, 2015 http://dx.doi.org/10.1080/09658211.2015.1059858

How motor practice shapes memory: retrieval but not extra study can cause

Tobias Tempel and Christian Frings

Fachbereich I – Psychologie, University of Trier, Trier, Germany

(Received 5 September 2014; accepted 2 June 2015)

We investigated the retrieval specificity of retrieval-induced forgetting (RIF) of motor sequences. In two experiments, participants learned sequential finger movements, each consisting of the movement of two fingers of either the left or the right hand. In the learning phase, these motor sequences were graphically presented and were to be learned as responses to simultaneously presented letter stimuli. Subsequently, participants selectively practiced half the items of one hand. A final test then assessed memory for all initially learned items. We contrasted different kinds of selective practice with each other. Whereas retrieval practice required retrieving motor sequences in response to letter stimuli from the learning phase, extra study was an extension of the learning phase, that is, participants performed motor sequences in response to the same animation graphic display as in the learning phase again accompanied by the letter stimulus. All practice conditions strengthened the practiced items, but only retrieval practice resulted in RIF. Thus, the strengthening of items through practice did not suffice to induce forgetting of related motor sequences. Retrieval was a necessary component for practice to shape memory for body movements by impairing the subsequent recall of motor sequences that were related to the practiced motor sequences.

Keywords: Retrieval-induced-forgetting; Body movement; Recall.

Doing sports or playing a musical instrument 1992). Here, we focus on comparing the effects of requires practice—the way to mastery involves practice with and without retrieval. thorough exercises. Repeating the same body Practice seems to be especially effective if it movements over and over again aims at the acquire- involves memory retrieval. The testing effect is ment of fluent motor sequences that can quickly be the phenomenon that retrieval improves memory retrieved from memory. There are various ways of for the retrieved information more than mere rep- practicing body movements, of course. Exercises etition (Allen, Mahler & Estes, 1969; Bjork, 1975; Downloaded by [Central Michigan University] at 22:38 06 November 2015 may focus on exact repetition of movement patterns Roediger & Karpicke, 2006). Whereas most or on the generalisation and transfer of skills. Feed- studies concerned with the benefits of retrieval back may be given permanently or sparsely, relied on verbal material, recent studies also expli- immediately or after a delay. The investigation of citly focused on testing effects in motor action. For such variables suggests that comparing the effi- example, Boutin, Panzer, and Blandin (2013) ciency of different training techniques depends on demonstrated that testing improved the generalis- the outcome of interest with some benefiting pri- ation of a dynamic arm movement skill (see marily the speed of acquisition whereas others also, Boutin, Panzer, Salesse, & Blandin, 2012). rather benefit retention (cf. Schmidt & Bjork, Kromann, Jensen, and Ringsted (2009) showed

Address correspondence to: Tobias Tempel Fachbereich I – Psychologie, University of Trier, 54286 Trier, Germany. E-mail: [email protected]

© 2015 Taylor & Francis 2 TEMPEL AND FRINGS

that testing at the end of a resuscitation course for we found that RIF of motor sequences also medical students enhanced the retention of resus- occurs in indirect memory tests that required the citation skills. execution of movements overlapping with the However, practice has side effects. Whereas learned items only in the respective motor pro- practice is beneficial for the memory of the prac- grammes (Tempel & Frings, 2014b, 2015), ticed material, it can impair related . suggesting that the processes underlying RIF Retrieval-induced forgetting (RIF) is the phenom- affected motor programmes. In those indirect enon that selectively retrieving information from a tests, participants had to execute the same motor set of information stored in memory can cause the sequences as in the learning phase while entering forgetting of non-retrieved information from this novel stimuli via the keyboard. The performance set. RIF is analysed in the retrieval-practice para- in this task overlapped with the preceding tasks digm (Anderson, Bjork, & Bjork, 1994) that con- of learning and retrieval-practice trials solely sists of three main phases. In the learning phase, regarding the to-be-executed movements. participants study several sets of items in combi- However, it is not clear so far whether these pro- nation with a shared (category-)cue that defines cesses depended on memory retrieval or if other the specific set of items (e.g., flower—rose, kinds of motor practice not involving retrieval flower—tulip, fruit—banana, fruit—cherry). In equally would entail forgetting. The previous the subsequent retrieval-practice phase, partici- experiments were not able to demonstrate pants are cued to recall half of the studied items whether retrieval attempts were necessary for from half of the sets (e.g., flower—ro__). In the inducing forgetting because the effects of retrieval test phase then, recall performance for all items practice were never compared with the effects of is tested. The recall of practiced items (Rp+ retrieval-free practice. Perhaps, the term RIF items; e.g., rose) and unpracticed items from prac- was in reality inappropriate since Rp– items ticed sets (Rp− items; e.g., tulip) is compared to were impaired because the strengthened Rp+ the recall of unpracticed items from unpracticed items interfered at test blocking access to Rp− sets (Nrp items; e.g., banana, cherry). Rp+ items items. Therefore, we here scrutinised if selective typically profit from retrieval practice and are motor practice also entails similar forgetting better recalled than Nrp items. RIF manifests effects when it does not involve memory retrieval. itself in significantly worse recall of Rp− items as The contribution of retrieval to practicing body compared to Nrp items. movements only seldom has been scrutinised, Recently, the retrieval-practice paradigm has although it has been suggested that retrieval is a been adapted for investigating motor memory crucial element in many training techniques (cf. (Tempel & Frings, 2013). Participants first Schmidt & Bjork, 1992). Athletes or musicians learned 12 sequential finger movements (SFM), do not solely rely on restudying but include the each consisting of 2 consecutive key presses in retrieval of motor sequences as a natural com- response to a letter stimulus. Six items involved ponent in their exercises. Therefore, it is important two fingers of the left hand, six involved two to know whether retrieval practice entails specific fingers of the right hand (e.g., left ring finger, left consequences that do not follow on other retrie-

Downloaded by [Central Michigan University] at 22:38 06 November 2015 index finger). After retrieval practice on three val-free ways of practice. With regard to verbal items of one hand, a final recall test assessed material, it has been shown that not every kind memory for all items. Rp+ and Rp− items per- of practice induces forgetting of related infor- tained to one hand, and Nrp items pertained to mation. RIF has been shown to be retrieval- the other hand. Rp− items were tested before specific in several studies comparing selective Rp+ items in order to preclude any output inter- retrieval practice with selective extra study ference by Rp+ items (cf. Anderson et al., 1994; (Bäuml, 2002; Ciranni & Shimamura, 1999; Stau- Roediger & Schmidt, 1980). Rp− items were digl, Hanslmayr, & Bäuml, 2010; for a meta-analy- accordingly compared to the first three Nrp sis see: Murayama, Miyatsu, Buchli, & Storm, items tested (Nrp1), whereas Rp+ items were com- 2014). Only retrieval practice impaired memory pared to the last three Nrp items tested (Nrp2). for related items. In contrast, extra study instead Significantly fewer Rp− than Nrp1 items were of retrieval practice did not induce forgetting of recalled. Also, response times (RTs) to Rp− related items. This finding suggests that RIF is items were significantly longer than RTs to Nrp1 not merely a product of interference at test items (see also Reppa, Worth, Greville, & Saun- arising from the strengthened Rp+ items. ders, 2013; Tempel & Frings, 2014a). In addition, However, properties of phenomena in verbal HOW MOTOR PRACTICE SHAPES MEMORY 3

memory cannot be readily transferred to motor EXPERIMENT 1 memory. Since different systems are likely involved in and processing verbal In Experiment 1, we contrasted two conditions. In versus motor content (e.g., Krakauer & Shadmehr, both conditions, participants first learned 12 SFM 2006), processes of interference and inhibition as responses to letters. During a learning trial a may operate differently as well. In general, letter appeared at the centre of the computer current memory research unnecessarily neglects screen together with an animation graphic display motor action. A notable exception is work by of the corresponding two-finger-movement. Anderson (2003) who explicitly linked inhibition Immediately after this animation, participants exe- in memory with behavioural inhibition in motor cuted the just displayed SFM by pressing corre- action, both achieving a kind of response override sponding keys. Six items were executed with (see also Anderson & Weaver, 2009). Yet, empiri- fingers of the left and six with fingers of the right cal investigations of retrieval dynamics in motor hand. One experimental condition then involved memory are scarce. Hence, more studies are selective retrieval practice of items. Participants needed examining in which regard motor had to retrieve three items of one hand in response memory operates in similar or disparate ways. to their letter stimuli. A letter appeared on the Retrieval specificity is one of the key predic- screen and the participant entered the correspond- tions of the inhibitory account of RIF (Anderson, ing item while his or her fingers rested on the same 2003; Anderson et al., 1994; Anderson & Spell- keys as in the learning phase. We expected to repli- man, 1995; Storm & Levy, 2012; Wimber, Alink, cate an RIF effect (Tempel & Frings, 2013). The Charest, Kriegeskorte, & Anderson, 2015). This selective retrieval practice of three items of one theory assumes that during retrieval practice of hand (Rp+) should induce forgetting of the three one item the other items of this set interfere in a non-retrieved items of this hand (Rp−)ascom- competition for conscious recollection. The retrie- pared to items of the other hand (Nrp). The other val attempt is necessary for this competition to condition involved selective extra study. Partici- arise. In order to resolve this interference, the pants selectively practiced the same items as in Rp+ items are strengthened while simultaneously the first condition, but, instead of retrieval practice, the Rp− items are inhibited. Translated to motor they received additional learning trials for three sequences, the inhibitory account implies that inhi- items of one hand by means of the same format bition resolves interference arising between motor as in the learning phase, that is, they performed programmes because RIF has been shown to the SFM in response to the same animation occur in indirect memory tests overlapping with graphic display as before accompanied by the cor- motor sequences acquired in the learning phase responding letter. The participants’ task in the only in the to-be-executed movements (Tempel extra study phase, hence, was identical to the pre- & Frings, 2014b, 2015). In contrast, extra study ceding learning trials. They entered the displayed might not affect motor programmes because no SFM and were instructed to memorise it in interference between motor programmes may response to the simultaneously presented letter arise during extra study. stimulus. Thus, participants in both conditions

Downloaded by [Central Michigan University] at 22:38 06 November 2015 We investigated if selective practice of a subset selectively practiced a subset of items, but the of body movements stored in memory also has to kind of practice differed by only involving retrieval involve retrieval for inducing forgetting of non- in one condition. The final test phase was identical practiced movements of the same set. Thus, we in both conditions. We expected no RIF to occur in tested whether retrieval dynamics that have been the extra study condition. In contrast, Rp+ proven to work in declarative memory affect enhancement should occur in both conditions. motor memory in the same way. Perhaps, motor practice without retrieval suffices to impair later recall of related non-practiced movements. Selec- Method tively executing some movements repeatedly might strengthen these movements in a way that Participants they block access to related movements. To this end, we compared practice without retrieval Sixty-four psychology students at the Univer- versus retrieval practice and therefore were able sity of Trier (53 women, mean age = 21.6, SD = to analyse for the first time whether retrieval at 4.9) participated in the experiment. They received practice is a precondition for RIF in motor memory. course credit for their participation. 4 TEMPEL AND FRINGS

Design items appeared once in random order, whereupon an instruction screen informed the participant that The study had a 2 (retrieval practice, extra the items just presented would appear again − study) × 4 (item type: Rp+, Rp , Nrp1, Nrp2) several times in the following. Eleven blocks mixed design with repeated measures on the last each containing the 12 items in a random order factor. were presented. The selective practice phase differed between Material the two experimental conditions. In the retrieval- practice condition, participants had to retrieve The experiment was conducted using Dell Opti- three items of one hand. The items were cued by plex 755 PCs with Eizo FlexScan S1901 monitors their letter stimuli. A letter appeared on the and standard German QWERTZ keyboards. The screen and the participant entered the correspond- software PXLab (Irtel, 2007) served for running ing item while his or her fingers again rested on the the experiment. The items consisted of 12 two- marked keys (cf. Figure 1, middle section). No finger movements. Six two-finger movements con- feedback about the accuracy of the performed sisted of fingers of the left, the other six of fingers two-finger-movement was given. In the extra of the right hand. Participants learned these move- study condition, participants received additional ments as responses to letters. The letters a to f learning trials for three items of one hand by were the stimuli for the left‐hand movements, means of the same format as in the preceding the letters u to z for the right-hand movements phase, that is, they performed the SFM in response (see Appendix A). During the learning phase a to the same animation graphic display as letter appeared at the centre of the computer before accompanied by the corresponding letter. screen together with an animation of the corre- Thus, the participants’ task in the extra study sponding two-finger-movement (cf. Figure 1). phase was identical to the preceding learning The fingers of the left hand were placed on the trials. They entered the displayed SFM and were keys Q, W, E, and R. The fingers of the right instructed to memorise it in response to the hand were placed on the keys U, I, O,andP. simultaneously presented letter stimulus. Below the letter a display of the two hands demon- Whereas instructions in the retrieval-practice con- strated which fingers should be moved by showing dition explained that participants had to retrieve two consecutively flashing fingers (first finger was items of either the left or the right hand in coloured yellow, second finger was coloured blue, response to their letter stimuli, instructions in 200 ms per flash). After the display of the two the extra study condition stated that additional hands disappeared participants could perform learning trials for items of either the left or the movement. If the performed sequence was right hand would follow. In both conditions, coun- incorrect a feedback appeared, displaying: Fehler! terbalancing of the practiced items resulted in four (English: Error!). practice sets (abc, def, uvw, and xyz). These four sets were counterbalanced between participants. Procedure The items were practiced five times in both

Downloaded by [Central Michigan University] at 22:38 06 November 2015 conditions. The experiment consisted of four phases (learn- As typical for the retrieval-practice paradigm, ing, selective practice, distractor task, final recall). we increased the retention interval by running a Instructions were given on the screen and summar- distractor phase. The distractor phase comprised ised by an experimenter. After having read the an aesthetic judgement task. Using a mouse the instructions for the learning phase the participant participants disclosed abstract paintings on the clicked an on-screen button in order to start with computer screen by clicking on-screen buttons, the learning phase. Participants had 10 seconds and subsequently rated how appealing these paint- to place their fingers with exception of the ings were to them. Sixteen paintings were rated. thumbs on eight keys (Q, W, E, R, U, I, O, and The mean time taken for this task was 3.2 P) marked by labels hiding the inscription. Then, minutes (SD = 0.5). the learning of the 12 items began. Participants The final test phase presented participants with had to consecutively press two keys in response the 12 letter stimuli in the same fashion as during to a displayed letter accompanied by an animation the selective practice phase in the retrieval-prac- graphic display showing which digits should be tice condition (cf. Figure 1, lower section). The moved (cf. Figure 1, upper section). First, all 12 items of one hand were presented in succession, HOW MOTOR PRACTICE SHAPES MEMORY 5

Figure 1. The upper section depicts a trial in the learning phase. It starts with a display of the letter stimulus together with a drawing of the two hands. After 400 ms the first finger illuminates yellow for 200 ms and then the second finger illuminates blue for 200 ms. Downloaded by [Central Michigan University] at 22:38 06 November 2015 Subsequently, the hand display disappears and the participant can enter the SFM just illustrated. In the selective retrieval-practice, a stimulus is given and the participant is supposed to perform the corresponding SFM without receiving feedback about his or her response. In contrast, extra study trials required the execution of SFM in response to the same animation as in the learning phase. The lower section depicts a trial in the final recall phase. A stimulus is given and the participant is supposed to perform the correspond- ing SFM. SFM, sequential finger movement.

that is, the test was blocked by hand. The test one of two sequences thereby counterbalancing either started with the left or the right hand. It which items constituted Nrp1 items and Nrp2 was counterbalanced if the test sequence started items. with the practiced or with the non-practiced hand. Within the items of the practiced hand the Rp− items were tested before the Rp+ items in Results order to avoid effects of output interference (cf. Anderson et al., 1994). Within the items of the Mean retrieval success in the selective practice non-practiced hand the items were presented in phase of the retrieval-practice condition was 6 TEMPEL AND FRINGS

49% (SD = 23%). For analyses of RTs, we Discussion excluded outliers exceeding three SD above the mean. RIF occurred as a consequence of selective retrie- val practice, whereas selectively practicing SFM RIF without retrieval did not induce forgetting of related SFM. Still, both kinds of practice strength- A 2 (condition: retrieval practice, extra ened Rp+ items. Indeed, the facilitation for Rp+ study) × 2 (item type: Rp−, Nrp1) ANOVA exam- items did not occur with regard to the number of ined RIF. With regard to RTs, the main effect recalled items but only for RTs. Therefore, the of item type was not significant, F(1, 62) = 3.16, strengthening might seem somewhat weak. p = .080, whereas the main effect of condition However, it has been shown that RIF also occurs approached significance, F(1, 62) = 3.90, p = .053. with relatively weak Rp+ enhancement, as well More importantly, the interaction was significant, as that RIF even occurs with impossible retrie- h2 F(1, 62) = 6.47, p = .014, p = .09. Separate t-tests val-practice trials, that is, in the absence of any per condition showed that RTs for Rp– items strengthening (Storm, Bjork, Bjork, & Nestojko, were significantly longer than RTs for Nrp1 2006; Storm & Nestojko, 2010). Moreover, RIF items in the retrieval-practice condition, t(31) = also occurred in RTs but not with regard to the 2.87, p = .007, dz = 0.49, but RTs for Rp– and number of recalled items in the present exper- Nrp1 items did not significantly differ in the iment. Hence, the results show that strengthening extra study condition, t(31) = 0.58, p = .566 Rp+ items does not suffice to induce forgetting of (Figure 2). With regard to the number of recalled related motor sequences. However, it is possible items, the main effect of item type was not signifi- that the extra study condition did not induce for- cant, F(1, 62) = 2.42, p = .125, neither was the getting because it did not affect motor pro- main effect of condition, F(1, 62) = 1.98, p = .165. grammes. Previous investigations found evidence There was a marginal interaction, F(1, 62) = 3.38, for retrieval practice to affect motor programmes p = .071. (Tempel & Frings, 2014b, 2015). If RIF is a conse- quence of interference during retrieval practice, Rp+ enhancement extra study might not have induced forgetting because competition between motor programmes A 2 (condition: retrieval practice, extra was less strong than during retrieval practice. study)×2 (item type: Rp+, Nrp2) ANOVA exam- Perhaps, extra study did not suffice to entail com- ined Rp+ enhancement. With regard to RTs, the petition between motor programmes of practiced main effect of item type was significant, F(1, 62) and non-practiced items during practice or at test h2 = 17.51, p < .001, p = .22, whereas the main causing forgetting. Therefore, we examined effect of condition was not significant, F(1, 62) = whether a different kind of practice focusing 0.06, p = .801. The interaction was not significant more on the motor programme would not entail either, F(1, 62) = 1.98, p = .164. With regard to RIF either but only strengthen practiced items in the number of recalled items, the main effect of a second experiment.

Downloaded by [Central Michigan University] at 22:38 06 November 2015 item type was not significant, F(1, 62) = 1.12, p = .293, neither was the main effect of condition, F(1, 62) = 2.94, p = .092, nor the interaction, F(1, EXPERIMENT 2 62) = 1.68, p = .200. Again, retrieval practice was contrasted with extra Correlation of RIF and Rp+ enhancement study. However, the kind of study was changed. Instead of only entering a SFM once in response For additional analyses, we computed RIF as to the corresponding animation, participants had the difference in RTs for Rp– items and Nrp1 to immediately repeat the execution of SFM 10 items (Rp–—Nrp1) and Rp+ enhancement as times in response to its animation. Thus, the prac- the difference in RTs for Nrp2 and Rp+ items tice focused especially on executing the motor (Nrp2 — Rp+). There was no correlation response. Moreover, there were two extra study between RIF and Rp+ enhancement in the retrie- conditions comprising a different amount of prac- val-practice condition (r = −.081, p = .661), nor tice cycles. A further change pertained to the item between the corresponding measures in the extra material. RIF and Rp+ enhancement occurred in study condition (r = −.055, p = .765). RTs in Experiment 1, but they did not with HOW MOTOR PRACTICE SHAPES MEMORY 7

Figure 2. The upper section shows the percentages of items recalled in the final test phase of Experiment 1 as a function of exper- imental condition and item type. The lower section shows mean response time (in ms) in the final recall test. Error bars depict standard error of the mean. Rp+, practiced items; Rp−, unpracticed items of the practiced hand; Nrp1, RIF baseline; Nrp2, Rp+ enhancement baseline.

regard to the number of recalled items. Although, Method we had found previously that RTs are, in fact, a more reliable dependent variable (Tempel & Participants Frings, 2013) to assess both effects, we changed to items that might be suited better for showing Ninety-six psychology students at the Univer- RIF and Rp+ enhancement also in the number sity of Trier (75 women, mean age = 22.5, SD = of recalled items. For this purpose, we turned to 2.8) participated in the experiment. They received material used in Experiment 3 of the study by course credit for their participation. Tempel and Frings (2013) where both effects occurred with regard to the number of recalled Design items. Again, the items were six SFM with fingers of the right and six SFM with fingers of Participants were randomly assigned to one of the left hand. The hands served as categories, three conditions. The retrieval-practice condition

Downloaded by [Central Michigan University] at 22:38 06 November 2015 that is, the three Rp+ items and the three Rp– involved the retrieval of Rp+ items during the items pertained to the same hand, whereas Nrp1 selective practice phase, whereas the two extra and Nrp2 items pertained to the other hand. The study conditions involved selectively practicing three SFM of one item type all started with the Rp+ items by entering the SFM in response to same finger. This regularity may allow for retrieval the corresponding animation 10 times in a row. dynamics operating within sets of items to have a The short study condition comprised two practice stronger impact also on the successful recall and cycles. The long study condition comprised four not only affect retrieval latency. Confining all practice cycles. Item type (Rp–, Rp+, Nrp1, items of one item type to the same starting finger Nrp2) was manipulated within subjects. probably reduces inter-item integration between item types which is known to counteract the Material and procedure dynamics causing RIF (Anderson, Green, & McCulloch, 2000; Anderson & McCulloch, 1999). The items again were SFM. However, we used We expected to replicate the finding of Exper- different item sets. For both hands, all two-finger iment 1 that only retrieval practice would impair movements started with one out of two fingers. subsequent recall of Rp– items. Combining these starting fingers with all three 8 TEMPEL AND FRINGS

remaining fingers resulted in the six items used for number of recalled items, the main effect of item each hand. We counterbalanced if the starting type was not significant, F(1, 93) = 1.39, p = .242, fingers were the index fingers and the ring neither was the main effect of condition, F(2, 93) fingers or the middle fingers and the pinkies (see = 0.96, p = .386. The interaction was significant, F h2 Appendix B). A further difference pertained to (2, 93) = 4.50, p = .014, p = .09 (Figure 3). Separ- the keys used for input. The fingers of the right ate t-tests per condition showed that significantly hand were placed on the keys 7, 8, and 9, and + fewer Rp– than Nrp1 items were recalled in the of the numeric keypad on the right side of the key- retrieval-practice condition, t(31) = 3.63, p = .001, board, instead of the keys U, I, O, and P. The pro- dz = 0.64, but there was no significant difference cedure was identical to Experiment 1, with the between Rp– and Nrp1 items in the short study exception of the practice phase in the extra study condition, t(31) = 1.14, p = .263, or in the long conditions. Participants had to execute a SFM 10 study condition, t =0. times in a row in response to its corresponding letter accompanied by the same animation Rp+ enhancement graphic display as in the learning phase. First, the animation graphic depicting the SFM appeared A 3 (condition: retrieval practice, short extra together with the corresponding letter stimulus study, long extra study)×2 (item type: Rp+, in the same fashion as in the learning phase. Nrp2) ANOVA examined Rp+ enhancement. Then, the participants had to execute the just dis- With regard to RTs, the main effect of item type h2 played SFM 10 times. After entering the SFM 10 was significant, F(1, 93) = 11.98, p = .001, p = .11, times, a next SFM was practiced. Instructions whereas the main effect of condition was not, F additionally emphasised that participants were (2, 93) = 1.80, p = .172. The interaction was not sig- supposed to use this extra study opportunity to nificant either, F(2, 93) = 1.68, p = .192. RTs for Rp memorise the practiced items especially good for + items were shorter than RTs for Nrp2 items. the later test. Whereas the short study condition With regard to the number of recalled items, the comprised two practice cycles, the long practice main effect of item type was significant, F(1, 93) h2 condition comprised four cycles. = 5.39, p = .022, p = .06, whereas the main effect of condition was not, F(2, 93) = 1.63, p = .202. The interaction was not significant either, F(1, Results 93) = 0.82, p = .443. More Rp+ than Nrp2 items were recalled. Mean retrieval success in the selective practice phase of the retrieval-practice condition was Correlation of RIF and Rp+ enhancement 57% (SD = 29%). For analyses of RTs, we excluded outliers exceeding three SD above the We computed two measures each for RTs and mean. for the number of recalled items. RIF was com- puted as the difference in RTs for Rp– and Nrp1 –– RIF items (Rp Nrp1), as well as the difference in

Downloaded by [Central Michigan University] at 22:38 06 November 2015 the number of recalled Nrp1 and Rp– items A 3 (condition: retrieval practice, short extra (Nrp1 – Rp–). Rp+ was computed as the difference study, long extra study)×2 (item type: Rp–, in RTs for Nrp2 and Rp+ items (Nrp2 – Rp+), as Nrp1) ANOVA examined RIF. With regard to well as the difference in the number of recalled RTs, neither the main effect of item type was sig- Rp+ and Nrp2 items (Rp+ – Nrp2). With regard nificant, F(1, 93) = 1.28, p = .261, nor the main to RTs, there was no significant correlation effect of condition, F(1, 93) = 0.60, p = .551, between RIF and Rp+ enhancement in the retrie- whereas the interaction was significant, F(2, 93) val-practice condition (r =.071, p =.701), nor in h2 = 3.46, p = .036, p = .07. Separate t-tests per con- the short study condition (r = .330, p = .065), or in dition showed that RTs for Rp– items were signifi- the long study condition (r =.254, p = .160). With cantly longer than RTs for Nrp1 items in the regard to the number of recalled items, there was retrieval-practice condition, t(31) = 2.82, p = .008, no significant correlation between RIF and Rp+ dz = 0.50, but RTs for Rp– and Nrp1 items did enhancement either in the retrieval-practice con- not significantly differ in the short study condition, dition (r = −.174, p = .341), nor in the short study t(31) = 0.42, p = .680, or in the long study con- condition (r = −.044, p = .813), or in the long study dition, t(31) = 0.43, p = .671. With regard to the condition (r = .025, p = .891). HOW MOTOR PRACTICE SHAPES MEMORY 9

Figure 3. The upper section shows the percentages of items recalled in the final test phase of Experiment 2 as a function of exper- imental condition and item type. The lower section shows mean response time (in ms) in the final recall test. Error bars depict standard error of the mean. Rp+, practiced items; Rp−, unpracticed items of the practiced hand; Nrp1, RIF baseline; Nrp2, Rp+ enhancement baseline.

Discussion practiced half of the items of one hand. After a dis- tractor task, finally, memory for all initially RIF again only occurred in the retrieval-practice learned SFM was assessed. Different variants of condition. Neither the short nor long extra study selective practice were compared. Experiment 1 condition impaired the recall of items belonging contrasted retrieval practice with extra study by to the same hand as the practiced items. This means of the same format as in the preceding result replicates the finding of Experiment learning phase, whereas Experiment 2 contrasted 1. With the use of other item material, RIF and retrieval practice with extra study that focused Rp+ enhancement also occurred with regard to especially on practicing motor programmes. All the number of recalled items and not only in practice conditions strengthened the practiced RTs. We assume that confining all SFM of one items, but only retrieval practice resulted in RIF. item type to start with the same finger reduced The present findings prove the power of retrieval

Downloaded by [Central Michigan University] at 22:38 06 November 2015 inter-item integration and, moreover, might have in shaping motor memory. When motor practice allowed for facilitation of Rp+ items to occur does not involve memory retrieval, it does not also with regard to the number of recalled items negatively affect later access to body movements because this regularity might have permitted a that are related to the practiced movements. relatively stronger strengthening of the association Thus, retrieval is essential to trigger dynamics of motor sequences to their respective letter causing RIF. These dynamics may involve inhi- stimuli. bition. Retrieval specificity is one of the key pre- dictions of the inhibitory account of RIF that assumes that during retrieval practice of one GENERAL DISCUSSION item the other items of this set interfere in a com- petition for conscious recollection. In order to Two experiments compared the effects of selective resolve this interference, the Rp+ items are retrieval practice and extra study of motor strengthened while simultaneously the Rp– items sequences. Participants first learned SFM. Half are inhibited. An alternative model assumes that of these pertained to the left and the other half Rp+ items block Rp– items, that is, recall of Rp– to the right hand. Subsequently, participants suffers from stronger interference in the test 10 TEMPEL AND FRINGS

phase than Nrp items (Raaijmakers & Jakab, preventing attempts to retrieve SFM from long- 2013). However, blocking cannot explain the term memory via their associated letter stimuli. present results because extra study strengthened Most importantly, of course, the significant differ- practiced items as did retrieval practice. This ences between the retrieval-practice conditions strengthening occurred in the absence of forget- and the extra study conditions indicate that extra ting of SFM of the same hand. Hence, the prac- study did not represent an equivalent retrieval ticed items did not block access to the non- condition, even though it is not possible to practiced items. In Experiment 1, there was even preclude rudimentary retrieval attempts with a tendency for more Rp– items to be recalled com- certainty. pared to Nrp1 items. In addition, there was no sig- Although we found that RIF of motor nificant correlation between Rp+ enhancement sequences is retrieval-specific, as it has been and RIF. found to be with regard to other learning material, Recently, Jonker, Seli, and MacLeod (2013) it also depends on peculiar characteristics of motor challenged the inhibitory account by demonstrat- memory. For example, we have previously found ing that also a context change between a selective RIF of motor sequences to occur in tests indirectly extra study phase and a subsequent test phase can testing memory by requiring the execution of induce RIF-like effects. Therefore, they argue that motor sequences in response to novel test cues the assumption of shifts in mental context occur- (Tempel & Frings, 2014b, 2015). This finding ring automatically when participants proceed shows that RIF affected motor programmes repre- from the learning phase to the retrieval-practice senting sequences in a format closely correspond- phase could account for most RIF effects reported ing to parameters of movement execution. In in the literature, whereas the non-occurrence of addition, the occurrence of RIF required the corresponding forgetting effects subsequent to organised into memory sets according to selective extra study resulted from the absence features inherent in the movements, different of mental context shifts. For future studies, it effectors (Tempel & Frings, 2014a) or movement might be interesting to investigate if mental direction (Tempel & Frings, 2015). Therefore, it context shifts also are able to affect the recall of is important to take into consideration general motor sequences. Of course, this probably would prerequisites of RIF, such as retrieval specificity, require adjusted context manipulations that are as well as peculiarities of motor memory in the able to impact motor programmes. The necessary investigation of retrieval dynamics. groundwork for examining the contribution of In recent years, memory researchers have mental context to RIF of motor sequences unfor- strongly advocated the beneficial consequences tunately is missing so far. of testing on retention (e.g., Karpicke & Blunt, The repetition of learning trials over the course 2011; Karpicke & Roediger, 2008). The current of the learning phase in the present study may conclusion seems to be that testing is one of the suggest that in learning trials occurring at the most powerful tools for enhancing memory. end of the learning phase participants might Although by far the most studies have been con- have tried to retrieve SFM before the animation cerned with verbal materials, there are a few first

Downloaded by [Central Michigan University] at 22:38 06 November 2015 graphic display appeared. Such retrieval attempts investigations now that also reported testing could have occurred during later blocks of the effects in motor action (Boutin et al., 2012, 2013; learning phase as well as during selective extra Kromann et al., 2009). Our present findings study. Thus, there might have been a retrieval imply limits for the application of testing to component to the extra study conditions. enhance retention because the observed RIF However, retrieval was neither required nor effects can be considered a side effect of testing. reinforced at study trials but the animation With regard to verbal materials, it has been graphic display started very shortly after the pres- argued that protecting influences, such as the entation of a letter stimulus. This short interval meaningful integration of facts in school textbooks hardly sufficed for accessing the corresponding to be learned, probably preclude RIF in many con- SFM in memory. Instead, the presentation texts (Chan, 2009; Chan, McDermott, & Roediger, format suggests that retrieval rather was pre- 2006). However, integration hardly can affect vented. Moreover, extra study only pertained to motor action because it involves the activation of three items. From the second extra study cycle semantic representations that do not possess cor- on, therefore, items probably could have been responding counterparts in motor memory. Extra accessed in short-term memory more easily, study did not induce forgetting but nevertheless HOW MOTOR PRACTICE SHAPES MEMORY 11

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Downloaded by [Central Michigan University] at 22:38 06 November 2015 Storm, B. C., Bjork, E. L., Bjork, R. A., & Nestojko, J. F. Right v Ring finger Index finger (2006). Is retrieval success a necessary condition for Right w Pinkie Ring finger retrieval-induced forgetting?. Psychonomic Bulletin Right x Middle finger Pinkie & Review, 13, 1023–1027. doi:10.3758/BF03213919 Right y Pinkie Index finger Storm, B. C., & Levy, B. J. (2012). A progress report on Right z Ring finger Middle finger the inhibitory account of retrieval-induced HOW MOTOR PRACTICE SHAPES MEMORY 13

APPENDIX B. ITEMS IN EXPERIMENT 2

Set A Set B

Hand Stimulus First finger Second finger First finger Second finger

Left a Index finger Pinkie Middle finger Index finger Left b Index finger Ring finger Middle finger Pinkie Left c Index finger Middle finger Middle finger Ring finger Left d Ring finger Middle finger Pinkie Ring finger Left e Ring finger Index finger Pinkie Middle finger Left f Ring finger Pinkie Pinkie Index finger Right u Ring finger Middle finger Pinkie Ring finger Right v Ring finger Index finger Pinkie Middle finger Right w Ring finger Pinkie Pinkie Index finger Right x Index finger Pinkie Middle finger Index finger Right y Index finger Ring finger Middle finger Pinkie Right z Index finger Middle finger Middle finger Ring finger Downloaded by [Central Michigan University] at 22:38 06 November 2015