2018), but not decisions pointing to attentional systems) rather than on a deeper based on visual evidence (Katz, Yates, Pillow, & Huk, 2016). Atten Percept Psychophys (2019) 81:2288–2303 2291 These apparently contradictory findings have fueled a heat- 1996; Gersch, Kowler, & Dosher, 2004;Hoffman & ed and persistent debate, now in its fifth decade, on whether Subramaniam, 1995; Peterson, Kramer, & Irwin, 2004). the posterior parietal cortex is involved in guiding “attention” These studies demonstrate there is preferential processing of or whether it reflects the individual’s “intention.” As is typical stimuli at the goal of a saccade just before the onset of the eye for the dominant analytical approach to psychological science, movement (presumably because “attention” has been shifted researchers have tried to resolve this debate by defining the to the goal location). concepts of attention and intention in ways that make them Other research has extended the study of these action- appear mutually exclusive: “Attention” is what restricts the attention interactions to manual actions, showing that plan- inflow of sensory information to cognition, what enters con- ning and performing reaching and grasping movements prior- scious thought for further processing, whereas “intention” is itizes the processing of the target objects of these movements the output of cognition, the will (free or otherwise) to perform (e.g., Pratt & Abrams, 1994; Rizzolati, Riggio, & Sheliga, a specific action. Defined in this way, the two appear like 1994; Tipper, Lortie, & Baylis, 1992; see also Wu, 2014). distinct concepts that must be dissociable through careful ex- Even when the eyes remain fixated, perceptual discrimination perimental design. And yet, after decades of work by some of is better at the to-be-reached goal than non-goal locations the world’s most accomplished neuroscientists, a clear disso- (Baldauf & Deubel, 2008; Baldauf, Wolf, & Deubel, 2006; ciation of the function of posterior parietal cortex remains Deubel & Schneider, 2003; Deubel, Schneider, & Paprotta, elusive. A prominent review expressed this frustration many 1998; Khan, Song, & McPeek, 2011). The “attentional im- years ago, suggesting that “current hypotheses concerning pa- pact” or prioritized processing associated with intended future rietal function may not be the actual dimensions along which movements goes beyond mere spatial prioritization because the parietal lobes are functionally organized; on this view, other studies have shown that moving or planning to move what we are lacking is a conceptual advance that leads us to also facilitates the detection of action-related features of the test better hypotheses” (Culham & Kanwisher, 2001). object targeted by the movement. For example, preparing for a To escape this rather uncomfortable state of affairs, some grasping movement facilitates the detection of size oddballs, researchers have argued for a more integrative view on atten- while preparing for a pointing movement facilitates the detec- tion and intention. A particularly promising approach is the tion of location oddballs (Fagioli, Hommel, & Schubotz, 2007; pre-motor theory, which argues that shifts of attention are see also Craighero, Fadiga, Rizzolatti, & Umiltà, 1999). Other triggered by sub-threshold saccadic commands in oculomotor studies (Bekkering & Neggers, 2002; Moher, Anderson, & areas and, conversely, shifts of attention in space lead to action Song, 2015; Tipper, Meegan, & Howard, 2002; Weir et al., planning (e.g., Rizzolatti, Riggio, Dascola, & Umilta, 1987). 2003; Welsh & Pratt, 2008; Welsh & Zbinden, 2009;see also Support for the former idea has come from a large number of Gallivan, Barton, Chapman, Wolpert, & Flanagan, 2015; observations including: (a) behavioral studies showing that Glazebrook, Welsh, & Tremblay, 2016; Yoxon, Constable, & attention and eye movements are strongly linked behaviorally Welsh, 2019 [this issue]) show that the processing of specific (e.g., Deubel & Schneider, 1996; Kowler, Anderson, Dosher, object features can be prioritized depending on the relative (i.e., &Blaser, 1995; Sheliga, Riggio, & Rizzolatti, 1995); (b) task/action-specific) salience of those features for the to-be- fMRI studies of visual attention showing activation in eye- performed action. That is, the same feature (e.g., orientation) movement areas for attention tasks (e.g., Beauchamp, Petit, can be prioritized in one action context (e.g., grasping), but not Ellmore, Ingeholm, & Haxby, 2001; Corbetta et al., 1998; another action context (e.g., pointing). Hence, it is neither phys- Nobre, Sebestyen, & Miniussi, 2000)and for movement acti- ical stimulus properties nor action goals alone that generate vation (decoding) in retinotopically defined visual cortex dur- selectivity, but rather selectivity is shaped by the reciprocal ing movement tasks (Gallivan, Chapman, Gale, Flanagan, & and iterative interactions between these factors. These findings Culham, 2019); (c) stimulation studies showing that activation thus suggest that multiple functional and neural systems are of neurons in the superior colliculus (SC), frontal eye field involved in selective attention. (FEF), and lateral intraparietal area (LIP) can change the focus In addition to the interactive nature of stimulus properties of attention (Cavanaugh & Wurtz, 2004;Cutrell&Marrocco, and action goals in determining selection and prioritization of 2002; Moore & Fallah, 2001; Muller, Philiastides, & locations and features, it does not seem that selection stops Newsome, 2005); and (d) neurological studies of patients with solely within any putative attentional system. Indeed, neural attentional disorders following damage to the frontal cortex, activity related to multiple simultaneously active intentions to parietal cortex, or midbrain (e.g., Husain & Kennard, 1996; act at potential target locations, as well as the selection of the Posner, Cohen, & Rafal, 1982; Posner, Rafal, Choate, & final target, has been identified in various structures more Vaughan, 1985; Sapir, Soroker, Berger, & Henik, 1999). Of commonly associated with the planning and execution of ac- particular relevance to the present purpose are behavioral stud- tions, such as the dorsal premotor area, the parietal reach re- ies revealing that perceptual discrimination at the goal location gion, and the motor cortex (Cisek & Kalaska, 2005; Klaes, of an upcoming saccade is improved (Deubel & Schneider, Westendorff, Chakrabarti, & Gail, 2011; Pesaran, Nelson, & 2292 Atten Percept Psychophys (2019) 81:2288–2303 Andersen, 2008; Scherberger & Andersen, 2007;Song& selection. Given that this would make it no longer apparent McPeek, 2010; Thura & Cisek, 2014). Behaviorally, the pres- that such a reduced version of “attention” has anything to do ence of multiple co-existing response representations and the with other “attentional” functions like integration, orientation, dynamic selection of target from non-target stimuli and ac- or vigilance, this would eventually call for dropping the con- tions is also expressed through the spatiotemporal character- cept – and the same argument holds for “intention.” On the istics of reaching and grasping movements. Specifically, in- positive side, this would prevent researchers from trying to stead of the efficient straight and direct movements that one find commonalities in processes and substrates that are unlike- might anticipate if attentional selection had been completed ly to be found. On the negative side, however, there is no prior to the intention to act, the trajectories of hand and eye theoretical justification to pick just these functional aspects movements veer towards or away from non-target stimuli de- but not others. What looks like a definitional issue thus be- pending on the timing and salience of the non-target stimuli comes a theoretical bias that is lacking justification. (e.g., Chapman et al., 2010; Gallivan & Chapman, 2014; Alternatively, one might search for hybrid approaches that Howard & Tipper, 1997;Moheretal., 2015; Neyedli & allow for additional components and factors. A typical ap- Welsh, 2012; Song & Nakayama, 2006, 2008; Welsh, 2011; proach of this sort was the resource theory of attention, which Welsh & Elliott, 2004; Wispinski, Gallivan, & Chapman, in triggered heated debates in the 1970s and 1980s (Kahneman, press). Thus, the characteristics of the physically executed 1973; Navon, 1984). While the first approaches were simple action actually reflect the “attentional” state of the target and and elegant by assuming one kind of resource that needs to be non-target stimuli. Collectively, these data indicate that atten- distributed over all mental work, the attempt to integrate an tion, selection, and intention are not readily separated in a set increasing number of unpredicted findings led to the invention of discrete serial processes, but are more dynamic and contin- of increasing numbers and types of separate resources. In the uous in nature and embedded within a densely interconnected, end, this made systematic predictions impossible (Navon, parallel processing system. 1984), which is the main reason why this approach no longer While more work needs to be done to synthesize these plays an important role – except in the field of ego-depletion, neural and behavioral observations into a coherent framework, where history seems to repeat itself (Friese, Loschelder, it seems clear (to us) that the conceptual distinction between Gieseler, Frankenbach, & Inzlicht, in press). The main reason attention and intention is not sufficient to account for the va- why hybrid approaches that simply lump together different riety of findings discussed here. The distinction fails to pro- factors are not overly successful rests in the fact that the re- vide a meaningful contribution or framework for sorting the spective factors are not truly integrated into a coherent available findings into useful categories to stimulate further framework. theorizing, and it also clashes with the demonstration of so A similar tendency can be seen with respect to selective attention, where Awh, Belopolsky, and Theeuwes (2012)have many interactions between input processing and output gen- eration. But what is the solution to this and the many other tried to integrate findings that are no longer consistent with the conceptual problems we are encountering in thinking about historical distinction between endogenous attention, which human attention (e.g., controlled vs. automatic processing; represents the prioritized processing of stimuli to which the facilitation vs. inhibition, etc.)? agent “wants” to attend, and exogenous attention, which rep- resents the prioritized processing of stimuli that are unrelated to the present action and goals. The history of distinguishing A failed analytic solution: Selection between endogenous and exogenous attention is very similar and reward history to the distinction between attention and intention. Each started out by trying to improve definitions about what the concept As noted above, the dominant analytical approach to psycho- referred to, only to be faced later with the inability to system- logical functioning begins with an exhaustive search for a atically sort the available findings into two distinct categories. definition of a concept, including the borders of where it dif- In a nutshell, endogenous attention is sometimes too automat- fers from other concepts. So, to understand “attention,” one ic and exogenous attention is sometimes too dependent on the would tend to first define how it differs from “intention,” current prioritized stimulus feature or action goal to make this “decision-making,”“motivation,” etc. In view of a failure of dichotomy fruitful and tenable (Awh et al., 2012;Folk, this approach, as is obvious for the case of attention versus Remington, & Johnston, 1992;Hommel & Wiers, 2017). intention, two reactions are to be expected. First, one might Awh et al. suggest solving this problem by adding a third consider the previous attempts to define attention and inten- variable – selection history – to the list of factors. In particular, tion as flawed and try to improve the definitions by further the idea is that goals (the factor responsible for endogenous reducing the conceptual overlap between the two concepts. attention), salience (the factor responsible for exogenous at- For instance, one may further reduce the concept of attention tention), and selection history (a factor that does not seem to fit to mere input selection and the concept of intention to output the previous dichotomy and is associated with previous Atten Percept Psychophys (2019) 81:2288–2303 2293 selections and rewards generated by the selections) all con- Indeed, prioritizing shape and orientation when grasping ob- tribute to selectivity by sending their output to an integrative jects makes more sense than prioritizing color because those priority map. Although this approach may account for many features are more likely to determine a successful or an unsuc- of the available findings, we are not convinced that it really cessful grasp. Hence, establishing a bias for shape and orien- solves the problem, but rather provides a patch that holds tation over color when grasping would be a functional adap- concepts together and, in the end, prevents or misdirects the tation. However, this influence implies that selection history search for suitable solutions. Instead, we suggest that a com- affects how goals impact (endogenous) attention. Along the plete dismantling of the concept of attention is required. same lines, the relative salience of the visual dimension At first there does not seem to be anything wrong with the changes substantially during the first years of life (e.g., idea that structures, or a singular structure, in the human cen- Suchman & Trabasso, 1966), which at least opens the possi- tral nervous system are devoted to collecting and integrating bility that selection history impacts salience. information that affects prioritized processing. One candidate While these arguments are fully consistent with Awh structure is the superior colliculus, which is thought to reflect a et al.’s suggestion to consider selection history as a third factor priority map of stimuli in the visual field (Fecteau & Munoz, involved in attentional control, they also imply that the 2006). Importantly, and as required from the view that selec- resulting three factors are not independent but strongly over- tivity for behavioral relevance is the purview of the entire lapping and intertwined – both empirically and conceptually. moving body, the superior colliculus is involved not just in As we have tried to explain, these conceptual-overlap prob- eye movements, but in orienting movements of the eye, head, lems are unlikely to be resolved by more definitions. Rather, body, and hand (Gandhi & Katnani, 2011; Stuphorn, what is needed is a theory that not only assumes that integra- Hoffmann, & Miller, 1999). Emphasizing this point, Song tion takes place but that explains how that integration works. and colleagues (Song, Rafal, & McPeek, 2011; Song & Another reason why just adding selection history as an McPeek, 2015) found that the superior colliculus plays a caus- additional factor raises more questions than answers is that al role in target selection during manual reaching tasks, the concept itself is unclear, particularly in its overlap with supporting the idea that the superior colliculus is part of a other related factors beyond exogenous and endogenous con- general-purpose target selection/orientation system trol. One such factor that is intertwined with selection history (Nummela & Krauzlis, 2010; Song et al., 2011). On the other is reward history. It is uncontroversial that previously hand, there is no need to assume that the superior colliculus is rewarded stimuli receive preferential processing (Anderson, the only map that integrates relevant information to steer at- Laurent, & Yantis, 2011; Anderson & Yantis, 2012), suggest- tention, nor is it necessary to assume that all available infor- ing that reward history is important in determining salience. mation is integrated into that one map. As we argue below, the Conventionally, stimuli must have been selected in order for human brain can be considered to have many sources of se- the organism to have received a reward, thus conflating the lectivity and, in the end, it is the brain as a whole that does the two concepts. Awh et al. (2012) appear to acknowledge this integration. Given that this integration is the explanandum tension, considering both previous reward and previous stim- (the to-be-explained phenomenon), postulating the existence ulus selections to be exemplars of “selection history” while of one map that has no other function than achieving this facing the fact that selection history and reward history cannot integration seems to be one more attempt to “explain” apsy- be identical (given that previous selections might not have chological phenomenon by positing the existence of a dedi- received reward). However, the fit of reward history into this cated system whose only purpose is to somehow create that joint category is much less obvious than this theoretical inte- phenomenon. gration suggests since reward history is itself likely composed Apart from this more general meta-theoretical problem, of many differentiable factors. adding one more factor to a model that just assumes that inte- For instance, the preference for rewarding stimuli is stron- gration takes place without explaining how that can be done is ger when the larger expected gain of the stimulus is due to an unlikely to guide further research. In the case of Awh et al. increased probability of receiving a reward than when there is (2012), one reason is that selection history overlaps consider- a lower cost of failure (Neyedli & Welsh, 2015b). This finding ably with goal-induced endogenous selectivity and salience- suggests that what participants take to be rewarding is itself induced exogenous selectivity. For instance, the fact that plan- multifaceted – not just the magnitude of the reward, but also ning and carrying out particular kinds of actions systematical- how likely it is that they will receive the reward. Furthermore, ly facilitates the processing of particular object features (e.g., across a series of reach-decision experiments, a multitude of of size and orientation for grasping, location for reaching: biasing factors have been observed including: reward value Bekkering & Neggers, 2002; Craighero et al., 1999; Fagioli and probability (Chapman, Gallivan, & Enns, 2015a), the best et al., 2007) is unlikely to be genetically determined, but rather option in a decision set (Wispinski, Truong, Handy, & the consequence of learning and experience of selecting dif- Chapman, 2017), current level of accumulated wealth ferent features for grasping over the lifespan (Hommel, 2010). (Neyedli & Welsh, 2015a), the number of targets and not the 2294 Atten Percept Psychophys (2019) 81:2288–2303 perception of them (Milne et al., 2013), and how the number A synthetic approach of choice-options is represented (Chapman et al., 2014). The problem here is similar to the one Awh et al. (2012)tried to A synthetic approach is valuable only insofar as it synthesizes solve: the definition of the concept under investigation is too elements that actually correspond to real biological processes restricted. In their case, endogenous and exogenous control at both neural and functional levels, and it is a significant were insufficient to account for the variety of pheomenena challenge to figure out what those processes are. One power- being ascribed to the concept of attention, so they added se- ful strategy for doing so – for keeping our synthesis close to lection history. Selection history is itself decomposable into biological reality – is to use evolution as a guide. This guid- (at least) selection and reward history, and reward history is ance can be done through a procedure of “phylogenetic refine- itself decomposable even further. Thus, the nature and the ment,” whereby one progressively elaborates a theory about influence of reward is itself dependent on numerous contex- how neural and behavioral processes evolved along a given tual factors and the expression of the confluence of these fac- lineage, always respecting constraints about the neural modi- tors is not easily captured in a unitary construct. fications and behavioral adaptations that appeared at each The picture gets even more complicated when one con- stage (see Cisek, 2019 [this issue]). For this reason, here we siders studies comparing the impact of rewards of equal step away from the concept of “attention” and take a brief magnitude but in opposite directions (e.g., positive/gain detour into the history of how the relevant neural circuits vs. negative/loss). For example, loss aversion, made fa- evolved in the lineage that leads to homo sapiens (humans). mous by the work of Kahneman and Tversky (1979), While it is often very difficult to know why a given modifica- shows that people treat potential losses as being more aver- tion took place in evolution, establishing what was the se- sive than equivalent gains are rewarding. Interestingly, quence of modifications can be constrained by a wealth of when decisions between positively and negatively reward- comparative and developmental data, leading to strong and ing stimuli are made rapidly (Chapman et al., 2015b), testable hypotheses about how neural circuits and behavioral asymmetries in choice behavior are also observed, but go abilities evolved together. opposite to loss aversion. That is, in these situations, par- The evolutionary history of spatial interaction along the ticipants appear to be disproportionately drawn toward op- primate lineage is a long and complex tale (Fig. 1). A major tions giving gains, while the aversive impact of loss-related advance occurred during the Cambrian epoch, over 500 mil- choices is attenuated. These findings highlight an addition- lion years ago (Mya), with the elaboration of visually guided al complication – that different biases are likely to operate orientation behaviors. Our simple chordate ancestors pos- on different timelines. For instance, in the study of the sessed a visual escape circuit that involved projections from asymmetryindecisionstogo for agoodoptionversus a single photosensitive patch in the rostral neural tube to a avoid a bad one, participants were biased toward positively midbrain structure called the tectum, which projected to the valanced targets 100 ms earlier than they were biased to spinal cord to generate locomotion (Lacalli, 1996, 2018). In move away from negative ones (Chapman et al., 2015b). the lineage leading to vertebrates, the photosensitive patch This finding echoes related work showing that more time is split into two lateral eye patches on both sides of the head required to select optimally between visuomotor choices (Butler, 2000). Because these eye patches projected contra- when they differ based on negative value information com- laterally to the tectum, which projected ipsilaterally to the pared to when they differ based on the probability of re- spinal cord, the circuit caused our ancestors to turn away from ward (Neyedli & Welsh, 2015b). salient visual stimuli such as the shadow from an approaching Taken together, these and other findings strongly suggest predator (Fig. 2a). As the eye patches expanded, they folded that selection history is tightly interconnected with reward into cups and formed a lens (Lamb, 2013), resulting in a two- history and that neither of these concepts are particularly well dimensional retina that provided a topographic mapping of understood. Hence, adding selection history to the two other external stimuli. The tectum expanded in parallel, with a not well understood concepts of exogenous and endogenous matched topographic map of space in its superficial layers attention is unlikely to help much in understanding the mech- and gradients of downstream projections in its deep layers. anisms underlying human selective attention. To be clear, we The result was an “action map” of oriented escape responses are not advocating an alternative theory at this point, but rather to threatening stimuli at specific locations in the external we suggest an alternative theoretical perspective: Let us re- world. place the analytical approach, which seeks to explain complex Microstimulation studies reveal the presence of an orga- phenomena by first carefully defining them and then nized map of oriented escape responses in the tectum of lam- subdividing them into simpler elements, with a synthetic ap- prey (Saitoh, Menard, & Grillner, 2007), a jawless fish whose proach that considers how simple mechanisms and functional ancestors diverged from ours about 550 Mya. These studies processes, each of which is itself behaviorally relevant, can also reveal the presence of another action map, which lies together give rise to complex phenomena. within the rostral region of the tectum. This map is sensitive Atten Percept Psychophys (2019) 81:2288–2303 2295 Fig. 1 A reduced phylogenetic tree of bilaterally symmetric animals, (paleobiodb.org). Small rectangles indicate the estimated latest timing exclusively emphasizing the lineage that leads to humans. Branch of innovations described in the boxes. Note that many branch points points represent some of the divergences between different lineages, and lineages are omitted for clarity. Silhouettes along the right are from with timing estimated on the basis of molecular clock analyses (Erwin phylopic.org et al., 2011). Thick lines indicate the presence of relevant fossil data to space in front of the animal, and projects mostly contra- accomplished through lateral inhibition that produces “win- laterally to the spinal cord, thereby producing orientation and ner-take-all” dynamics (Grossberg, 1973; Mysore & approach actions (Jones, Grillner, & Robertson, 2009; Knudsen, 2011; Wang, 2002). Kardamakis, Saitoh, & Grillner, 2015). It is this latter tectal How is this related to attention? A few sentences after that sub-circuit that is most relevant to attention and selection. In famous phrase we quoted above, James wrote that attention the avoidance circuit, multiple stimuli can engage multiple “implies a withdrawal from some things in order to deal ef- escape actions that can simply be averaged downstream to fectively with others.” That withdrawal from some stimuli to produce adaptive avoidance behavior (Fig. 2b). In contrast, interact with another stimulus is indeed accomplished, quite averaging cannot work in an approach circuit, because the literally, within the approach circuit of the rostral tectum. And average response to two stimuli will cause the animal to miss while these simple circuits for governing interactive behavior both of them (Fig. 2c). Consequently, the approach circuit may seem far removed from the higher cognition of humans, must select between actions, such that one completely sup- they are indeed the precursors to the mechanisms that control presses the other. This kind of selection could be what has been called “selective attention.” The tectum is 2296 Atten Percept Psychophys (2019) 81:2288–2303 Fig. 2 Circuits for avoidance and approach in a hypothetical early stimulus. (b) Spatial averaging of escape directions (numbered arrows) vertebrate. (a) In the avoidance circuit, visual information from the away from two threatening stimuli (black stars) is an effective response. lateral eyes arrives in the contralateral tectum, which projects (c) For approach actions, spatial averaging is maladaptive, making ipsilaterally to the midbrain locomotor regions. Thus, if a stimulus falls winner-take-all dynamics necessary. B and C reused with permission on the left eye, the locomotion will tend to turn to the right until from Cisek (2019) stimulation is balanced and the body is oriented away from the homologous to the human superior colliculus, which, as and each projected to a specific set of relevant effectors. In a discussed earlier, is strongly implicated in both orienting gaze sense, each circuit was an “action map” analogous to the much through eye and head movements and in controlling covert older tectal systems for approach and avoidance, but guiding attention when gaze is stationary (Basso & May, 2017). the much wider repertoire of task-specific interactions avail- Though much has developed in the central nervous system able in the mammalian niche. and the world since our lineage diverged from lamprey in As the behavioral repertoire of mammals continued to ex- the early Cambrian, both the approach and avoidance circuits pand, so did the dorsomedial neocortex, and there was a dif- of the tectum are still present in fish (Herrero, Rodriguez, ferentiation and specialization of action-specific maps of sen- Salas, & Torres, 1998) and in mammals (Comoli et al., 2012). sory space. In primates, expansion of the parietal cortex was Eventually, our ancestors left the seas and some of them, particularly dramatic, yielding a variety of idiosyncratic rep- the amniotes, adapted to a fully terrestrial lifestyle. This adap- resentations of space particular to the needs of different action tation was accompanied by an expansion and lamination of types (Andersen, Snyder, Bradley, & Xing, 1997; Stein, 1992) the telencephalic pallium, an integrative olfactory, visual, and (see Fig. 3). For example, visually guided reaching actions somatosensory region that would eventually give rise to the involve the medial intraparietal cortex (Cui & Andersen, cerebral cortex (Aboitiz & Montiel, 2015;Striedter, 2005). In 2007;Kalaska &Crammond, 1995), which represents targets all mammals, the neocortex consists of two sheets (Finlay & within reach with respect to the direction of gaze and the Uchiyama, 2015), a dorsomedial sector that is spatially topo- position of the hand (Buneo, Jarvis, Batista, & Andersen, graphic and a ventrolateral sector that is non-topographic. In 2002; Gallivan, Cavina-Pratesi, & Culham, 2009), and is in- primates, the former includes a medial and dorsolateral pre- terconnected with frontal regions controlling reaching, such as frontal cortex, cingulate regions, all of the premotor, motor, the dorsal premotor cortex (Johnson et al., 1996;Wise, sensorimotor, and parietal cortex, as well as the retrosplenial Boussaoud, Johnson, & Caminiti, 1997). Grasp control in- cortex. The latter includes parts of the lateral prefrontal cortex, volves the anterior intraparietal area (Baumann, Fluet, & orbitofrontal cortex, and all of the limbic cortex and the tem- Scherberger, 2009), which is sensitive to object shape and is poral lobe. Most relevant to the issue of attention is the interconnected with grasp-related frontal regions such as the dorsomedial sector of the neocortex, which is organized into ventral premotor cortex (Nakamura et al., 2001;Rizzolatti & a set of fronto-parietal circuits dedicated to different classes of Luppino, 2001). The control of gaze involves the lateral species-typical actions (Graziano, 2016; Kaas & Stepniewska, intraparietal area (Snyder, Batista, & Andersen, 2000), which 2016). In early mammals (300 Mya), this system was proba- represents space in a retinotopic frame (Colby & Duhamel, bly quite limited and consisted simply of medial circuits con- 1996; Snyder, Grieve, Brotchie, & Andersen, 1998), and is cerned with locomotion and lateral circuits concerned with interconnected with frontal regions controlling gaze, such as head and mouth movements (Kaas, 2017). Each of these cir- the frontal eye fields and the superior colliculus (Paré & cuits processed sensory information in an idiosyncratic man- Wurtz, 2001) – taking advantage of the tectal orientation/ ner specialized for its specific type of action (e.g., space near approach system that has been steering animals since the the legs for locomotion, space near the snout for ingestion) Cambrian epoch. Atten Percept Psychophys (2019) 81:2288–2303 2297 Fig. 3 The primate cerebral cortex contains a set of parallel sensorimotor relevant given the current behavioral context. AIP anterior intraparietal streams in the dorsomedial regions (blue arrows), each involved in a area, FEF frontal eye fields, IT inferotemporal cortex, LIP lateral specific type of action using specific representations of space. All of intraparietal area, LPFC lateral prefrontal cortex, MIP medial these use information on object identity and outcome value, computed intraparietal area, OFC orbitofrontal cortex, PMd dorsal premotor in the ventrolateral regions (red arrows), to select the actions most cortex, PMv ventral premotor cortex, V1 primary visual cortex In many situations, different fronto-parietal action streams goals and avoid negative outcomes. The phylogenetic per- will compete against each other. For example, one must make spective suggests that the primate posterior parietal cortex is an all-or-none decision as to whether to burrow at the roots of part of a topographically organized dorsomedial neocortical one tree or instead walk over to another tree. In other situa- system for visually guided interactions oriented with respect tions, however, different fronto-parietal streams will be coor- to objects in the world (Cisek, 2007). This system is organized dinated. For example, when a head/snout orientation system as parallel sensorimotor streams, each contributing to a spe- points at a target, that target is then made available to other cific type of action within the animal’s behavioral repertoire, behaviors, such as burrowing or biting. This availability be- whose activity is orchestrated through selective invigoration, comes particularly important in primates, which evolved from energization, or drive from the basal ganglia and other struc- tree-climbing insect eaters and developed large eyes with a tures (Cisek & Thura, 2018; Grillner, Robertson, & central, high-resolution fovea, and acquired a taste for fruit. Stephenson-Jones, 2013). Within each of these fronto- In such animals, the system for controlling the orientation of parietal action streams, target selection occurs through gaze takes on an executive role for many other visually guided winner-take-all dynamics taking place in an idiosyncratic spa- behaviors. Selecting a target for gaze becomes part of tial reference frame specific to each given type of action (e.g., selecting what to reach for or which branch to grasp to climb. retinotopic for eye movements). One of those streams is con- It comes to serve much of the role traditionally ascribed to cerned with orienting gaze through eye and head movements, “selective attention.” Indeed, it has long been proposed that and appears to have an executive role simply because so many selective attention, both overt and covert, is closely related to of the other streams rely on high-resolution visual information the gaze orientation system and involves the same neural that is derived from the fovea. When an animal (including structures (Corbetta et al., 1998; Rizzolatti et al., 1987), in- humans) is placed in a laboratory situation and trained/ cluding the posterior parietal cortex, the frontal eye fields, and instructed to perform just one isolated aspect of complex nat- the superior colliculus. ural behavior, what the researcher will observe in this region is And so, this brief foray into the long journey of primate activity that appears to be related to what the researcher has evolution has brought us back to selective attention and to the defined as “attention,”“intention,” or “decision-making,” de- posterior parietal cortex, but with a different perspective on pending on the particular task variables that are being exper- both. The question is not whether the posterior parietal cortex imentally manipulated. But it does not follow from these cor- plays a role in something called “attention” or something else relations that there exists anything in the brain that can be called “intention,” but how the posterior parietal cortex fits meaningfully delineated as an “attentional system” (or, for that within a broader system that enables animals to select and matter, an “intention system” or “decision system”). The key control interactions with their environment to achieve their insight is that the posterior parietal cortex is not part of an 2298 Atten Percept Psychophys (2019) 81:2288–2303 “attention system” but, rather, that selective attention phenom- In an analytic approach to science, one runs the risk of ena are part of what the posterior parietal cortex produces as it becoming a slave to the concepts that have been generated. goes about its business of controlling goal-directed action. Many researchers have taken terms like “attention,”“inten- tion,” and “decision making” from everyday language and expect this linguistic categorization to somehow map to iden- Summary and conclusions tifiable mechanisms in the brain or functions. Of course, when one starts to peer into actual neural functions, there is no clear We thus conclude that selectivity emerged through evolution delineation, only a set of processes that interact to create se- as a design feature to enable efficient goal-directed action. lectivity in the end. These processes interact not because they Such selectivity became necessary as the action repertoire of belong to a dedicated system, but because the human brain the given line of organisms that led to humans increased. This and body evolved this way and selectivity was a necessary means that selectivity is an emerging property arising from a feature to achieve efficient behavior. Further, everything an myriad underlying processes, and the simple fact that humans individual does throughout their life (distant and recent past) (and other species showing selective attention) evolved the creates, reinforces, and shapes selection: Turning to the left way they did, with selective attention being one of many makes us ignore stimuli on the right, picking one apple makes byproducts, next to “selective intention” and “selective deci- us overlook the others, saying one word prevents us from sion making.” Here, we have primarily emphasized selection uttering any other. And each of the different selections results mechanisms in the superior colliculus and parietal cortex, but in all ranges of rewards, from positive gains to negative losses. similar arguments can be made for other selection Selection and reward are thus inherent ingredients of all our mechanisms in other brain regions. For example, Krauzlis lives and the way we lead them (Allport, 1987). et al. (2014) suggest how some types of “attentional” phenom- To produce selective behavior, multiple, inter-related pro- ena could be products of value-based selection mechanisms of cesses integrate numerous sources of information. One of the the basal ganglia. If selectivity is a design feature that emerged challenges is that these processes unfold over different as the repertoire of behaviors increased in number and sophis- timeframes (e.g., Chapman et al., 2015b; Welsh, Neyedli, & tication (avoidance, approach, saccade, eat, reach, grasp, use Tremblay, 2013). Therefore, in a laboratory setting, if these tools…), it would seem futile to search for a single dedicated processes are only observed during one point or snapshot functional or neural subsystem generating selection. We feel during the selection process, the observation could appear to that this futility is the reason that attention research has so reflect “attention” or “intention” or “decision making and re- many longstanding and rather fruitless debates about the true ward.” The synthetic approach proposed here also rectifies origins and processes of selective attention. These debates are and makes explicit that reward and selection history are commonly binary in nature because the debates start with the intertwined subjects, but likely reflect multiple processes that assumption of one cause or singular central core system. As contribute to goal-oriented behavior. For example, the syn- this one cause is then increasingly challenged by additional thetic approach can account for harm avoidance. research findings, another, commonly opposite cause is Specifically, harmful stimuli should receive priority process- established… and the process and debate continues. We sus- ing for detection, yet the organism should move away from pect that none of these debates will come to an end, simply these stimuli. The primitive neural circuits for reward/ because the proponents of all camps are “correct” in some way approach and harm/avoid processes diverge early in evolu- and in some cases given that selectivity is a feature of the tionary history, providing a process-based account for diver- system that has emerged from the interaction of many factors gent findings regarding positive and negative value-based across evolution. stimuli. Likewise, the synthetic approach explains why sensi- And yet, we strongly feel that these debates do not move tivity to different features of objects depends on the action our field forward; that they do not really increase our under- context (Bekkering & Neggers, 2002; Craighero et al., 1999; standing of how “attentional phenomena” are generated. Fagioli et al., 2007; Welsh & Pratt, 2008) – because the con- Pursuing the analytic approach and trying to use one concept text determines which action-centered parietal stream, with its like “attention” to explain all of these results (that is, as a idiosyncratic representation of the external world, is being singular explanans) is problematic – the term invariably gets selectively invigorated at a given time. spread so thin, across so many different findings, that it ends One of the great conundrums in experimental psychology up being too vague to have any empirical punch. Researchers and neuroscience is exactly how all of these streams of infor- are right to pursue these as multiple explananda, but would be mation diverge from initial sensory areas and then converge to wrong to seek or be forced to rule out only one explanans. produce action. Working backwards from what researchers Rather, inasmuch as it is possible, one should seek to identify observe in behavior, it is known that generally only one the key mechanisms and processes at work and explain each goal-directed movement is performed at a time, though more in turn. than one might be simultaneously represented (e.g., Cisek & Atten Percept Psychophys (2019) 81:2288–2303 2299 Kalaska, 2005). As discussed in the section on evolutionary delineate the phenomena we aim to explain were misleading, adaptations, we advocate for a parallel competitive structure and we feel that this is in particular true for the concept of with winner-take-all dynamics resolving to produce a single attention. Hence, we argue that, in contrast to James’s(1890) action for each action system (e.g., hand and eye). Of course, assertion, no one knows, or can ever know, exactly what at- much of the detail about how this occurs is an open question tention is. and beyond the scope of this article. What we hope to empha- Acknowledgements The authors would like to recognize the financial size here is the synthetic approach to understanding how com- support of the Psychonomic Society. The meeting that brought the au- plex sensory information is transformed into action. The cor- thors of this paper together was funded by the Leading Edge Workshop ollary argument is that progress is hindered when we appeal to Award from the Psychonomic Society (awarded to JHS and TNW). The or attempt to apply catch-all terms like “attention.” Thus, rath- research presented in this paper was supported by numerous grants in- cluding: Advanced Grant of the European Research Council (ERC-2015- er than saying that an individual “pays more attention to a AdG-694722) to BH; Operating Grants from the Canadian Institute of physically salient stimulus,” one should make an attempt to Health Research to PC; National Science Foundation grant BCS-1555006 understand the mechanism by which physical salience trans- to JHS; and Discovery Grants from the Natural Sciences and Engineering lates to more efficient processing and behavior. Instead of Research Council to CSC, PC, HFN and TNW. The authors would also like to thank the Brouwerij 't IJ pub for providing the atmosphere and arguing that rewarding stimuli “demand more attention,” pro- nourishment that stimulated and facilitated the discussions that led to this vide a description of how a particular reward is associated paper. with a particular target, and how, perhaps even more astound- ingly, the cognitive system/brain then recalls this association Open practices statement None of the data or materials for the exper- iments reported here are available, and none of the experiments was in a fraction of a second to guide behavior on a subsequent preregistered. trial. Experiment to figure out how and why visual informa- tion presented at a location selected for action is amplified, Open Access This article is distributed under the terms of the Creative rather than passing the finding off as “just attention.” Hence, Commons Attribution 4.0 International License (http:// turn to the mechanisms that we understand and try to re-create creativecommons.org/licenses/by/4.0/), which permits unrestricted use, the behavior that cognitive and neural scientists are interested distribution, and reproduction in any medium, provided you give appro- in. If that approach turns out to be successful, there will be no priate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. need for undefinable concepts like attention, either in describ- ing the explanandum or in describing the explanans. The synthetic approach we suggest here might appear re- ductionist. 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