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1、HIPPOCAMPUS 10:181186 (2000)Awareness Predicts the Magnitude of Single-Cue TraceEyeblink Conditioning(意识可以提前预知单信号刺激的眨眼反射)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. (不同情况下的眨眼反射已经证明成功的条件反射需要刺激的应急意识而delay反射不需要)Two experiments were carried out to determine whether awareness is also important for single-cue trace eyeblink conditioning.(开展了两个实验去判断意识是否也对单信号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. (在实验一中,参加一场中等注意力集中的测验的参与者比观看一部无声电影的人在条件反射过程中发出明显信号)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. (在实验2中,在先前预知到刺激的参与者比没有知道或后来知道的在条件反射过程发出更加明显信号)Th
6、ese results indicate that awareness is important for single-cue trace eyeblink conditioning, just as it is for differential trace conditioning. (这些结果表明意识是对单信号刺激的眨眼反射和trace条件反射有重要的意义的)The relationship between awareness and trace eyeblink conditioning is discussed in the light of these and other recen
7、t findings.(意识和trace眨眼反射的关系由于这些和其他的最近的发现而被讨论) 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(记忆是由基于不同大脑系统的一些不同能力构成)
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. (陈述性记忆提供的事实和事件的有意识回忆的能力,是依赖于海马和内侧颞叶结构。)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.(非陈述记忆是通过表现技能、表达习惯、起动和某些形式的经典条件作用和独立于这些大脑结构。)Human eyeblink classical conditioning provides a useful paradigm for exploring the distinction between declarative and nondeclarati
10、ve forms of memory. (人经典眨眼反射提供了一个有用的范式探索陈述性知识和程序形式的记忆之间的区别。)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,一个音(条件刺激;CS)出现在一阵空气(非条件刺激)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).(经过反复配对的CS和US,人开始在眨眼来对声音采取反应(条件反应;CR) In delay eyeblink conditioning, the tone remains on until the air puff has been delivered. (在延迟眨眼反射中,声音持续到一阵空气被释放)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). (延迟条件反射是非陈述性的,可以通过健忘症患者或实验动物尽管损害海马和相关的结构来获得)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,当跟踪的间隔足够长的时候,trace眨眼条件反射,在声音的结束一段时
14、间后发出空气冲击。健忘症患者与海马损伤的实验动物无法获得trace条件反射,)(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,在家兔海马病变1天(而不是28天)后trace条件反射消失了)(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)发现,对刺突发刺激的意识与成功的trace条件反射有关。在条件反射的最后(120试验与CS1和CS -微分调节),参与者被询问CS和US的关系。) Only those participants who had become aware of this relationship (i.e., that the CS predicted the US) exhibited successful conditioning. (只有那些已经意识到这种关系(即使CS预示着US)表现出成功的条件条件反射。)Awareness was unrela
17、ted to differential delay conditioning. (意识与delay条件反射无关)These results suggested that trace conditioning might be dependent on the hippocampus, because declarative knowledge about the stimulus contingencies must be available during conditioning.(这些结果表明,trace条件反射可能依赖于海马,因为对刺激的突发事件的陈述性知识必须在条件反射中可用)The
18、study by Clark and Squire (1998) followed a differential conditioning procedure. (Clark和Squire(1998)的研究是一个微分调节程序。)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-. (提出了两个听觉刺激(CS1和CS -),和调节测
19、量CS1 CRs的百分比减去CRs CS -的百分比。)Yet, most previous human and animal studies of eyeblink conditioning have used single-cue conditioning.(然而,大多数以前的人类和动物研究的眨眼反射用的是单信号刺激条件反射) 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. (在单信号刺激条件反射中,只有一个条件刺激(CS1),和条件反射的水平是由试验中出现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识的重要性进行trace条件反射延伸到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).(事实上,有可能trace条件反射额外需要要求微分跟踪调节(处理两个CSs和鉴别它们和非条件反射的关系)可能需要意识,而简单single-cue跟踪调节过程可能不会(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.(为了确定意识是跟踪单信号刺激眨眼反射的一个重要的因素和trace差别眨眼反射,我们进行了两个实验。) The first experiment compared the magnitude of
23、eyeblink conditioning in two groups of volunteers.(一个实验中比较了两组志愿者眨眼反射的强度。) One group watched a silent movie during the conditioning trials, and the other performed an attentiondemanding, secondary task. (一组看无声电影调节试验期间,和其他执行attentiondemanding,次要任务。)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.(如果意识到刺激突发事件对单信号刺激trace条件反射很重要,那么看电影的人应该更加有意识,应该比成为条件的人更分心。) 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.(在第二个实验中,参与者观看了无声电影, 在早期会话,会话中,结束时被问及CS和US之间的在调节。) 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.(如果意识对single-cuetrace条件反射很重要,那么那些意识到CS和US之间的关系早在会话中条件应该在更大程度上比那些后来意识到会话或从来没有意识到。)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.(参与者(10人,14个女性)志愿者或员工在圣地亚哥退伍军人医疗中心。他们平均为69.1岁(范围、47 - 78)和平均16.3年的教育。他们获得WAIS-R信息和词汇内部氧化物分数为23.8和60.3,分别。)Apparatus and ProcedureParticipants were assigned to one of two groups. One group (n 5 14) watched a silent movie (The Gold Rush) during conditioning. (参与者被分成两组。一组(n 5 14)看无声电影淘金热)
29、The other group (n 5 10) performed an attention-demanding, secondary task. (另一组(n 5 10)执行一个要求很苛刻,次要任务。)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).(他们观看数字出现
30、在电脑屏幕上(1 s时间每隔1.5秒), 只要连续(出现三个奇怪的数字就按钮(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. (参与者被告知他们参加一项研究分心影响学习和记忆,他们将如何被音调和空气冲击。)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).(给予知情同意后,受试者坐在舒适的椅子上在一个漆黑的房间里,大约距离电视监控(电影组)或电脑显示器(分心任务组)0.7m) One hundred twenty trace conditioning trials were then administe
32、red, with an intertrial interval of 1015 s. (。一百二十跟踪调节试验被实施,间隔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的声音,持续时间250毫秒后通过耳机发出;1000 ms CS的终止后,US。)The US was a 100-ms, 3-psi
33、 airpuff delivered to the left eye through specially designed goggles.(US是通过特别设计的眼镜发出100 ms,3磅airpuff给左眼。) The goggles also included an infrared reflective sensor for recording eyeblinks (眼镜还包括红外线反射传感器来记录眨眼)(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.(调节会话后,要求被试完成一项或真或假的问卷调查,询问方面调节会话。) For the Movie group, 10 questions concerned the content of the silent movie that participants watched during conditioning. (电影组,参与者被问10个问题有关无声电影的内容。)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.(六个问题问参与者识别刺激是否已经呈现,四个问题问参与者是如何回应US,和四个问题要求参与者对CS的反应。) Finally, the seven
36、 critical questions asked about the relationship between the CS and the US (Appendix A).(最后,七个关键问题问及CS和US之间的关系)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). (图1显示的百分比CRs
37、发出的每组在前10的试验条件(图1)和6块20试验(图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很明显在最初几个试验条件,先前报道在人类志愿者为single-cue延迟条件和single-cue跟踪调节)Performance did not noticeably improve after the first few trials. (前几个试验后性能没有明显改善)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).(这一发现也比赛一直在报道之前人类single-cue调节) 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,参与者在电影集团发出CRs比分散组的参与者。这种差异是明显的在会议的前10个试验(46.9 6 6 4.8%和27.8 4.8% CRs电影和干扰组,分别;t(22)5 2.53,P,0.05),并在所有120试验条件(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. (图1 c显示了意味着认识分数,根据七个关键问题问及CS和US之间的关系。)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). (。参与者分散组的得分低于电影组的4.50(4.50 6 2.4 vs 6.
43、1 6),但这种差异没有达到意义(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.( 这种无意义的差异是由于两位参与者电影组明显逆转CS和US之间的关系,正确得分只有1 7七个关
44、键问题。) 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14参与者在电影集团获得满分7分的,而只有3的10分散组的参与者获得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).(参与者在电影集团表现良好的10个问题关于电影的内容(平均数量的正确答案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).(干扰组的参与者表现良好在二级分心任务(平均百分比字符串正确确定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).(两组人能够识别CS和US(5.9和5.6的6个问题正确的电影和干扰组,分别),和两组获得的知识关于他们如何回应US(3.1 6
48、0.2和2.9 6 0.3正确4问题;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,无论是集团收购了重要的知识如何回应了CS(2.0 6 0.1和1.8 6 0.1正确的四个问题)。)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,(实验1的发现表明,条件刺激与非条件刺激联系中的意识,对单信号眨眼条件反射很重要) just as in the case of differential trace conditioning (就像在不同trace刺激的案列中一样)(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.(完成一个需要集中注意力的任务时,会减少单信号条件刺激,并且减少条件刺激与非条件刺激联系中的意识)Despite the difference between the two groups in performance(尽管两组在表现上有所不同), most of the participants did acquire considerable awareness ab
51、out the relationship between the CS and US(大多数的参与者确实需要相当大的意识,来控制条件刺激与非条件刺激的联系) (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的参与者在7次临时试验中,成功6或7次;包括4/10的注意力分散组的参与者)). However, awareness was assessed only
52、at the completion of all 120 conditioning trials(但是,意识只在120次条件试验中被监测到). Accordingly, it was not possible to determine when in the conditioning session awareness developed(因此,在意识逐渐形成的条件反射环节中,它不可能被决定了). Some participants could have become aware towards the end of the 120 trials(一些参与者已经变得有意识倾向,在120次实验
53、结束后)individuals would have been identified as aware but would not in fact have been aware during the course of the conditioning session(在条件反射形成过程中,个体可能被认为是意识到的,但是实际上是没有意识到的)The second experiment sought to determine the importance of awareness during the conditioning process itself(第二个实验探求在调整过程中,意识的重
54、要性). Conditioning trials were presented while participants watched a silent movie, as in experiment 1,(与实验1一样,参与者看无声电影进行条件刺激) but awareness was tested after 10 conditioning trials, after 60 trials, and after 120 trials(但是在10条件刺激、60或120刺激后,意识被监测到)。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(有趣的问题是,是否在条件放射过程中,意识到的参与者会比迟意识到的、完全没意识到的参与者,产生更多更强的条件刺激反应).EXPERIMENT 2, METHODS(试验2 方法)Participants(参与者)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名女士参加了实验2,而未参加实验1). Their average age was 66.2 years (range, 5175), and they averaged 14.9 years of education(平均年龄66.2岁,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(测智力的相关的).Apparatus and Procedure(仪器与过程)The procedure was the same as for experiment 1, except that all participants watched the silent movie(实验过程与实验1一样,除了所有的参与者都需要看无声电影) (The Gold Rush). Additionally, the seven critical true or false questions from experiment 1(此外,有7个来
58、自实验1中的判断题)FIGURE 2. A: Percentage of participants who emitted CRs during of the first 10 trials(在前10次试验中,表现经典条件放射的参与者的百分数). 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次试验中,基于他们对给出的7个是非判断题的意识,而被归为意识或者无意识的参与者,). Bracketsshow SEM.(Appendix(附录) 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次试验条件刺激后还是没有提前给警告,再60次试验,直到120次试验完成). 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(在实验最后,17/20的参与者被给予10个关于电影类容的是非判断题).EXPERIMENT 2, RESULTS(试验2 结果)Fi
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