Motor Cognition

1. Motor Cognition Overview • Many people believe processes used to plan and enact a movement can be used in problem solving and reasoning • Moreover, the processes involved in perceiving action are also involved in movement • This lecture will introduce key ideas involved in motor cognition and memory for action needed • in planning, and producing our own actions; anticipating, perceiving, and interpreting the actions of others • In remembering actions

2. Motor Cognition Terminology • Movement is a voluntary displacement of a body part in space • Action – a series of movements performed to achieve a goal

3. Motor Cognition Perception-action cycle • Refers to the transformation of perceived patterns (often visually) into coordinated patterns of movement • Examples. Returning a tennis ball, picking up a mug of coffee, walking on uneven terrain • According to this perspective much of human behavior involves the interconnection of perception and movement • The mediating link between perception and action is that a shared representation; that is, the coding of perception and action is shared in the brain

4. Motor Cognition Motor processing in the brain • Neuroanatomy • Area M1, the primary cortex. Neurons in this region control fine motor movements, and send fibers out to muscles themselves • Premotor area (PM) sets up program for motor sequences, and send input into M1 • Supplementary motor area (SMA) sets up and executes motor plans

5. Motor Cognition Motor processing in the brain • Neuroanatomy • Think of these 3 areas as a hierarchy. M1 is at the bottom of the hierarchy, and it enables specific movements; PM higher up and it enables sets of movements; at top SMA represents overarching plans of action

6. Primary motor cortex (M1) Posterior parietal cortex Supplementary motor cortex (SMA) Premotor cortex

7. Motor Cognition Some evidence for roles of three areas • Mushiake (1991) • Recorded single-cell activity in monkeys in M1, PM, and SMA • 2 conditions of interest were IT (internally triggered) and VT (visually triggered) • In both conditions monkeys were required to touch 3 pads on a panel; in the IT condition monkeys needed to remember sequence and then touch the panels in the remembered order; in the VT condition monkeys touched panels as they were illuminated

8. Motor Cognition Some evidence for roles of three areas • M1 cells were active in the premovement and movement periods consistent with the hypothesis that M1 is involved in movement • More SMA cells were active in the IT than the VT condition, consistent with the idea that SMA is important during planning since planning is more important in IT than VT condition • More neurons were active in the VT than the IT condition during the premovement and movement periods

9. Motor Cognition Some evidence for roles of three areas • Conclusions • Movement planning and production involves a number of neuropsychological processes with different functions. These processes occur in different brain regions, and often occur simultaneously • Planning and then producing a response involves different neural processes than responding to environmental cues

10. Motor Cognition Summary • We have reviewed the role of just 3 brain regions involved in motor cognition • Other regions are also involved with their involvement depending on the precise nature of the task; in fact it’s been said that it takes the whole brain to make a cup of coffee

11. Motor Cognition Shared representation • A considerable amount of data suggests that we can efficiently represent the actions made by other people (e.g., through viewing) • It has been hypothesized this occurs (and brain-activation studies support this) because the representation of perceived action and produced action is shared • These representations enable us to interpret the actions of others, respond appropriately, and efficiently learn how to physically produce actions that were viewed

12. Motor Cognition • Mirror Neurons • Mirror neurons—refers to neurons that fire when organism (monkey) performs a specific grasping movement, and when that same grasping movement is observed being performed by experimenter or monkey

13. Motor Cognition • Mirror Neurons • Human evidence • A variety of techniques have been used to provide evidence for motor neurons in humans • A transcranial magnetic stimulation study showed increased excitability in the motor system during observation of actions performed by another person, but only for the brain regions involved in the muscles used by the other person

14. Motor Cognition Apraxia • Definition • An impaired ability to generate skilled actions that cannot be attributed to basic sensory or motor disturbance • Generally conceptualized to reflect a disruption of a distributed praxis network • Research has often focused on transitive actions -- actions that involve manipulation of a tool or object • Examples– use of a hammer, spatula, • Apraxia frequently observed after neurological damage • praxis network was thought to be left lateralized

15. Motor Cognition Diagram of praxis network

16. Component Model ApproachRoy and Square, 1994; Roy, 1996 Sensory/Perceptual System Visual/Gestural Information Auditory/Verbal Information Visual Tool/Object Information Pantomime Conceptual System Knowledge of Action Knowledge of Tool/Object Function Production System Response Selection Image Generation Delayed Imitation Working Memory Concurrent Imitation Response Organization/Control

17. Component Model ApproachRoy and Square, 1994; Roy, 1996 Sensory/Perceptual System Visual/Gestural Information Auditory/Verbal Information Visual Tool/Object Information Main Responsibilities: Analyzing visual gestural information Identifying key features of tools and objects for use

18. Component Model ApproachRoy and Square, 1994; Roy, 1996 Sensory/Perceptual System Visual/Gestural Information Auditory/Verbal Information Visual Tool/Object Information Conceptual System Knowledge of Action Knowledge of Tool/Object Function Main Responsibilities: Understanding tools, objects, and gestures different types of conceptual knowledge are dissociable

19. Motor Cognition • Conceptual knowledge • It appears that different types of conceptual knowledge associated with action are dissociable from each other as demonstrated in the next slides

20. Motor Cognition • Conceptual knowledge • Function knowledge vs manipulation knowledge • Buxbaum and Saffran (2002) investigated function and manipulation knowledge in apraxic and non-apraxic patients with LHD (aside, px were apraxic to gestural tests including tests of pantomime) • Function knowledge – which two items are most similar in function (e.g., stapler, cellophane tape, pen) • Manipulation knowledge – which two items are most similar in manner of manipulation (e.g., piano, typewriter, stove)

21. Motor Cognition • Conceptual knowledge • Results apraxic patients were more impaired in manipulation test, but than function test • Kellenbach et al. (2003) used PET to investigate brain activation associated with function and manipulation judgments • Results showed that when participants made conceptual judgments about function, there was activation of a left network consisting of the ventral premotor cortex and the posterior middle temporal gyrus • When participants made manipulation judgment an additional region, and additional region, the intraparietal sulcus, was activated

22. Motor Cognition • Conceptual knowledge The intraparietal sulcus plays an important role in skilled object use (Heilman, 1993)

23. Motor Cognition • Conceptual knowledge Visual-gestural knowledge Beauchamp (2002) in neuroimaging study showed that bilateral regions of the middle temporal cortex were activated when tool motion (i.e., gestural motion) was viewed in comparison to human motion (i.e., person jogging on the spot)

24. Motor Cognition • Conceptual knowledge Beauchamp (2003) in neuroimaging study showed that middle temporal gyrus responded more strongly to tool motion videos and point-light displays of tool motion

25. Motor Cognition • Point-light displays (aside) • Animals and humans move in ways that are distinctive and different from the way in which nonhumans move. These patterns of movement are called biological motion • Johansson (1973) developed the ‘point-light display’ technique to investigate movement. • Small light sources attached wrists, knees, ankles, shoulders, and heads of actors who performed various movements (e.g., walking, running, dancing)

26. Motor Cognition • Conceptual knowledge Park and Roy (in prep) showed that patients with LHD, but not RHD were strongly impaired on function and manipulation tests but that patients with LHD and RHD were impaired on tests of visual-gestural knowledge

27. Motor Cognition • Conclusions • These studies suggest that different types of conceptual knowledge associated with action are dissociable from each other • Three types of knowledge have been studied in depth. These are knowledge of: • Function • Manipulation • Visual-gestural knowledge of tool motion

28. Motor Cognition • Imitation in this model can be accomplished in two different ways • 1. directly from perception to action; and • 2. indirectly through long-term memory • Evidence to support this comes from studies which have shown: • The general observation that meaningless actions can be imitated accurately in cognitively unimpaired individuals • Findings of dissociation between imitation and pantomime (e.g., Goldenberg & Hagmann, 1997; Ochipa et al., 1994); stronger lateralization to pantomime than to imitation • (interpret on basis of model)

29. Motor Cognition • What is acquired when we view purposeful action • It would appear that we derive the goal of the action • (e.g., see a person reaching hand across table, grasping a mug of coffee, and moving is toward lips would be described as …drinking a cup of coffee. In other words viewed actions tend to be described in terms of the goal of the action

30. Motor Cognition Imitation • Imitation -- ability to understand the intent of a viewed action and then to reproduce it • This needs to be distinguished from mimicry, which is reproduction of a behavior without understanding (e.g., a parrot mimics human speech) • Meltzoff and Moore (1977) showed that newborn infants can imitate viewed action (sticking out tongue; opening mouth etc.) • By 6 months of age infants can imitate actions on objects (e.g., shaking a rattle)

31. Motor Cognition Imitation • As infants grow older deferred imitation abilities increase (i.e., memory for imitated action) • data show that infants as young as 18 months appear to represent intentions of actions not just the action itself • E.g., children who watched an actor try to pull apart a dumbbell but failed, were more likely to try and pull apart the dumbbell than if they watched a mechanical device attempt to pull apart a dumbbell

32. Motor Cognition • Components of imitation • Decety et al. (1997) in neuroimaging studies compared brain activation of subjects as they viewed meaningless actions either intending to recognize or to imitate the viewed action • Additional brain regions activated when intending imitate meaningless actions: supplementary motor area (SMA), the middle frontal gyrus, the premotor cortex, the anterior cingulate, and the superior and inferior parietal cortices in both hemispheres.

33. Motor Cognition • Conclusion: intentions (top-down) processes of participant influenced observation of action. Regions activated during observation also are the ones involved in action generation. • Observing an action with the intention to perform that action involves regions similar to those used to generate the action • when intending to recognize an action activated regions were the memory encoding structures (the parahippocampal gyrus)

34. Motor Cognition • When actions are viewed — separating intention from means – a neuroimaging perspective • Several people have proposed that actions are often understood in terms of the intentions (goals) they achieve and the means used (movements) to achieve these goals (e.g., Heider) • Chaminade (2002) investigated using neuroimaging the neural regions associated with goals and means

35. Motor Cognition • When actions are viewed — separating intention from means – a neuroimaging perspective • Participants saw an actor make Lego constructions: participants viewed the goal alone (hand moving away from block in specified location); the means alone (the hand grasping and moving the block); or the entire action. All participants imitated action just observed

36. Motor Cognition • When actions are viewed — separating intention from means – a neuroimaging perspective • Findings– when participants imitated action or means, the medial prefrontal cortex was activated; this region appears to play a critical role in inferring other people’s intentions • When participants imitated goal the left premotor cortex activated • Conclusion • In normally functioning adults imitating a gesture activates neural regions associated with the intentions underlying the action

37. Motor Cognition Mental simulation • Since imagery and perception activate similar brain regions it seems reasonable to hypothesize that one way to reason is to simulate (or imagine) the consequence of performing a planned action

38. Motor Cognition • Simulation theories of action understanding • It has been proposed that actions of others are understood by putting yourself in their place (either by observation or imagination) • This permits you to derive their intentions and generate an action plan (but how can you do this since intentions and actions are internal and unobservable?)

39. Motor Cognition Mental simulation • It has been shown that practicing with mental imagery can help participants in their performance of the task • It has been shown that mental imagery practice has positive effects on complex motor skill learning (e.g., putting a golf ball) • Yue & Cole (1992) showed compared finger strength of two groups– group 1 performed repeated isometric exercises; group 2 received motor imagery instruction and imagined making finger movements without actually making them • Both groups had increased finger strength– group 1 by 30% and group 2 by 22% • Conclusion– possible to increase strength without actually repeated muscle activation

40. Motor Cognition Mental simulation • It has also been shown that viewing an action can facilitate enactment of that action • Priming refers to the facilitation of processing by previous performance of a task • Motor priming refers to the facilitative effect that watching a movement or action has on making a similar motor response. Motor priming has been observed in a variety of experimental situations

41. Motor Cognition Mental simulation • fMRI studies have shown that the neural difference between motor imagery and motor performance is not a matter of “what”, but “how much” • i.e., similar brain regions are activated, but the level of activation is significantly lower; in one study imagery activation was 30% of that found in actual execution (Roth, 1996)

42. Motor Cognition Mental simulation • Ruby & Decety (2001) Nature Neuroscience investigated the question of agency • Background—if viewing an action activates regions involved in performing an action, how do people distinguish actions they perform vs those they observe (i.e., attribution of action to self or another agent) • Previous studies have shown that the first-person perspective (imagining oneself) is associated with activation of inferior parietal, premotor, and SMA on left side

43. Motor Cognition Mental simulation • This study asked what areas are activated when we imagine not ourselves performing an action, but another person performing that action • Method • Participants were scanned while they simulated everyday actions (e.g., winding a watch) with right hand (all Ps were right handed) • Ps instructed to imagine themselves perform the action or to imagine another person performing the action • Perspectives initiated by presenting a photo or from a spoken sentence describing the action

44. Motor Cognition Conclusions • First person perspective versus imaging another person acting was associated with activation of common neural resources • Consistent with notion that a common code is used to perceive, imagine, and produce actions • However, specific regions are activated when imagining oneself performing an action versus another person. These regions may be used to determine agency

45. Motor Cognition Mental simulation • Results • Both self perspective and other perspective activated common regions – supplementary motor area (SMA), premotor cortex, precuneus (an area located in parietal lobe), and occipital-temporal lobe • However, when compared to the first-person perspective, the third-person perspective selectively activated the frontopolar cortex, the precuneus, and the right inferior portion of the parietal lobe • See figure in next slide

46. Ruby & Decety (2001)

47. Motor Cognition & Memory • Memory for action • Subject-performed task (SPT) paradigm requires participant(P) to perform actions according to verbal instructions given by experimenter (e.g., roll the ball, fold the paper, lift the pen) at study • At test P’s memory for these actions is tested • Control condition– P hears instructions but does not perform actions • Result—memory for enacted action phrases is superior to that for events encoded without enactment Presentation relies on Nilsson (2000) In Craik and Tulving Oxford Handbook of memory

48. Motor Cognition & Memory • Memory for action--theories • Non-strategic encoding theory of Cohen • This theory proposed that enacted actions are encoded nonstrategically unlike verbal and other types of events

49. Motor Cognition & Memory • Memory for action--theories • Multimodal theory of Backman and Nilsson (1984, 1985) • Enactment during encoding automatically leads to multimodal processing, which produces a rich encoding of information (multimodal because there is auditory, visual, and haptic input) in SPT condition • subsequently proposed that physical (perceptual) properties were encoded nonstrategically, whereas verbal components were encoded strategically. • SPTs contained verbal and physical properties whereas VTs contained verbal component only (Backman et al. 1986)

50. Motor Cognition & Memory • Memory for action--theories • (Backman et al. 1989) proposed that the physical component of the dual code is encoded incidentally and retrieved implicitly, whereas verbal component is encoded intentionally and retrieved explicitly