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工作记忆 (查看源代码)

2020年8月9日 (日) 17:05的版本

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无编辑摘要

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=== The multicomponent model ===

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=== 多组件模型 The multicomponent model ===

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=== Working memory as part of long-term memory ===

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=== 作为长期记忆一部分的工作记忆 Working memory as part of long-term memory ===

{{Annotated image|caption=The central executive of working memory is retrieving memory from long-term memory.|image=WorkingMemory Label Free.jpg|width=320|height=179|image-width=320|image-left=0|image-top=0|annotations={{Annotation|130|15|Central Executive|font-weight=bold|font-size=10}}

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奥伯奥尔 Oberauer 通过添加第三个组件扩展了考恩 Cowan 的模型，第三个组件是一个更窄的注意焦点，一次只能容纳一个块（chunk）。一元素焦点系统嵌入在四元素焦点系统中，用于选择要处理的单个块。例如，在考恩 Cowan 的“注意力焦点”中，四个数字可以同时出现在脑海中。当个人希望对每个数字进行处理时（例如，将数字2加到每个数字）就需要对每个数字进行独立处理，因为大多数个人不能同时进行多个数学处理。奥伯奥尔 Oberauer 的注意力组件选择其中一个数字进行处理，然后将注意力的焦点转移到下一个数字，直到所有数字都处理完毕。

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== Capacity ==

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== 容量 Capacity ==

Working memory is widely acknowledged as having limited capacity. An early quantification of the capacity limit associated with short-term memory was the "[[The Magical Number Seven, Plus or Minus Two|magical number seven]]" suggested by Miller in 1956.<ref name="miller">{{Cite journal|author=Miller GA |title=The magical number seven plus or minus two: some limits on our capacity for processing information |journal=Psychological Review |volume=63 |issue=2 |pages=81–97 |date=March 1956 |pmid=13310704 |doi=10.1037/h0043158|citeseerx=10.1.1.308.8071 }} Republished: {{Cite journal|author=Miller GA |title=The magical number seven, plus or minus two: some limits on our capacity for processing information. 1956 |journal=Psychological Review |volume=101 |issue=2 |pages=343–52 |date=April 1994 |pmid=8022966 |doi=10.1037/0033-295X.101.2.343}}</ref> He claimed that the information-processing capacity of young adults is around seven elements, which he called "chunks", regardless of whether the elements are digits, letters, words, or other units. Later research revealed this number depends on the category of chunks used (e.g., span may be around seven for digits, six for letters, and five for words), and even on features of the [[chunking (psychology)|chunks]] within a category. For instance, span is lower for long than short words. In general, memory span for verbal contents (digits, letters, words, etc.) depends on the phonological complexity of the content (i.e., the number of phonemes, the number of syllables),<ref>{{Cite journal|last=Service|first=Elisabet|date=1998-05-01|title=The Effect of Word Length on Immediate Serial Recall Depends on Phonological Complexity, Not Articulatory Duration|journal=The Quarterly Journal of Experimental Psychology Section A|volume=51|issue=2|pages=283–304|doi=10.1080/713755759|issn=0272-4987}}</ref> and on the lexical status of the contents (whether the contents are words known to the person or not).<ref>{{Cite journal|first1=Charles |last1=Hulme |first2=Steven |last2=Roodenrys |first3=Gordon |last3=Brown |first4=Robin |last4=Mercer |date=November 1995 |title=The role of long-term memory mechanisms in memory span |journal=British Journal of Psychology |volume=86 |issue=4 |pages=527–36 |doi=10.1111/j.2044-8295.1995.tb02570.x}}</ref> Several other factors affect a person's measured span, and therefore it is difficult to pin down the capacity of short-term or working memory to a number of chunks. Nonetheless, Cowan proposed that working memory has a capacity of about four chunks in young adults (and fewer in children and old adults).<ref>{{Cite journal|first1=Nelson |last1=Cowan |year=2001 |title=The magical number 4 in short-term memory: A reconsideration of mental storage capacity |journal=Behavioral and Brain Sciences |volume=24 |issue=1 |pages=87–185 |doi=10.1017/S0140525X01003922 |pmid=11515286|doi-access=free }}</ref>

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=== Measures and correlates ===

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=== 测量和关联 Measures and correlates ===

Working memory capacity can be tested by a variety of tasks. A commonly used measure is a dual-task paradigm, combining a [[memory span]] measure with a concurrent processing task, sometimes referred to as "complex span". Daneman and Carpenter invented the first version of this kind of task, the "[[reading span]]", in 1980.<ref>{{Cite journal|first1=Meredyth |last1=Daneman |first2=Patricia A. |last2=Carpenter |date=August 1980 |title=Individual differences in working memory and reading |journal=Journal of Verbal Learning & Verbal Behavior |volume=19 |issue=4 |pages=450–66 |doi=10.1016/S0022-5371(80)90312-6}}</ref> Subjects read a number of sentences (usually between two and six) and tried to remember the last word of each sentence. At the end of the list of sentences, they repeated back the words in their correct order. Other tasks that do not have this dual-task nature have also been shown to be good measures of working memory capacity.<ref>{{Cite journal|last2=Süss|first2=H.-M.|last3=Schulze|first3=R.|last4=Wilhelm|first4=O.|last5=Wittmann|first5=W. W.|date=December 2000|title=Working memory capacity—facets of a cognitive ability construct|journal=Personality and Individual Differences|volume=29|issue=6|pages=1017–45|doi=10.1016/S0191-8869(99)00251-2|first1=K.|last1=Oberauer}}</ref> Whereas Daneman and Carpenter believed that the combination of "storage" (maintenance) and processing is needed to measure working memory capacity, we know now that the capacity of working memory can be measured with short-term memory tasks that have no additional processing component.<ref>{{Cite journal|last1=Unsworth|first1=Nash|last2=Engle|first2=Randall W.|title=On the division of short-term and working memory: An examination of simple and complex span and their relation to higher order abilities.|journal=Psychological Bulletin|volume=133|issue=6|pages=1038–1066|doi=10.1037/0033-2909.133.6.1038|pmid=17967093|year=2007}}</ref><ref>{{Cite journal|last=Colom, R. Abad, F. J. Quiroga, M. A. Shih, P. C. Flores-Mendoza, C.|year=2008|title=Working memory and intelligence are highly related constructs, but why?|journal=Intelligence|volume=36|issue=6|pages=584–606|doi=10.1016/j.intell.2008.01.002}}</ref> Conversely, working memory capacity can also be measured with certain processing tasks that don't involve maintenance of information.<ref>{{Cite journal|last=Oberauer, K. Süß, H.-M. Wilhelm, O. Wittmann, W. W.|year=2003|title=The multiple faces of working memory - storage, processing, supervision, and coordination|doi=10.1016/s0160-2896(02)00115-0|journal=Intelligence|volume=31|issue=2|pages=167–193|url=https://www.zora.uzh.ch/id/eprint/97155/1/intelligence.pdf}}</ref><ref>{{Cite journal|last=Chuderski|first=Adam|date=2013-09-25|title=The relational integration task explains fluid reasoning above and beyond other working memory tasks|journal=Memory & Cognition|language=en|volume=42|issue=3|pages=448–463|doi=10.3758/s13421-013-0366-x|issn=0090-502X|pmc=3969517|pmid=24222318}}</ref> The question of what features a task must have to qualify as a good measure of working memory capacity is a topic of ongoing research.

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=== Experimental studies of working-memory capacity ===

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=== 工作记忆容量的试验研究 Experimental studies of working-memory capacity ===

There are several hypotheses about the nature of the capacity limit. One is that a limited pool of cognitive resources is needed to keep representations active and thereby available for processing, and for carrying out processes.<ref name=":0">{{Cite journal|author=Just, M. A.|author2=Carpenter, P. A. |title=A capacity theory of comprehension: individual differences in working memory |journal=Psychological Review |volume=99 |issue=1 |pages=122–49 |date=January 1992 |pmid=1546114 |doi=10.1037/0033-295X.99.1.122|url=http://repository.cmu.edu/cgi/viewcontent.cgi?article=1730&context=psychology }}</ref> Another hypothesis is that memory traces in working memory decay within a few seconds, unless refreshed through rehearsal, and because the speed of rehearsal is limited, we can maintain only a limited amount of information.<ref>{{Cite journal|doi=10.3758/BF03198549|author=Towse, J. N.|author2=Hitch, G. J.|author3=Hutton, U.|title=On the interpretation of working memory span in adults |journal=Memory & Cognition |volume=28 |issue=3 |pages=341–8 |date=April 2000 |pmid=10881551|doi-access=free }}</ref> Yet another idea is that representations held in working memory interfere with each other.<ref>{{Cite journal|vauthors=Waugh NC, Norman DA |title=Primary Memory |journal=Psychological Review |volume=72 |issue= 2|pages=89–104 |date=March 1965 |pmid=14282677 |doi=10.1037/h0021797}}</ref>

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====Decay theories====

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==== 衰变理论 Decay theories====

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====Resource theories====

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==== 资源理论 Resource theories====

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====Interference theories====

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==== Interference theories====

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==== Limitations ====

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==== 极限 Limitations ====

None of these hypotheses can explain the experimental data entirely. The resource hypothesis, for example, was meant to explain the trade-off between maintenance and processing: The more information must be maintained in working memory, the slower and more error prone concurrent processes become, and with a higher demand on concurrent processing memory suffers. This trade-off has been investigated by tasks like the reading-span task described above. It has been found that the amount of trade-off depends on the similarity of the information to be remembered and the information to be processed. For example, remembering numbers while processing spatial information, or remembering spatial information while processing numbers, impair each other much less than when material of the same kind must be remembered and processed.<ref>{{Cite journal|doi=10.1016/j.jml.2006.07.009 |title=The relationship between processing and storage in working memory span: Not two sides of the same coin |date=February 2007 |first1=Yukio |last1=Maehara |first2=Satoru |last2=Saito |journal=Journal of Memory and Language |volume=56 |issue=2 |pages=212–228}}</ref> Also, remembering words and processing digits, or remembering digits and processing words, is easier than remembering and processing materials of the same category.<ref>{{Cite journal|doi=10.1076/anec.6.2.99.784 |title=Selection from Working Memory: on the Relationship between Processing and Storage Components |date=June 1999 |last1=Li |first1=Karen Z.H. |journal=Aging, Neuropsychology, and Cognition |volume=6 |issue=2 |pages=99–116}}</ref> These findings are also difficult to explain for the decay hypothesis, because decay of memory representations should depend only on how long the processing task delays rehearsal or recall, not on the content of the processing task. A further problem for the decay hypothesis comes from experiments in which the recall of a list of letters was delayed, either by instructing participants to recall at a slower pace, or by instructing them to say an irrelevant word once or three times in between recall of each letter. Delaying recall had virtually no effect on recall accuracy.<ref>{{Cite journal|doi=10.3758/BF03196705|vauthors=Lewandowsky S, Duncan M, Brown GD |title=Time does not cause forgetting in short-term serial recall |journal=Psychonomic Bulletin & Review |volume=11 |issue=5 |pages=771–90 |date=October 2004 |pmid=15732687 |url=http://pbr.psychonomic-journals.org/cgi/pmidlookup?view=long&pmid=15732687|doi-access=free }}</ref><ref>{{Cite journal|vauthors=Oberauer K, Lewandowsky S |title=Forgetting in immediate serial recall: decay, temporal distinctiveness, or interference? |journal=Psychological Review |volume=115 |issue=3 |pages=544–76 |date=July 2008 |pmid=18729591 |doi=10.1037/0033-295X.115.3.544|url=https://api.research-repository.uwa.edu.au/files/1546099/11204_PID11204.pdf }}</ref> The [[interference theory]] seems to fare best with explaining why the similarity between memory contents and the contents of concurrent processing tasks affects how much they impair each other. More similar materials are more likely to be confused, leading to retrieval competition.

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== Development ==

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== 发展 Development ==

The capacity of working memory increases gradually over childhood<ref name="ReferenceA">{{cite journal | doi = 10.1037/0012-1649.40.2.177 | last1 = Gathercole | first1 = S. E. | last2 = Pickering | first2 = S. J. | last3 = Ambridge | first3 = B. | last4 = Wearing | first4 = H. | year = 2004 | title = The structure of working memory from 4 to 15 years of age | journal = Developmental Psychology | volume = 40 | issue = 2| pages = 177–190 | pmid = 14979759 | citeseerx = 10.1.1.529.2727 }}</ref> and declines gradually in old age.<ref>{{cite journal | doi = 10.1037/0894-4105.8.4.535 | last1 = Salthouse | first1 = T. A. | year = 1994 | title = The aging of working memory | journal = Neuropsychology | volume = 8 | issue = 4| pages = 535–543 }}</ref>

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=== Childhood ===

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=== 童年 Childhood ===

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=== Aging ===

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=== 老化 Aging ===

Working memory is among the cognitive functions most sensitive to decline in [[old age]].<ref name="Hertzog 2003">{{cite journal |vauthors=Hertzog C, Dixon RA, Hultsch DF, MacDonald SW |title=Latent change models of adult cognition: are changes in processing speed and working memory associated with changes in episodic memory? |journal=Psychol Aging |volume=18 |issue=4 |pages=755–69 |date=December 2003 |pmid=14692862 |doi=10.1037/0882-7974.18.4.755 }}</ref><ref name="Park, D. C. 2002">{{cite journal |vauthors=Park DC, Lautenschlager G, Hedden T, Davidson NS, Smith AD, Smith PK |title=Models of visuospatial and verbal memory across the adult life span |journal=Psychol Aging |volume=17 |issue=2 |pages=299–320 |date=June 2002 |pmid=12061414 |doi= 10.1037/0882-7974.17.2.299 }}</ref> Several explanations have been offered for this decline in psychology. One is the processing speed theory of cognitive aging by Tim Salthouse.<ref>{{cite journal | doi = 10.1037/0033-295X.103.3.403 | last1 = Salthouse | first1 = T. A. | year = 1996 | title = The processing speed theory of adult age differences in cognition | journal = Psychological Review | volume = 103 | issue = 3| pages = 403–428 | pmid = 8759042 | citeseerx = 10.1.1.464.585 }}</ref> Drawing on the finding of general slowing of cognitive processes as people grow older, Salthouse argues that slower processing leaves more time for working-memory contents to decay, thus reducing effective capacity. However, the decline of working-memory capacity cannot be entirely attributed to slowing because capacity declines more in old age than speed.<ref name="Park, D. C. 2002" /><ref>{{cite journal | doi = 10.1016/0010-0277(95)00689-3 | last1 = Mayr | first1 = U. | last2 = Kliegl | first2 = R. | last3 = Krampe | first3 = R. T. | year = 1996 | title = Sequential and coordinative processing dynamics in figural transformation across the life span | journal = Cognition | volume = 59 | issue = 1| pages = 61–90 | pmid = 8857471 }}</ref> Another proposal is the inhibition hypothesis advanced by [[Lynn Hasher]] and Rose Zacks.<ref>Hasher, L., & Zacks, R. T. (1988). Working memory, comprehension, and aging: A review and new view. In G. H. Bower (Ed.), ''The psychology of learning and motivation'', ''Vol. 22'', (pp. 193–225). New York: Academic Press.</ref> This theory assumes a general deficit in old age in the ability to inhibit irrelevant, or no-longer relevant, information. Therefore, working memory tends to be cluttered with irrelevant contents that reduce the effective capacity for relevant content. The assumption of an inhibition deficit in old age has received much empirical support<ref>Hasher, L., Zacks, R. T., & May, C. P. (1999). Inhibitory control, circadian arousal, and age. In D. Gopher & A. Koriat (Eds.), ''Attention and Performance'' (pp. 653–675). Cambridge, MA: MIT Press.</ref> but, so far, it is not clear whether the decline in inhibitory ability fully explains the decline of working-memory capacity. An explanation on the neural level of the decline of working memory and other cognitive functions in old age has been proposed by West.<ref>{{cite journal | doi = 10.1037/0033-2909.120.2.272 | last1 = West | first1 = R. L. | year = 1996 | title = An application of prefrontal cortex function theory to cognitive aging | journal = Psychological Bulletin | volume = 120 | issue = 2| pages = 272–292 | pmid = 8831298 }}</ref> She argued that working memory depends to a large degree on the [[pre-frontal cortex]], which deteriorates more than other brain regions as we grow old. Age related decline in working memory can be briefly reversed using low intensity transcranial stimulation, synchronizing rhythms in bilateral frontal and left temporal lobe areas.<ref>{{Cite news|url=https://www.theguardian.com/science/2019/apr/08/scientists-use-electrical-pulses-reverse-memory-decline-ageing|title=Scientists reverse memory decline using electrical pulses|last=Devlin, H.|date=2019-04-08|work=The Guardian|access-date=2019-04-09|language=en-GB|issn=0261-3077}}</ref>

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== Training ==

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== 训练 Training ==

{{further|Working memory training|Neurobiological effects of physical exercise#Cognitive control and memory}}

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== In the brain ==

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== 脑内 In the brain ==

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=== Neural mechanisms of maintaining information ===

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=== 信息维持的神经机制 Neural mechanisms of maintaining information ===

The first insights into the neuronal and neurotransmitter basis of working memory came from animal research. The work of Jacobsen<ref>{{Cite journal|author=Jacobsen CF|title= Studies of cerebral function in primates |journal=Comparative Psychology Monographs |volume=13 |issue=3 |pages=1–68 |year=1938 |oclc=250695441 }}</ref> and Fulton in the 1930s first showed that lesions to the PFC impaired spatial working memory performance in monkeys. The later work of [[Joaquin Fuster]]<ref>{{Cite journal|author=Fuster JM |title=Unit activity in prefrontal cortex during delayed-response performance: neuronal correlates of transient memory |journal=Journal of Neurophysiology |volume=36 |issue=1 |pages=61–78 |date=January 1973 |pmid=4196203 |doi=10.1152/jn.1973.36.1.61 }}</ref> recorded the electrical activity of neurons in the PFC of monkeys while they were doing a delayed matching task. In that task, the monkey sees how the experimenter places a bit of food under one of two identical-looking cups. A shutter is then lowered for a variable delay period, screening off the cups from the monkey's view. After the delay, the shutter opens and the monkey is allowed to retrieve the food from under the cups. Successful retrieval in the first attempt – something the animal can achieve after some training on the task – requires holding the location of the food in memory over the delay period. Fuster found neurons in the PFC that fired mostly during the delay period, suggesting that they were involved in representing the food location while it was invisible. Later research has shown similar delay-active neurons also in the posterior [[parietal cortex]], the [[thalamus]], the [[Caudate nucleus|caudate]], and the [[globus pallidus]].<ref>{{Cite journal|vauthors=Ashby FG, Ell SW, Valentin VV, Casale MB |title=FROST: a distributed neurocomputational model of working memory maintenance |journal=Journal of Cognitive Neuroscience |volume=17 |issue=11 |pages=1728–43 |date=November 2005 |pmid=16269109 |doi=10.1162/089892905774589271|citeseerx=10.1.1.456.7179 }}</ref> The work of [[Patricia Goldman-Rakic|Goldman-Rakic]] and others showed that principal sulcal, dorsolateral PFC interconnects with all of these brain regions, and that neuronal microcircuits within PFC are able to maintain information in working memory through recurrent excitatory glutamate networks of pyramidal cells that continue to fire throughout the delay period.<ref>{{Cite journal|author=Goldman-Rakic PS|title= Cellular basis of working memory |journal=Neuron |volume=14 |issue= 3 |pages=447–485 |year=1995 | pmid = 7695894 | doi = 10.1016/0896-6273(95)90304-6 }}</ref> These circuits are tuned by lateral inhibition from GABAergic interneurons.<ref>{{Cite journal|vauthors=Rao SG, Williams GV, Goldman-Rakic PS |title= Destruction and creation of spatial tuning by disinhibition: GABA(A) blockade of prefrontal cortical neurons engaged by working memory |journal=Journal of Neuroscience |volume=20 |pages=485–494 |year=2000|pmid=10627624 |pmc= 6774140 |issue=1|doi= 10.1523/JNEUROSCI.20-01-00485.2000 }}</ref> The neuromodulatory arousal systems markedly alter PFC working memory function; for example, either too little or too much dopamine or norepinephrine impairs PFC network firing<ref>{{Cite journal|doi=10.1016/j.tics.2010.05.003|author1=Arnsten AFT |author2=Paspalas CD |author3=Gamo NJ |author4=Y. Y |author5=Wang M |title= Dynamic Network Connectivity: A new form of neuroplasticity|journal=Trends in Cognitive Sciences|volume=14 |pages=365–375 |year=2010|issue=8|pmid=20554470|pmc=2914830}}</ref> and working memory performance.<ref>{{Cite journal|doi=10.1146/annurev.neuro.051508.135535|vauthors=Robbins TW, Arnsten AF |title= The neuropsychopharmacology of fronto-executive function: monoaminergic modulation |journal=Annu Rev Neurosci|volume=32 |pages=267–287 |year=2009|pmid=19555290|pmc=2863127}}</ref>

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=== Localization in the brain ===

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=== 脑内定位 Localization in the brain ===

Localization of brain functions in humans has become much easier with the advent of [[brain imaging]] methods ([[Positron emission tomography|PET]] and [[fMRI]]). This research has confirmed that areas in the PFC are involved in working memory functions. During the 1990s much debate has centered on the different functions of the ventrolateral (i.e., lower areas) and the [[Dorsolateral prefrontal cortex|dorsolateral (higher) areas of the PFC]]. A human lesion study provides additional evidence for the role of the [[dorsolateral prefrontal cortex]] in working memory.<ref>{{cite journal|last2=Koenigs|first2=Michael|last3=Grafman|first3=Jordan|year=2013|title=Dorsolateral prefrontal contributions to human working memory|journal=Cortex|volume=49|issue=5|pages=1195–1205|doi=10.1016/j.cortex.2012.05.022|pmid=22789779|last1=Barbey|first1=Aron K.|pmc=3495093}}</ref> One view was that the dorsolateral areas are responsible for spatial working memory and the ventrolateral areas for non-spatial working memory. Another view proposed a functional distinction, arguing that ventrolateral areas are mostly involved in pure maintenance of information, whereas dorsolateral areas are more involved in tasks requiring some processing of the memorized material. The debate is not entirely resolved but most of the evidence supports the functional distinction.<ref>{{Cite journal|author=Owen, A. M.|title=The functional organization of working memory processes within human lateral frontal cortex: the contribution of functional neuroimaging |journal=The European Journal of Neuroscience |volume=9 |issue=7 |pages=1329–39 |date=July 1997 |pmid=9240390 |doi=10.1111/j.1460-9568.1997.tb01487.x}}</ref>

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=== Effects of stress on neurophysiology ===

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=== 神经生理学的压力效果 Effects of stress on neurophysiology ===

Working memory is impaired by acute and chronic psychological stress. This phenomenon was first discovered in animal studies by Arnsten and colleagues,<ref>{{Cite journal|doi=10.1126/science.280.5370.1711|author=Arnsten, A. F.|title=The biology of being frazzled |journal=Science |volume=280 |issue=5370 |pages=1711–2 |date=June 1998 |pmid=9660710}}</ref> who have shown that stress-induced [[catecholamine]] release in PFC rapidly decreases PFC neuronal firing and impairs working memory performance through feedforward, intracellular signaling pathways.<ref>{{Cite journal|author=Arnsten, AF |title=Stress signalling pathways that impair prefrontal cortex structure and function |journal=Nature Reviews Neuroscience |volume=10 |issue=6 |pages=410–22 |date=June 2009 |pmid=19455173|pmc=2907136 |doi=10.1038/nrn2648}}</ref> Exposure to chronic stress leads to more profound working memory deficits and additional architectural changes in PFC, including dendritic atrophy and spine loss,<ref>{{Cite journal|author=Radley, J. J.|author2= Rocher, A. B.|author3=Miller, M.|author4= Janssen, W. G.|author5=Liston, C.|author6=Hof, P. R.|author7=McEwen, B. S.|author8=Morrison, J. H.|title=Repeated stress induces dendritic spine loss in the rat medial prefrontal cortex |journal=Cereb Cortex |volume=16 |issue=3 |pages=313–20 |date=Mar 2006 |pmid=15901656 |doi=10.1093/cercor/bhi104|doi-access=free}}</ref> which can be prevented by inhibition of protein kinase C signaling.<ref>{{Cite journal|author=Hains, A. B.|author2=Vu, M. A.|author3=Maciejewski, P. K.|author4= van Dyck, C. H. |authorlink4=Christopher H. van Dyck |author5=Gottron, M.|author6= Arnsten, A. F. |title=Inhibition of protein kinase C signaling protects prefrontal cortex dendritic spines and cognition from the effects of chronic stress |journal=Proceedings of the National Academy of Sciences|volume=106 |issue=42 |pages=17957–62 |date=Oct 2009 |pmid=19805148|pmc=2742406 |doi=10.1073/pnas.0908563106|bibcode=2009PNAS..10617957H }}</ref> [[fMRI]] research has extended this research to humans, and confirms that reduced working memory caused by acute stress links to reduced activation of the PFC, and stress increased levels of [[catecholamine]]s.<ref>{{Cite journal|vauthors=Qin S, Hermans EJ, van Marle HJ, Luo J, Fernández G |title=Acute psychological stress reduces working memory-related activity in the dorsolateral prefrontal cortex |journal=Biological Psychiatry |volume=66 |issue=1 |pages=25–32 |date=July 2009 |pmid=19403118 |doi=10.1016/j.biopsych.2009.03.006}}</ref> Imaging studies of medical students undergoing stressful exams have also shown weakened PFC functional connectivity, consistent with the animal studies.<ref>{{Cite journal|vauthors=Liston C, McEwen BS, Casey BJ |title=Psychosocial stress reversibly disrupts prefrontal processing and attentional control |journal=Proceedings of the National Academy of Sciences|volume=106 |issue=3 |pages=912–7 |date=Jan 2009 |pmid=19139412|pmc=2621252 |doi=10.1073/pnas.0807041106|bibcode=2009PNAS..106..912L }}</ref> The marked effects of stress on PFC structure and function may help to explain how stress can cause or exacerbate mental illness.

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=== Effects of alcohol on neurophysiology ===

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=== 神经生理学的酒精效果 Effects of alcohol on neurophysiology ===

Alcohol abuse can result in brain damage which impairs working memory.<ref name="pmid21466500">{{cite journal |vauthors=van Holst RJ, Schilt T |title=Drug-related decrease in neuropsychological functions of abstinent drug users |journal=Curr Drug Abuse Rev |volume=4 |issue=1 |pages=42–56 | date=March 2011 |pmid=21466500 |doi= 10.2174/1874473711104010042}}</ref> Alcohol has an effect on the [[Blood-oxygen-level dependent|blood-oxygen-level-dependent]] (BOLD) response. The BOLD response correlates increased blood oxygenation with brain activity, which makes this response a useful tool for measuring neuronal activity.<ref>{{cite journal | author = Jacobus J.|author2=Tapert S. F. | year = 2013 | title = Neurotoxic Effects of Alcohol in Adolescence | journal = [[Annual Review of Clinical Psychology]] | volume = 9 | issue = 1| pages = 703–721 | doi = 10.1146/annurev-clinpsy-050212-185610 | pmc = 3873326 | pmid=23245341}}</ref> The BOLD response affects regions of the brain such as the basal ganglia and thalamus when performing a working memory task. Adolescents who start drinking at a young age show a decreased BOLD response in these brain regions.<ref>{{cite journal | vauthors = Weiland BJ, Nigg JT, Welsh RC, Yau WY, Zubieta JK | displayauthors=etal | year = 2012 | title = Resiliency in adolescents at high risk for substance abuse: flexible adaptation via subthalamic nucleus and linkage to drinking and drug use in early adulthood | journal = Alcohol. Clin. Exp. Res. | volume = 36 | issue = 8| pages = 1355–64 | doi=10.1111/j.1530-0277.2012.01741.x| pmc = 3412943 | pmid=22587751}}</ref> Alcohol dependent young women in particular exhibit less of a BOLD response in parietal and frontal cortices when performing a spatial working memory task.<ref>{{cite journal | vauthors = Tapert SF, Brown GG, Kindermann SS, Cheung EH, Frank LR, Brown SA | year = 2001 | title = fMRI measurement of brain dysfunction in alcohol-dependent young women | journal = Alcohol. Clin. Exp. Res. | volume = 25 | issue = 2| pages = 236–45 | doi=10.1111/j.1530-0277.2001.tb02204.x | pmid=11236838}}</ref> Binge drinking, specifically, can also affect one's performance on working memory tasks, particularly visual working memory.<ref>{{cite journal | vauthors = Ferrett HL, Carey PD, Thomas KG, Tapert SF, Fein G | year = 2010 | title = Neuropsychological performance of South African treatment-naive adolescents with alcohol dependence | journal = Drug Alcohol Depend | volume = 110 | issue = 1–2| pages = 8–14 | doi=10.1016/j.drugalcdep.2010.01.019| pmc = 4456395 | pmid=20227839}}</ref><ref>{{cite journal | vauthors = Crego A, Holguin SR, Parada M, Mota N, Corral M, Cadaveira F | year = 2009 | title = Binge drinking affects attentional and visual working memory processing in young university students | journal = Alcohol. Clin. Exp. Res. | volume = 33 | issue = 11| pages = 1870–79 | doi=10.1111/j.1530-0277.2009.01025.x| pmid = 19673739 | hdl = 10347/16832 | hdl-access = free }}</ref> Additionally, there seems to be a gender difference in regards to how alcohol affects working memory. While women perform better on verbal working memory tasks after consuming alcohol compared to men, they appear to perform worse on spatial working memory tasks as indicated by less brain activity.<ref>{{cite journal | vauthors = Greenstein JE, Kassel JD, Wardle MC, Veilleux JC, Evatt DP, Heinz AJ, Yates MC | year = 2010 | title = The separate and combined effects of nicotine and alcohol on working memory capacity in nonabstinent smokers | journal = [[Experimental and Clinical Psychopharmacology]] | volume = 18 | issue = 2| pages = 120–128 | doi = 10.1037/a0018782 | pmid = 20384423 }}</ref><ref>{{cite journal | vauthors = Squeglia LM, Schweinsburg AD, Pulido C, Tapert SF | year = 2011 | title = Adolescent binge drinking linked to abnormal spatial working memory brain activation: Differential gender effects | journal = Alcoholism: Clinical and Experimental Research | volume = 35 | issue = 10| pages = 1831–1841 | doi = 10.1111/j.1530-0277.2011.01527.x | pmc = 3183294 | pmid=21762178}}</ref> Finally, age seems to be an additional factor. Older adults are more susceptible than others to the effects of alcohol on working memory.<ref>{{cite journal | vauthors = Boissoneault J, Sklar A, Prather R, Nixon SJ | year = 2014 | title = Acute effects of moderate alcohol on psychomotor, set shifting, and working memory function in older and younger social drinkers | journal = Journal of Studies on Alcohol and Drugs | volume = 75 | issue = 5| pages = 870–879 | doi = 10.15288/jsad.2014.75.870 | pmc = 4161706 | pmid=25208205}}</ref>

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== Genetics ==

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== 基因 Genetics ==

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=== Behavioral genetics ===

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=== 行为基因 Behavioral genetics ===

Individual differences in working-memory capacity are to some extent [[heritable]]; that is, about half of the variation between individuals is related to differences in their genes.<ref name=":1">{{Cite journal|last1=Engelhardt|first1=Laura E.|last2=Mann|first2=Frank D.|last3=Briley|first3=Daniel A.|last4=Church|first4=Jessica A.|last5=Harden|first5=K. Paige|last6=Tucker-Drob|first6=Elliot M.|title=Strong genetic overlap between executive functions and intelligence.|journal=Journal of Experimental Psychology: General|volume=145|issue=9|pages=1141–1159|doi=10.1037/xge0000195|pmc=5001920|pmid=27359131|year=2016}}</ref><ref name="Ando 615–624">{{Cite journal|last1=Ando|first1=Juko|last2=Ono|first2=Yutaka|last3=Wright|first3=Margaret J.|title=Genetic Structure of Spatial and Verbal Working Memory|journal=Behavior Genetics|language=en|volume=31|issue=6|pages=615–624|doi=10.1023/A:1013353613591|pmid=11838538|issn=0001-8244|year=2001}}</ref><ref>{{Cite journal|last1=Blokland|first1=Gabriëlla A. M.|last2=McMahon|first2=Katie L.|last3=Thompson|first3=Paul M.|last4=Martin|first4=Nicholas G.|last5=de Zubicaray|first5=Greig I.|last6=Wright|first6=Margaret J.|date=2011-07-27|title=Heritability of Working Memory Brain Activation|journal=Journal of Neuroscience|volume=31|issue=30|pages=10882–10890|doi=10.1523/jneurosci.5334-10.2011|pmid=21795540|pmc=3163233}}</ref> The genetic component of variability of working-memory capacity is largely shared with that of fluid intelligence.<ref name="Ando 615–624"/><ref name=":1" />

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=== Attempts to identify individual genes ===

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=== 识别个别基因的尝试 Attempts to identify individual genes ===

Little is known about which genes are related to the functioning of working memory. Within the theoretical framework of the multi-component model, one candidate gene has been proposed, namely [[ROBO1]] for the hypothetical [[phonological loop]] component of working memory.<ref>{{Cite journal|last=Bates|first=Timothy|date=2011|title=Genetic Variance in a Component of the Language Acquisition Device: ROBO1 Polymorphisms Associated with Phonological Buffer Deficits|journal=Behavior Genetics|volume=41|issue=1|pages=50–7|doi=10.1007/s10519-010-9402-9|pmid=20949370}}</ref>

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== Role in academic achievement ==

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== 在学术成就方面的角色 Role in academic achievement ==

Working memory capacity is correlated with learning outcomes in literacy and numeracy. Initial evidence for this relation comes from the correlation between working-memory capacity and reading comprehension, as first observed by Daneman and Carpenter (1980)<ref>{{Cite journal|title = Individual differences in working memory and reading|journal = Journal of Verbal Learning and Verbal Behavior|date = 1980-08-01|pages = 450–466|volume = 19|issue = 4|doi = 10.1016/S0022-5371(80)90312-6|first1 = Meredyth|last1 = Daneman|first2 = Patricia A.|last2 = Carpenter}}</ref> and confirmed in a later meta-analytic review of several studies.<ref>{{Cite journal|last1=Daneman|first1=Meredyth|last2=Merikle|first2=Philip M.|title=Working memory and language comprehension: A meta-analysis|journal=Psychonomic Bulletin & Review|language=en|volume=3|issue=4|pages=422–433|doi=10.3758/BF03214546|pmid=24213976|issn=1069-9384|year=1996|doi-access=free}}</ref> Subsequent work found that working memory performance in primary school children accurately predicted performance in mathematical problem solving.<ref>{{Cite journal|last1=Swanson|first1=H. Lee|last2=Beebe-Frankenberger|first2=Margaret|year=2004|title=The Relationship Between Working Memory and Mathematical Problem Solving in Children at Risk and Not at Risk for Serious Math Difficulties|journal=Journal of Educational Psychology|volume=96|issue=3|pages=471–491|doi=10.1037/0022-0663.96.3.471}}</ref> One longitudinal study showed that a child's working memory at 5 years old is a better predictor of academic success than IQ.<ref>{{Cite journal|vauthors=Alloway TP, Alloway RG |title=Investigating the predictive roles of working memory and IQ in academic attainment |journal=Journal of Experimental Child Psychology |volume=106|issue=1|pages= 20–9|year=2010|pmid=20018296 |doi=10.1016/j.jecp.2009.11.003|url=https://www.pure.ed.ac.uk/ws/files/11958608/Investigating_the_predictive_roles_of_working_memory_and_IQ_in_academic_attainment.pdf }}</ref>

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== Relation to attention ==

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== 与注意力的关系 Relation to attention ==

There is some evidence that optimal working memory performance links to the neural ability to focus attention on task-relevant information and to ignore distractions,<ref>{{Cite journal|date=March 2009|title=Neural suppression of irrelevant information underlies optimal working memory performance|journal=The Journal of Neuroscience|volume=29|issue=10|pages=3059–66|doi=10.1523/JNEUROSCI.4621-08.2009|pmc=2704557|pmid=19279242|author=Zanto, T. P.|author2=Gazzaley, A.}}</ref> and that practice-related improvement in working memory is due to increasing these abilities.<ref>{{cite journal|last2=Zanto|first2=T. P.|last3=Rutman|first3=A. M.|last4=Clapp|first4=W. C.|last5=Gazzaley|first5=A.|year=2009|title=Practice-related improvement in working memory is modulated by changes in processing external interference|journal=Journal of Neurophysiology|volume=102|issue=3|pages=1779–89|doi=10.1152/jn.00179.2009|pmc=2746773|pmid=19587320|last1=Berry|first1=A. S.}}</ref> One line of research suggests a link between the working memory capacities of a person and their ability to control the orientation of attention to stimuli in the environment.<ref name="attention09">{{Cite journal|vauthors=Fukuda K, Vogel EK |title=Human variation in overriding attentional capture |journal=The Journal of Neuroscience |volume=29 |issue=27 |pages=8726–33 |date=July 2009 |pmid=19587279 |pmc=6664881 |doi=10.1523/JNEUROSCI.2145-09.2009}}</ref> Such control enables people to attend to information important for their current goals, and to ignore goal-irrelevant stimuli that tend to capture their attention due to their sensory [[salience (neuroscience)|saliency]] (such as an ambulance siren). The direction of attention according to one's goals is assumed to rely on "top-down" signals from the pre-frontal cortex (PFC) that biases processing in [[posterior cortex|posterior cortical areas]].<ref>{{Cite journal|vauthors=Desimone R, Duncan J |title=Neural mechanisms of selective visual attention |journal=Annual Review of Neuroscience |volume=18 |pages=193–222 |year=1995 |pmid=7605061 |doi=10.1146/annurev.ne.18.030195.001205}}</ref> Capture of attention by salient stimuli is assumed to be driven by "bottom-up" signals from subcortical structures and the primary sensory cortices.<ref>{{Cite journal|vauthors=Yantis S, Jonides J |title=Abrupt visual onsets and selective attention: voluntary versus automatic allocation |journal=Journal of Experimental Psychology. Human Perception and Performance |volume=16 |issue=1 |pages=121–34 |date=February 1990 |pmid=2137514 |url=http://content.apa.org/journals/xhp/16/1/121 |doi=10.1037/0096-1523.16.1.121|citeseerx=10.1.1.211.5016 }}</ref> The ability to override "bottom-up" capture of attention differs between individuals, and this difference has been found to correlate with their performance in a working-memory test for visual information.<ref name="attention09" /> Another study, however, found no correlation between the ability to override attentional capture and measures of more general working-memory capacity.<ref>{{Cite journal|last1=Mall|first1=Jonathan T.|last2=Morey|first2=Candice C.|last3=Wolff|first3=Michael J.|last4=Lehnert|first4=Franziska|date=2014-01-09|title=Visual selective attention is equally functional for individuals with low and high working memory capacity: Evidence from accuracy and eye movements|journal=Attention, Perception, & Psychophysics|language=en|volume=76|issue=7|pages=1998–2014|doi=10.3758/s13414-013-0610-2|pmid=24402698|issn=1943-3921|url=http://orca.cf.ac.uk/105362/1/Morey.%20Visual%20selective.pdf}}</ref>

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== Relationship with neural disorders ==

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== 与神经系统疾病的关系 Relationship with neural disorders ==

An impairment of working memory functioning is normally seen in several neural disorders:

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