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1、HIPPOCAMPUS 10:181186 (2000)Awareness Predicts the Magnitude of Single-Cue TraceEyeblink Conditioning(意識可以提前預知單信號刺激的眨眼反射)Joseph R. Manns1 Robert E. Clark,2 andLarry R. Squire13* 1 Department of Psychology, University of California at San Diego, San Diego, California 2 Department of Psychiatry, Unive
2、rsity of California at San Diego, San Diego, California 3Veterans Affairs Medical Center, San Diego, andDepartment of Neurosciences, University of California at San Diego, San Diego, CaliforniaABSTRACT(摘要): Studies of differential eyeblink conditioning (CS1 and CS-) have demonstrated that successful
3、 conditioning requires awareness of the stimulus contingencies and that delay conditioning does not. (不同情況下的眨眼反射已經證明成功的條件反射需要刺激的應急意識而delay反射不需要)Two experiments were carried out to determine whether awareness is also important for single-cue trace eyeblink conditioning.(開展了兩個實驗去判斷意識是否也對單信號trace眨眼反射很重
4、要) In experiment 1, participants who performed a secondary, attention-demanding task emitted significantly fewer conditioned eyeblink responses than participants who watched a silent movie during the conditioning session. (在實驗一中,參加一場中等注意力集中的測驗的參與者比觀看一部無聲電影的人在條件反射過程中發(fā)出明顯信號)In experiment 2, participan
5、ts who became aware of the stimulus contingencies early in the conditioning session emitted significantly more conditioned responses during the remainder of the session than participants who became aware later in the session or who never became aware. (在實驗2中,在先前預知到刺激的參與者比沒有知道或后來知道的在條件反射過程發(fā)出更加明顯信號)Th
6、ese results indicate that awareness is important for single-cue trace eyeblink conditioning, just as it is for differential trace conditioning. (這些結果表明意識是對單信號刺激的眨眼反射和trace條件反射有重要的意義的)The relationship between awareness and trace eyeblink conditioning is discussed in the light of these and other recen
7、t findings.(意識和trace眨眼反射的關系由于這些和其他的最近的發(fā)現而被討論) Hippocampus2000;10:181186.© 2000 Wiley-Liss, Inc.KEY WORDS:declarative memory; amnesia; hippocampus; classical conditioningINTRODUCTIONMemory is composed of several different abilities that depend upon different brain systems(記憶是由基于不同大腦系統的一些不同能力構成)
8、(Squire, 1992; Schacter and Tulving, 1994). Declarative memory provides the capacity for conscious recollection of facts and events and is dependent on the hippocampus and related medial temporal lobe structures. (陳述性記憶提供的事實和事件的有意識回憶的能力,是依賴于海馬和內側顳葉結構。)Nondeclarative memory is expressed through perfo
9、rmance as skills, habits, priming, and certain forms of classical conditioning and is independent of these brain structures.(非陳述記憶是通過表現技能、表達習慣、起動和某些形式的經典條件作用和獨立于這些大腦結構。)Human eyeblink classical conditioning provides a useful paradigm for exploring the distinction between declarative and nondeclarati
10、ve forms of memory. (人經典眨眼反射提供了一個有用的范式探索陳述性知識和程序形式的記憶之間的區(qū)別。)Typically, a tone (the conditioned stimulus; CS) is presented immediately before a puff of air (the unconditioned stimulus; US) is delivered to the eye. (通常的,一個音(條件刺激;CS)出現在一陣空氣(非條件刺激)吹到到眼之前。)After repeated pairings of the CS and US, indivi
11、duals begin to blink in response to the tone (the conditioned response; CR).(經過反復配對的CS和US,人開始在眨眼來對聲音采取反應(條件反應;CR) In delay eyeblink conditioning, the tone remains on until the air puff has been delivered. (在延遲眨眼反射中,聲音持續(xù)到一陣空氣被釋放)Delay conditioning is nondeclarative and can be acquired by amnesic pati
12、ents or experimental animals despite damage to the hippocampus and related structures (Norman et al., 1977; Weiskrantz and Warrington, 1979; Mauk and Thompson, 1987; Daum et al., 1989; Gabrieli et al., 1995; Clark and Squire, 1998). (延遲條件反射是非陳述性的,可以通過健忘癥患者或實驗動物盡管損害海馬和相關的結構來獲得)By contrast, in trace e
13、yeblink conditioning, an empty interval separates the end of the tone from the onset of the air puff. Amnesic patients or experimental animals with damage to the hippocampus are unable to acquire trace conditioning when the trace interval is sufficiently long (相比之下,當跟蹤的間隔足夠長的時候,trace眨眼條件反射,在聲音的結束一段時
14、間后發(fā)出空氣沖擊。健忘癥患者與海馬損傷的實驗動物無法獲得trace條件反射,)(McGlinchey-Berroth et al., 1997; Clark and Squire, 1998; Moyer et al., 1990; Solomon et al., 1986). In addition, in rabbits hippocampal lesions 1 day (but not 28 days) after trace conditioning abolish the conditioning (此外,在家兔海馬病變1天(而不是28天)后trace條件反射消失了)(Kim et
15、 al., 1995).Clark and Squire (1998) found that awareness of the stimulus contingencies was related to successful trace conditioning. At the end of the conditioning session (120 trials of differential conditioning with a CS1 and a CS-) , participants were asked about the relationship between the CS a
16、nd the US.(Clark和Squire(1998)發(fā)現,對刺突發(fā)刺激的意識與成功的trace條件反射有關。在條件反射的最后(120試驗與CS1和CS -微分調節(jié)),參與者被詢問CS和US的關系。) Only those participants who had become aware of this relationship (i.e., that the CS predicted the US) exhibited successful conditioning. (只有那些已經意識到這種關系(即使CS預示著US)表現出成功的條件條件反射。)Awareness was unrela
17、ted to differential delay conditioning. (意識與delay條件反射無關)These results suggested that trace conditioning might be dependent on the hippocampus, because declarative knowledge about the stimulus contingencies must be available during conditioning.(這些結果表明,trace條件反射可能依賴于海馬,因為對刺激的突發(fā)事件的陳述性知識必須在條件反射中可用)The
18、study by Clark and Squire (1998) followed a differential conditioning procedure. (Clark和Squire(1998)的研究是一個微分調節(jié)程序。)Two auditory stimuli were presented (a CS1 and a CS-), and conditioning was measured as the percentage of CRs to the CS1 minus the percentage of CRs to the CS-. (提出了兩個聽覺刺激(CS1和CS -),和調節(jié)測
19、量CS1 CRs的百分比減去CRs CS -的百分比。)Yet, most previous human and animal studies of eyeblink conditioning have used single-cue conditioning.(然而,大多數以前的人類和動物研究的眨眼反射用的是單信號刺激條件反射) In single-cue conditioning, only one conditioned stimulus is used (a CS1), and the level of conditioning is determined by the percent
20、age of trials in which a CR occurs. (在單信號刺激條件反射中,只有一個條件刺激(CS1),和條件反射的水平是由試驗中出現CR的百分比。)It is not known whether the importance of awareness for trace conditioning extends to the single-cue paradigm. (目前尚不清楚是否意識的重要性進行trace條件反射延伸到single-cue范式。)Indeed, it is possible that the extra demands of differentia
21、l trace conditioning (processing two CSs and appreciating their relationship to the US) might require awareness, whereas the simpler single-cue trace conditioning procedure may not (LaBar and Disterhoft, 1998).(事實上,有可能trace條件反射額外需要要求微分跟蹤調節(jié)(處理兩個CSs和鑒別它們和非條件反射的關系)可能需要意識,而簡單single-cue跟蹤調節(jié)過程可能不會(LaBar D
22、isterhoft,1998)。)To determine whether awareness is an important factor for trace single-cue eyeblink conditioning, as it is for trace differential eyeblink conditioning, we carried out two experiments.(為了確定意識是跟蹤單信號刺激眨眼反射的一個重要的因素和trace差別眨眼反射,我們進行了兩個實驗。) The first experiment compared the magnitude of
23、eyeblink conditioning in two groups of volunteers.(一個實驗中比較了兩組志愿者眨眼反射的強度。) One group watched a silent movie during the conditioning trials, and the other performed an attentiondemanding, secondary task. (一組看無聲電影調節(jié)試驗期間,和其他執(zhí)行attentiondemanding,次要任務。)If awareness of the stimulus contingencies is importa
24、nt for single-cue trace conditioning, then the group watching the movie should have become more aware and should have become conditioned to a greater extent than the group that was distracted.(如果意識到刺激突發(fā)事件對單信號刺激trace條件反射很重要,那么看電影的人應該更加有意識,應該比成為條件的人更分心。) In the second experiment, all participants watc
25、hed the silent movie and were asked about the relationship between the CS and the US early in the conditioning session, midway through the session, and at the end of the session.(在第二個實驗中,參與者觀看了無聲電影, 在早期會話,會話中,結束時被問及CS和US之間的在調節(jié)。) If awareness is important for single-cue trace conditioning, then those
26、 who become aware of the relationship between the CS and the US early in the session should condition to a greater extent than those who become aware later in the session or who never became aware.(如果意識對single-cuetrace條件反射很重要,那么那些意識到CS和US之間的關系早在會話中條件應該在更大程度上比那些后來意識到會話或從來沒有意識到。)EXPERIMENT 1, METHODSP
27、articipantsThe participants (10 men, 14 women) were volunteers or employees at the San Diego Veterans Affairs Medical Center. They averaged 69.1 years of age (range, 4778) and had an average of 16.3 years of education. They obtained WAIS-R Information and Vocabulary subscale scores of 23.8 and 60.3,
28、 respectively.(參與者(10人,14個女性)志愿者或員工在圣地亞哥退伍軍人醫(yī)療中心。他們平均為69.1歲(范圍、47 - 78)和平均16.3年的教育。他們獲得WAIS-R信息和詞匯內部氧化物分數為23.8和60.3,分別。)Apparatus and ProcedureParticipants were assigned to one of two groups. One group (n 5 14) watched a silent movie (The Gold Rush) during conditioning. (參與者被分成兩組。一組(n 5 14)看無聲電影淘金熱)
29、The other group (n 5 10) performed an attention-demanding, secondary task. (另一組(n 5 10)執(zhí)行一個要求很苛刻,次要任務。)They watched digits appearing on a computer screen (once every 1.5 s for a 1-s duration) and pressed a button whenever three odd digits appeared consecutively (Mulligan and Hartman, 1996).(他們觀看數字出現
30、在電腦屏幕上(1 s時間每隔1.5秒), 只要連續(xù)(出現三個奇怪的數字就按鈕(穆里根和哈特曼,1996)。)Participants were told that they were taking part in a study of how distraction affects learning and memory and that they would be distracted by tones and air puffs. (參與者被告知他們參加一項研究分心影響學習和記憶,他們將如何被音調和空氣沖擊。)After giving informed consent, participa
31、nts were seated in a comfortable chair in a darkened room, approximately 0.7 m from either a television monitor ( movie group) or a computer monitor (distraction task group).(給予知情同意后,受試者坐在舒適的椅子上在一個漆黑的房間里,大約距離電視監(jiān)控(電影組)或電腦顯示器(分心任務組)0.7m) One hundred twenty trace conditioning trials were then administe
32、red, with an intertrial interval of 1015 s. (。一百二十跟蹤調節(jié)試驗被實施,間隔10 - 15 s)The CS was an 85-dB, 1- kHz tone, 250 ms in duration, delivered through earphones; 1,000 ms after termination of the CS, the US was delivered. (CS是85 - db,1 - kHz的聲音,持續(xù)時間250毫秒后通過耳機發(fā)出;1000 ms CS的終止后,US。)The US was a 100-ms, 3-psi
33、 airpuff delivered to the left eye through specially designed goggles.(US是通過特別設計的眼鏡發(fā)出100 ms,3磅airpuff給左眼。) The goggles also included an infrared reflective sensor for recording eyeblinks (眼鏡還包括紅外線反射傳感器來記錄眨眼)(Clark and Squire, in press).Following the conditioning session, participants were given a tr
34、ue or false questionnaire that asked about aspects of the conditioning session.(調節(jié)會話后,要求被試完成一項或真或假的問卷調查,詢問方面調節(jié)會話。) For the Movie group, 10 questions concerned the content of the silent movie that participants watched during conditioning. (電影組,參與者被問10個問題有關無聲電影的內容。)The remaining questions were identic
35、al for both groups. Six questions asked participants to identify what stimuli had been presented, four questions asked how participants had responded to the US, and four questions asked how participants had responded to the CS.(六個問題問參與者識別刺激是否已經呈現,四個問題問參與者是如何回應US,和四個問題要求參與者對CS的反應。) Finally, the seven
36、 critical questions asked about the relationship between the CS and the US (Appendix A).(最后,七個關鍵問題問及CS和US之間的關系)EXPERIMENT 1, RESULTSFigure 1 shows the percentage of CRs emitted by each group during the first 10 trials of conditioning (Fig. 1A) and across 6 blocks of 20 trials (Fig. 1B). (圖1顯示的百分比CRs
37、發(fā)出的每組在前10的試驗條件(圖1)和6塊20試驗(圖1 b))CRs were evident during the first few trials of conditioning, as reported previously in human volunteers for both single-cue delay conditioning (Gabrieli et al., 1995, their Fig. 1; Carrillo et al., 1997, their Fig. 2) and single-cue trace conditioning (Woodruff-Pak a
38、nd Papka, 1996, their Fig. 1 ; McGlinchey-Berroth et al., 1997, their Fig. 3; Woodruff-Pak, 1999, his Fig. 1). (CRs很明顯在最初幾個試驗條件,先前報道在人類志愿者為single-cue延遲條件和single-cue跟蹤調節(jié))Performance did not noticeably improve after the first few trials. (前幾個試驗后性能沒有明顯改善)This finding also matches what has been reported
39、 previously in humans for single-cue conditioning (Gabrieli et al., 1995, her Fig. 1; Carrillo et al., 1997, their Fig. 2; Woodruff-Pak and Papka, 1996, their Fig. 1; McGlinchey-Berroth et al., 1997 , their Fig. 3).(這一發(fā)現也比賽一直在報道之前人類single-cue調節(jié)) Finally, participants in the Movie group emitted more
40、CRs than participants in the Distraction group. This difference was apparent during the first 10 trials of the session (46.9 6 4.8 % vs. 27.8 6 5.8% CRs for the Movie and Distraction groups, respectively; t(22) 5 2.53, P , 0.05), and also across all 120 trials of conditioning (45.3 6 3.8% vs. 30.4 6
41、 4.6% CRs; t(22) 5 2.5 , P , 0.05).(最后,參與者在電影集團發(fā)出CRs比分散組的參與者。這種差異是明顯的在會議的前10個試驗(46.9 6 6 4.8%和27.8 4.8% CRs電影和干擾組,分別;t(22)5 2.53,P,0.05),并在所有120試驗條件(45.3 6 6 3.8%和30.4 3.8% CRs;t(22)5 2.5,P,0.05)。)Figure 1C shows the mean awareness scores, based on the seven critical questions that asked about the r
42、elationship between the CS and US. (圖1 c顯示了意味著認識分數,根據七個關鍵問題問及CS和US之間的關系。)Participants in the Distraction group scored lower than participants in the Movie group (4.50 6 2.4 vs. 6.1 6 2.2), but this difference did not reach significance (t(22) 5 1.67, P 5 0.11). (。參與者分散組的得分低于電影組的4.50(4.50 6 2.4 vs 6.
43、1 6),但這種差異沒有達到意義(5 t(22)5 1.67,P 0.11)。)This nonsignificant difference was due to two participants in the Movie group who apparently reversed the relationship between the CS and the US and scored only 1 out of 7 correct on the seven critical questions.( 這種無意義的差異是由于兩位參與者電影組明顯逆轉CS和US之間的關系,正確得分只有1 7七個關
44、鍵問題。) Still, the awareness score for the Movie group was significantly above chance (t(13) 5 4.44, P , 0.01), whereas the awareness score for the Distraction group was not (t(9) 5 1.31, P 5 0.22). Moreover, 11 of the 14 participants in the Movie group obtained a perfect score of 7, whereas only 3 of
45、 the 10 participants in the distraction group obtained a score of 7 (X2 5 3.84, P 5,0.05).(此外,11日的14參與者在電影集團獲得滿分7分的,而只有3的10分散組的參與者獲得7分(X2 5 3.84,P 50.05)。)Participants in the Movie group performed well on the 10 questions concerning the content of the movie (mean number of correct answers 5 9.7 6 0.
46、2).(參與者在電影集團表現良好的10個問題關于電影的內容(平均數量的正確答案5 9.7 6 - 0.2)。) Participants in the Distraction group performed well on the secondary distraction task (mean percent of strings correctly identified 5 97.5 6 0.6).(干擾組的參與者表現良好在二級分心任務(平均百分比字符串正確確定5 97.5 6 - 0.6)。) Both groups were able to identify the CS and th
47、e US (5.9 and 5.6 correct out of 6 questions for the Movie and Distraction groups, respectively), and both groups acquired knowledge about how they had responded to the US (3.1 6 0.2 and 2.9 6 0.3 correct out of 4 questions; P , 0.05).(兩組人能夠識別CS和US(5.9和5.6的6個問題正確的電影和干擾組,分別),和兩組獲得的知識關于他們如何回應US(3.1 6
48、0.2和2.9 6 0.3正確4問題;P 0.05)。) Finally, neither group acquired significant knowledge about how they had responded to the CS (2.0 6 0.1 and 1.8 6 0.2 correct out of four questions).(最后,無論是集團收購了重要的知識如何回應了CS(2.0 6 0.1和1.8 6 0.1正確的四個問題)。)The findings of experiment 1 suggest that awareness of the relations
49、hip between the CS and the US is important for single-cue trace conditioning,(實驗1的發(fā)現表明,條件刺激與非條件刺激聯系中的意識,對單信號眨眼條件反射很重要) just as in the case of differential trace conditioning (就像在不同trace刺激的案列中一樣)(Clark and Squire, 1998, 1999). Engaging participants in an attention-demanding task inhibited the acquisi
50、tion of singlecue trace conditioning (Fig. 1A,B) and reduced awareness of the relationship between the CS and US.(完成一個需要集中注意力的任務時,會減少單信號條件刺激,并且減少條件刺激與非條件刺激聯系中的意識)Despite the difference between the two groups in performance(盡管兩組在表現上有所不同), most of the participants did acquire considerable awareness ab
51、out the relationship between the CS and US(大多數的參與者確實需要相當大的意識,來控制條件刺激與非條件刺激的聯系) (16 of the 24 participants scored 6 or 7 correct on the 7 critical questions, including 4 of the 10 participants in the Distraction group(16/24的參與者在7次臨時試驗中,成功6或7次;包括4/10的注意力分散組的參與者)). However, awareness was assessed only
52、at the completion of all 120 conditioning trials(但是,意識只在120次條件試驗中被監(jiān)測到). Accordingly, it was not possible to determine when in the conditioning session awareness developed(因此,在意識逐漸形成的條件反射環(huán)節(jié)中,它不可能被決定了). Some participants could have become aware towards the end of the 120 trials(一些參與者已經變得有意識傾向,在120次實驗
53、結束后)individuals would have been identified as aware but would not in fact have been aware during the course of the conditioning session(在條件反射形成過程中,個體可能被認為是意識到的,但是實際上是沒有意識到的)The second experiment sought to determine the importance of awareness during the conditioning process itself(第二個實驗探求在調整過程中,意識的重
54、要性). Conditioning trials were presented while participants watched a silent movie, as in experiment 1,(與實驗1一樣,參與者看無聲電影進行條件刺激) but awareness was tested after 10 conditioning trials, after 60 trials, and after 120 trials(但是在10條件刺激、60或120刺激后,意識被監(jiān)測到)。The question of interest was whether participants who
55、 became aware early in the conditioning session would produce more conditioned responses than participants who became aware only later in the session or who never became aware at all(有趣的問題是,是否在條件放射過程中,意識到的參與者會比遲意識到的、完全沒意識到的參與者,產生更多更強的條件刺激反應).EXPERIMENT 2, METHODS(試驗2 方法)Participants(參與者)Seven men an
56、d 13 women who did not participate in experiment 1 were recruited for experiment 2(7名男士和13名女士參加了實驗2,而未參加實驗1). Their average age was 66.2 years (range, 5175), and they averaged 14.9 years of education(平均年齡66.2歲,平均受教育14.9年). Their scores on the Information and Vocabulary subscales of the WAIS-R averag
57、ed 20.1 and 53.9, respectively(測智力的相關的).Apparatus and Procedure(儀器與過程)The procedure was the same as for experiment 1, except that all participants watched the silent movie(實驗過程與實驗1一樣,除了所有的參與者都需要看無聲電影) (The Gold Rush). Additionally, the seven critical true or false questions from experiment 1(此外,有7個來
58、自實驗1中的判斷題)FIGURE 2. A: Percentage of participants who emitted CRs during of the first 10 trials(在前10次試驗中,表現經典條件放射的參與者的百分數). B: Percent CRs across 6 blocks of 20 trials by participants who were classified as aware or unaware on the basis of their answers to the seven true or false questions given aft
59、er the first 10 trials(在10次試驗中,基于他們對給出的7個是非判斷題的意識,而被歸為意識或者無意識的參與者,). Bracketsshow SEM.(Appendix(附錄) A) were given without forewarning after the first 10 conditioning trials, again after 60 trials, and finally after all 120 trials were completed(在10次試驗條件刺激后還是沒有提前給警告,再60次試驗,直到120次試驗完成). At the end of the session, 17 of the 20 participants were also given the 10 true or false questions about the content of the movie from experiment 1(在實驗最后,17/20的參與者被給予10個關于電影類容的是非判斷題).EXPERIMENT 2, RESULTS(試驗2 結果)Fi
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