Short-Term Memoryq

G Vallar, University of Milan-Bicocca, Milan, Italy; and IRCCS Istituto Auxologico Italiano, Milan, Italy
Ó 2017 Elsevier Inc. All rights reserved.

Introduction 2
Functional Architecture 2
Behavioral Studies in Healthy Participants 2
Short-Term Forgetting 2
Immediate Serial Recall 3
Immediate Recall and the Recency Effect 4
STM Versus LTM: Short-Term Forgetting, Recency, and Span 4
Neuropsychological Evidence From Brain-Damaged Patients 8
Impairments of Phonological STM 8
Impairments of Spatial and Visual STM 12
Neurological Architecture 13
Phonological STM 13
Visual and Spatial STM 15
Short-Term Memory and Cognition 16
Long-Term Learning 16
Language Processing 17
Sentence Comprehension 17
Speech Production 17
Spatial Orientation and Navigation 17
Conclusions 17
Acknowledgments 18
References 18

Glossary
Articulatory suppression A paradigm whereby participants
are required to utter an irrelevant speech sound (eg, “the,
the, the”), while engaged in another task, such as the short-
term retention of a list of words. This concurrent
articulatory activity disrupts the process of verbal rehearsal,
and has little, if any, general interference effects.
Buffer/store A limited capacity system, concerned with
temporary retention; with reference to “short-term
memory” (STM), the term is frequently used as
a synonymous of “store.”
Double dissociation A pattern of impairment whereby:
(1) a patient or a group of patients exhibits a defective
performance in one task “A” (with respect to healthy
participants), but not in another task “B,” in which
performance level is within the normal range; (2)
another patient or group shows an opposite disorder
(impairment in task “B,” normal performance in task
“A”). In its “strong” or “classical” form the double
dissociation concerns tasks of comparable difficulty. The
double dissociation allows the inference that two
functions (eg, “short-” and “long-” term memory, STM,
LTM) are independent. If this pattern of impairment is
associated with different neuropathological correlates
(eg, defective performance in task “A,” but not in “B,”
in patients with parietal damage, impaired performance
in task “B,” but not in “A,” in patients with frontal
damage), the double dissociation is anatomo-
functional. The same logic may be applied to
neuroimaging activation, and to invasive and non-
invasive brain stimulation (NIBS) paradigms.
Recency effect A phenomenon whereby the final events in
a series are recalled better than the preceding ones; in
immediate free recall the recency effect reflects retention
in an STM system.
Rehearsal A process that supports temporary retention in
a “short-term store” (STS), reviving a memory trace;
rehearsal may involve translation between two different
representations or codes.
Similarity effect A phenomenon, whereby recall of lists
of items similar for some specific coding feature (eg,
phonological, visual, semantic similarity) differs from
that of lists including items dissimilar in that
particular feature. In STM paradigms similarity effects
typically disrupt performance, and provide indications
as to the representational format of the stored
material.

q
Change History: February 2016. G Vallar added abstract and Table 1, made some changes to the text, updated the figures and Further Reading section of the
entire article.

Reference Module in Neuroscience and Biobehavioral Psychology http://dx.doi.org/10.1016/B978-0-12-809324-5.03170-9 1

2 Short-Term Memory

Span A classical measure of STM. Participants are required
to recall a sequence of events, such as a list of digits, words,
visual patterns, or locations of objects, immediately after
presentation, and in the same order.
Store A store is a component of STM, with limited capacity,
which secures the temporary retention of a limited amount
of material (see also “buffer”).
Trace A trace refers to the event stored in memory, and is
frequently used in a physiological context.
Unattended speech Unattended speech is an interfering
auditory stimulation consisting of phonological material
presented to the participant, who is engaged in another
task, such as immediate memory span.
Working memory The term “working memory” (WM) is
widely used in the domain of cognitive psychology,
neuropsychology, and cognitive neuroscience. In a restricted
sense, WM refers to limited capacity systems involved in
temporary retention (STSs), but also emphasizing the
cognitive operations performed on the stored material. The
concept of WM has widened, however, to include the efficient
monitoring and coordination of mental activity during the
simultaneous execution of two or more tasks, the planning of
complex actions, and, more generally, the organization of
behavior. In the latter, broader, sense, the relationships of the
term WM with its original roots in the specific domain of STM
and memory in general are less close.

Introduction

The term “short-term memory” (STM) refers to a number of memory systems, with limited capacity, concerned with the temporary
(in the range of seconds) retention of a variety of materials. This article will present current knowledge concerning the functional
and anatomical organization of STM in humans, with a multidisciplinary approach. Behavioral, functional neuroimaging activa-
tion, and NIBS studies in healthy human participants, engaged in tasks assessing short-term retention, as well as neuropsychological
evidence from patients with brain damage will be considered. The article will be concerned with the more extensively investigated
aspects of STM: Verbal and visuo-spatial. Many of the data come from studies in the adult, but some pertinent evidence from
different populations (children, elderly participants) will be also mentioned.
The general idea that the faculty of memory is not unitary and monolithic, comprising instead multiple systems, dates back at
least to the “Essay concerning human understanding” by the British philosopher John Locke (1700), who suggested a broad distinc-
tion between two types of memory, one concerned with temporary retention and the other as a store-house of materials which have
been laid out of sight. This distinction between two memory systems, one shorter- and one longer-term with respect to the duration
of the trace, was revived by the North American psychologist William James (1890), who suggested the existence of a limited
capacity “Primary Memory,” embracing the present and the immediate past, and supporting consciousness. Psychological research
in the XIX century, and in the first half of the XX, was however mainly concerned with the diverse factors affecting learning and
retention, in the context of a basically unitary view of human memory. It was only in the 1950s that STM became the object of
systematic scientific investigation in healthy participants. Fig. 1 shows a model of STM and LTM, popular in the 1970s (Atkinson
and Shiffrin, 1971).

Functional Architecture
Behavioral Studies in Healthy Participants
The main empirical evidence suggesting the existence of a discrete limited-capacity system, concerned with short-term retention, was
provided in the 1950s and 1960s by the investigation in healthy human participants of three behavioral phenomena, which subse-
quently provided reliable paradigms for exploring also the behavior of pathological populations (Baddeley, 1976). The vast
majority of empirical studies focused on verbal material, such as words or letters, and, therefore, on the hypothetical construct
of “verbal” STM. Evidence was also provided, however, for the existence of a separate visual STS. Examples of stimuli used to inves-
tigate short-term retention are shown in Fig. 2.

Short-Term Forgetting
In the typical paradigm the participants’ task is to recall or recognize a few items after short time intervals (in the range of
seconds), filled by interfering activity. The accuracy of recall of a short list of stimuli (eg, trigrams of consonants or of words)
decreases dramatically in a few (less than 10) seconds if the participants’ repetition of the memory material (rehearsal), which
would prevent forgetting (for review and discussion of the mechanisms involved in short-term forgettingddecay of the
memory trace, as initially thought, interferencedsee Baddeley, 1976; Jonides et al., 2008) is prevented by a distracting activity
as counting backwards by threes (Fig. 3). A broadly similar forgetting occurs for visual and spatial (eg, the location of a dot,
random patterns), and auditory nonverbal stimuli (eg, a tone). This rapid short-term forgetting may be contrasted with the
long-lasting persistence in memory of other types of records, such as autobiographical events, which may last for years, up
to the entire life.
 Short-Term Memory 3

SENSORY REGISTERS SHORT-TERM STORE

(temporary
Visual working memory)
Environmental LONG-TERM STORE
Auditory CONTROL (permanent
input
PROCESSES memory store)
Haptic #rehearsal
#coding
... #decisions
#retrieval strategies

RESPONSE OUTPUT

Figure 1 Short-term and long-term memory. This flow-chart illustrates a functional architecture of human memory proposed in the late 1960s by
Richard Atkinson and Richard Shiffrin, which summarizes the main features of a number of models put forward at that time. The model draws
a distinction between short- and long-term memory systems (STM/LTM). The two components are organized serially, with temporary storage in STM
being a necessary condition for retention in LTM. The short-term store (STS) is a unitary system, which is not specific for sensory modality,
receiving input from different sensory registers, and includes a number of control processes, such as rehearsal. This store plays a central role in
cognitive activity, being equated with consciousness. Source: Redrawn from Atkinson, R.C., Shiffrin, R.M., 1971. The control of short-term memory.
Sci. Am. 225, 82–90.

(A) (B) (C)

(D) (E) (F)

(G) F, K, Q, R, X, W, Z
COW, DAY, BAR, FEW, HOT, PEN, CUP
(H) B, C, D, G, P, V, T
MAD, MAN, MAT, CAD, CAN, CAT, CAP

(I) SUM, HATE, HARM, WIT, BOND, YELD,

(J) ASSOCIATION, OPPORTUNITY, REPRESENTATIVE,
ORGANIZATION, CONSIDERABLE, IMMEDIATELY

Figure 2 Stimuli used in short-term memory tasks. (A–C) Patterns used to assess visual STM. (D–F) Stimuli used to assess visuo-spatial STM
(location of a dot in a square). (G–J) Stimuli used to assess verbal STM: phonologically dissimilar (G) and similar (H) sets of letters and words; short
(I) and long (J) sets of words. Sources: (A–C) Redrawn from Phillips, W.A., 1974. On the distinction between sensory storage and short-term
memory. Percept. Psychophys. 16, 283–290; (D–F) Reproduced from Dale, H.C., 1973. Short-term memory for visual information. Br. J. Psychol. 64,
1–8; (G and H set) From Vallar, G., Baddeley, A.D., 1984. Fractionation of working memory: neuropsychological evidence for a phonological short-
term store. J. Verbal Learn. Verbal Behav. 23, 151–161; Baddeley, A.D., 1966. Short-term memory for word sequences as a function of acoustic,
semantic and formal similarity. Q. J. Exp. Psychol. 18, 362–365; (I and J sets) From Baddeley, A.D., Thomson, N., Buchanan, M., 1975. Word length
and the structure of short-term memory. J. Verbal Learn. Verbal Behav. 14, 575–589.

Immediate Serial Recall

The ability of healthy participants to recall in their presentation order a sequence of events (memory span, review in Unsworth and
Engle, 2007), such as a list of digits, letters, words, a sequence of locations of objects, is limited (George Miller’s “magical” number
7  2 items, 1956). As for short-term forgetting, if a succession of different lists is presented, participants become unable to
remember the previous ones in a few minutes.
4 Short-Term Memory

100

CONTROL (CCC)
75 COUNTING BACKWARDS (CCC)
PATIENT PV (C)

PERCENT CORRECT
50

25

0
0 5 15
DELAY (sec)

Figure 3 Short-term retention of individual letters. Healthy participants: Control (recall of consonant trigrams “CCC” with no interpolated distracting
activity); Counting backwards (recall of CCC with interpolated distracting activity). Patient PV: recall of single letters “C” with no interpolated distracting
activity. In healthy participants the counting task has dramatic detrimental effects on short-term retention of CCC sequences. In the patient, who has an
auditory span of 2.5 digits, a C stimulus is forgotten in a few seconds, even under conditions of no interference. The errorless performance at the 0 s
delay rules out the possibility that the patient’s deficit reflects a perceptual impairment. Source: Data from Basso, A., Spinnler, H., Vallar, G., Zanobio,
M.E., 1982. Left hemisphere damage and selective impairment of auditory verbal short-term memory. A case study. Neuropsychologia 20, 263–274.

Immediate Recall and the Recency Effect

Healthy participants, presented with a sequence of events, exceeding their immediate memory span, such as a list of words or of
visual patterns, which they are required to recall in any order they wish (“free recall”), produce first and best the final, more recent,
stimuli. A comparatively minor advantage of the first one-two items is also present, giving to the recall curve a non-symmetrical, U-
shaped, form. A similar curve occurs when the participant produces the stimuli of the list in the presentation order (“serial recall”).
The recall superiority of the final items, a phenomenon termed the “recency effect” (Fig. 4), vanishes after a few seconds of
distracting activity. In a typical free recall experiment with verbal material, such as a list of 10–15 words, the recency effect
involves the final five to seven stimuli. The recency effect for verbal stimuli is minimally influenced by factors such as age, rate
of presentation, and word frequency, which, in turn, affect recall performance in the pre-recency positions (Glanzer, 1972). This
pattern is taken as evidence for separate STM and LTM processes, with the former being associated with the recency effect, and
the latter with recall in the preceding positions.

STM Versus LTM: Short-Term Forgetting, Recency, and Span

These three sets of observations (short-term forgetting, the recency effect in free recall, immediate memory span) illustrate the main
characteristics of STM: A retention system with limited capacity, able to hold recent (in the time range of seconds) events, in which
the memory trace decays in a few seconds, unless this is prevented through rehearsal.
The material stored in STM has a specific representational format. In the verbal domain this involves phonological codes, which
may be distinguished from the lexical-semantic representations stored in the semantic (for the original distinction between
“semantic” and “episodic” memory, see Tulving, 1972) component of LTM (for recent reviews and discussion, see Lambon Ralph,
2013; Zannino et al., 2015). In immediate verbal span, the participants’ performance is primarily affected by factors such as phono-
logical similarity and word length (see Fig. 5A). The phonological similarity effect is illustrated by the superior immediate serial
recall of sequences of dissimilar letters or words, compared to lists comprising similar items (Fig. 2, “G” vs. “H” sets). The word
length effect is illustrated by the superior immediate serial recall of sequences of short words, compared to lists comprising longer
items (Fig. 2, “I” vs. “J” sets).
The three paradigms of short-term forgetting, immediate memory span, and free recall provided the initial impetus to studies in
healthy participants, aimed at dissociating STM versus LTM processes. They also provided the basic tools for assessing this putative
dissociation in brain-damaged patients.
Each of these three phenomena subsequently proved to be considerably more complex than initially thought.
a. In short-term forgetting the counting backwards task has more complex effects than the mere disruption of the process of rehearsal,
which prevents decay of the short-term trace in the first few seconds after stimulus presentation (Vallar and Baddeley, 1982).
 Short-Term Memory 5

SHORT-TERM MEMORY

(A) AUDITORY INPUT (B) VISUAL INPUT

100 PV 100 PV
90 90
C C
80 80
70 70
Percent Correct

Percent Correct
60 60
50 50
40 40
30 30
20 20
10 10
0 0
1 2 3 4 5 6 7 8 9 10 11 12 1 2 3 4 5 6 7 8 9 10 11 12

SERIAL POSITION SERIAL POSITION

Figure 4 Immediate free recall of sequences of words. Average free recall performance with lists of 12 Italian words by patient PV (see caption to
Fig. 3) and unimpaired participants (C) (s.d.), by serial position of each stimulus in the presentation order, and modality of presentation [auditory (A),
visual (B)]. In this task participants are presented with lists of words, which they subsequently recall in any order they wish. Unimpaired participants
show a higher level of performance in the final positions of the list (recency effect), with recall accuracy steadily increasing in the last six serial posi-
tions (in this particular sequence from about the 6th to the final 12th stimulus). The recency effect is present with both auditory and visual presenta-
tion of the stimuli, and an auditory advantage (“modality effect”) is typically found. Unimpaired participants also show a minor advantage of the initial
one to two positions in the sequence. In immediate free recall, the recency effect reflects the output of STM systems, while recall from the preceding
positions is mainly based on LTM systems. With auditory presentation of the stimuli patient PV (see caption to Fig. 3) shows no recency, but this is
within the normal range with visual input. Patient PV also shows a preserved recall performance in the initial and middle positions of the recall
sequence. Source: Redrawn from Vallar, G., Papagno, C., 1986. Phonological short-term store and the nature of the recency effect: evidence from
neuropsychology. Brain Cogn. 5, 428–442.

In addition, subsequent forgetting involves other mechanisms, such as interference (Baddeley, 2003; Jonides et al., 2008). The
processes which maintain items for short time intervals may also involve a contribution from LTM processes, such as elaborative
rehearsal (Craik and Watkins, 1973), and redintegration of a short-term trace, on the basis of features such as the frequency of the
stimulus (Hulme et al., 1999).
b. Immediate memory span is influenced by lexical, syntactic, and semantic factors, which reflect LTM processes (Hurlstone et al.,
2014; Thorn and Page, 2009). Span is greater for words, compared to nonwords (Brener, 1940; Hulme et al., 1999), and for high
frequency than for low frequency words (Roodenrys et al., 2002; Watkins, 1977). Span is also greater for concrete than for
abstract words (Acheson et al., 2010; Campoy et al., 2015).
c. The recency effect is a general memory phenomenon, which may be observed also under conditions involving only LTM
processes, such as recall of remote autobiographical events. Furthermore, factors such as the relative temporal interval among the
events in the sequence influence the occurrence of the recency effect in long-term retention (Moreton and Ward, 2010).
Initially, in the early 1960s, attempts were made to explain the three phenomena (short-term forgetting, free recall, span) in
terms of the then dominant view of human memory as a unitary system, with a continuum between STM and LTM (Melton,
1963; Ruchkin et al., 2003, and commentaries therein). Such a monolithic account proved however untenable, mainly on the basis
of neuropsychological evidence from patients with brain damage (Shallice and Vallar, 1990; Vallar and Papagno, 2002), discussed
in the following section.
The functional architecture of the system concerned with the short-term retention of verbal material (“phonological short-term
memory,” PhSTM) has been investigated in further detail through the effects of phonological similarity and word length, and the
concurrent task of articulatory suppression, teasing apart “storage” versus “rehearsal” components. Suppression has three effects on
immediate retention in span tasks:
a. it brings about an overall reduction of memory performance;
b. it abolishes the effect of phonological similarity when the stimuli are presented visually, but not with auditory input;
a. it eliminates the effect of word length both with auditory and with visual presentation of the stimuli.
6 Short-Term Memory

(A) CONTROL SUBJECTS. AUDITORY INPUT

PHONOLOGICAL SIMILARITY EFFECT WORD LENGTH EFFECT
PERCENT PERCENT
CORRECT CORRECT
80 80

70 DISSIMILAR 70 SHORT

60 SIMILAR 60 LONG

50 50

40 40

30 30

20 20

10 10

0 0

LIST LENGTH 6 ITEMS LIST LENGTH 6 ITEMS

(B) PATIENT PV. AUDITORY INPUT

PHONOLOGICAL SIMILARITY EFFECT WORD LENGTH EFFECT
SHORT
LETTER SPAN WORD SPAN
4,5 4,5 LONG

4 DISSIMILAR 4

3,5 SIMILAR 3,5

3 3

2,5 2,5
2 2
1,5 1,5
1 1

0,5 0,5
0 0
(C) PATIENT PV
PHONOLOGICAL SIMILARITY EFFECT VISUAL INPUT
LETTER SPAN
5

4,5
DISSIMILAR
4

3,5 SIMILAR

2,5

1,5

0,5

0
BASELINE SUPPRESSION
 Short-Term Memory 7

Figure 6 A functional model of phonological short-term memory. Auditory-verbal material, after early acoustic and phonological analysis, (1) enters
the main retention component of the system, the “phonological short-term store” (PhSTS) (2), where material is coded in a phonological format. The
PhSTS is an input-system, to which auditory material has a direct and automatic access. The process of articulatory “Rehearsal” (ArReh) is
conceived as involving a recirculation of the memory trace between the PhSTS and a phonological output system, the “Phonological Output Buffer”
or “Phonological Assembly System” (3), primarily concerned with the articulatory programming of speech output, with a recurring translation
between input (“acoustic”) and output (“articulatory”) phonological representations. The Phonological Output Buffer (ArReh) provides also access of
visually presented verbal material to the PhSTS, after “phonological recoding” or “grapheme-to-phoneme conversion” (5). The model also illustrates
the multiple-component nature of STM, showing a visual STS (4), where material is likely to be encoded in terms of shape. Source: Redrawn from
Vallar, G., Di Betta, A.M., Silveri, M.C., 1997. The phonological short-term store-rehearsal system: patterns of impairment and neural correlates. Neu-
ropsychologia 35, 795–812.

This pattern suggests a fractionation of PhSTM into two components, both contributing to immediate retention in span tasks:
a. a passive input system, the “Phonological Short-Term Store” (PhSTS), to which auditory input has a direct access; this
phonological system is non-articulatory in nature, as witnessed by the persistence of the effect of phonological similarity during
suppression;
b. a more controlled process of “Articulatory Rehearsal” (ArReh), which has the two-fold function of refreshing the phonological
trace, preventing its decay, and of conveying visual-verbal information to the PhSTS, after Phonological Recoding (PhRec), or
grapheme-to-phoneme conversion.
Articulatory suppression, interfering with the operation of ArReh, reduces span performance, and prevents access of visual-verbal
material to the PhSTS, as witnessed by the disappearance of the effect of phonological similarity. The word length effect, being abol-
ished by suppression, is likely to reflect articulatory factors, such as the spoken duration and complexity of the stimuli, and may be
taken as an index of the activity of the ArReh process. A model of PhSTM is shown in Fig. 6, based on Vallar et al. (1997).

=
Figure 5 Coding in phonological short-term memory. (A) In immediate serial recall the memory performance of healthy participants is better when
the stimuli are phonologically dissimilar rather than similar (phonological similarity effect), and shorter rather than longer (word length effect).
Healthy participants exhibit these effects with both auditory and visual presentation of the stimuli. For each effect, the recall performance of healthy
participants with 6-item auditory sequences of similar and dissimilar letters, long (5-syllable) and short (2-syllable) words is shown. These effects are
taken as evidence of phonological coding in verbal STM. (B) Patient PV (see caption to Fig. 3) shows a dramatic reduction of auditory span perfor-
mance (less than 2.5 letters) and a preserved effect of phonological similarity, which indicates that auditory-verbal material is, as in healthy partici-
pants, coded in a phonological format, even though the capacity of the PhSTS is dramatically reduced. The absence of any effect of word length with
auditory input indicates that the process of ArReh is not used by PV. (C) Patient PV, at variance from healthy participants, shows neither the detri-
mental effect of phonological similarity on serial span with visual presentation of the letter stimuli, nor the detrimental effect of articulatory suppres-
sion. This pattern indicates that PV does not convey visually presented verbal material to the PhSTS via ArReh, for the purpose of short-term
retention, after recoding it into a phonological format. As the visually presented stimuli are stored by PV in a non-phonological format, presumably
visual, articulatory suppression, that engages ArReh, does not affect PV’s span performance. Source: Data from Vallar, G., Baddeley, A.D., 1984.
Fractionation of working memory: neuropsychological evidence for a phonological short-term store. J. Verbal Learn. Verbal Behav. 23, 151–161.
8 Short-Term Memory

In line with the PhSTS/ArReh fractionation illustrated in Fig. 6, unattended auditory speech disrupts immediate memory span for
verbal material, presented in both the visual and the auditory modality, preempting the storage capacity of the PhSTS. Suppression,
however, abolishes the disruptive effect of unattended speech on immediate retention of verbal material presented in the visual
modality. This effect takes place because suppression, interfering with the operation of ArReh, prevents visual-verbal material from
entering the PhSTS, corrupted by unattended speech (Salamé and Baddeley, 1982). Finally, some perceptual “phonological judgments”
are held to specifically involve the ArReh process, because the performance of healthy participants is slightly, but significantly, impaired
by suppression. The term “phonological judgment” refers to the conscious processing (“phonological awareness”) of a number of
phonological aspects of strings of letters, either words or nonwords. In most studies the material is presented visually, in order to prevent
the direct activation of acoustic-phonological representations. These judgments include deciding whether or not two letter strings have
identical initial sounds, or two words have stress in the same position (Burani et al., 1991; Vallar and Baddeley, 1984b).
In the non-verbal domain a similar distinction is drawn between visual and spatial STM and LTM systems, with the relevant
representational format being in terms of the spatial location or of the shape of the stimulus (Logie and Della Sala, 2005; Repovs
and Baddeley, 2006; Zimmer, 2008). Spatial and visual STM systems are likely to comprise storage and rehearsal (spatial and picto-
rial) components. In the case of spatial locations, rehearsal may be conceived in terms of planned movements (eg, ocular, manual
reaching, locomotion) towards a target coded in a spatial reference frame (eg, egocentric), although this may be not the only
possible format involved.
The relationships between knowledge stored in LTM and STM systems are schematically depicted in Fig. 7.

Neuropsychological Evidence From Brain-Damaged Patients

Studies in patients with memory disorders associated with brain damage support the independence of STM and LTM systems,
conjuring up a double dissociation of deficits. The hallmark of “global amnesia” (see the paradigmatic case of patient HM, Corkin,
2002), which is interpreted as a selective impairment of the declarative, or explicit, component of LTM, is the inability to learn
verbal and visuo-spatial material, as assessed by recall or recognition tasks (“anterograde amnesia”). The deficit of these patients
concerns also memory for past autobiographical events and facts, occurred prior to the onset of the disease (“retrograde amnesia”)
(Drachman and Arbit, 1966; Milner et al., 1968; Penfield and Milner, 1958; Scoville and Milner, 1957). This dramatic memory
deficit, which disrupts the patients’ everyday life, contrasts sharply with their unimpaired performance in the three paradigms
mentioned earlier. Short-term forgetting and immediate memory span are within the normal range, for both verbal and non-
verbal material (Drachman and Arbit, 1966; Sidman et al., 1968; Wickelgren, 1968). In immediate free recall, amnesic patients
show a preserved recency effect, associated with a defective performance in the pre-recency positions of the list (Baddeley and
Warrington, 1970). These converging results suggest a functional characterization of amnesia in terms of defective LTM, with
a preserved function of STM. Secondly, the pattern of impairment of amnesia is compatible with a serial organization of the
two systems, with temporary retention in STM being a necessary condition for long-term storage (see Fig. 1).
Since the late 1960s selective impairments of different components of STM have been reported. The more extensively investi-
gated areas concern PhSTM (or “auditory-verbal” STM, as the system was initially termed (Warrington and Shallice, 1969)), spatial
and visual STM.

Impairments of Phonological STM

Neuropsychological deficits of PhSTM comprise two different types of impairment: (1) a reduced capacity of the “store” component
of the system; (2) a dysfunction of ArReh.

Figure 7 LTM effects on STM systems. For spatial and visual STM, LTM effects are based on objects’ lexicon, visual semantics and spatial refer-
ence frames; for PhSTM, on lexical, semantic, and syntactic knowledge. Possible effects from “Episodic Memory” are not shown. Sources: Repro-
duced from Baddeley, A.D., 2003. Working memory: looking back and looking forward. Nat. Rev. Neurosci. 4, 829–839; Heyden, A., Sparr, G.,
Nielsen, M., Johansen, P. (Eds.), 2002. Computer Vision – ECCV 2002. In 7th European Conference on Computer Vision Copenhagen, Denmark, May
28–31, 2002 Proceedings, Part IV. Springer-Verlag, Berlin.
 Short-Term Memory 9

Disproportionately Reduced Capacity of the Phonological Short Term Store

Although it had long been known that patients with dysphasia may show a disproportionate impairment of repetition of verbal
material, such as a sentence, or a list of words or digits, it was only in the late 1960s that this deficit was interpreted as a specific
disorder of “auditory-verbal” STM (Warrington and Shallice, 1969), rather than as a linguistic impairment (“conduction aphasia,”
review in Ardila, 2010). The hallmark of this STM disorder is a selective and disproportionate deficit of immediate repetition of all
kinds of verbal material (digits, letters, words, sentences). Patients with this STM deficit, engaged in the three tasks discussed earlier,
exhibit this pattern of performance:
a. short-term forgetting is abnormally rapid (Fig. 3);
b. the recency effect in immediate free recall of auditory lists of words is abolished, or severely reduced (Fig. 4);
c. auditory-verbal span is disproportionately reduced, to an average of less than three items (digits, letters, or words, see Figs. 5B
and 8).
The retention deficit is not explained by impaired sensory or perceptual analysis. The patients’ immediate reproduction of indi-
vidual items is entirely spared, as well as their ability to perform tasks requiring phonological discriminations, under conditions of
minimal memory load. Speech production is also entirely preserved in a number of patients, ruling out interpretations in terms of
defective response (articulatory) processes (Shallice and Vallar, 1990; Vallar and Papagno, 2002). In addition, the memory deficit
does not improve when a response modality not requiring speech production (eg, recognition by pointing) is being used. These
features distinguish this “mnestic” disorder of repetition from a type of conduction aphasia (Buchsbaum et al., 2011), in which
both the repetition deficit and spontaneous speech are characterized by phonological errors (“phonemic paraphasias”). Taken
together, these findings suggest the impairment of an STM system.
In the vast majority of these patients the disorder is modality-specific, with level of performance being better when the material is
presented visually (see Fig. 8). This input-related dissociation has two main implications:
a. discrete phonological and visual STM components exist;
b. in the input–output processing chain, the PhSTS should be conceived as an “input,” perceptual, system, rather than as an
“output” production buffer store.
This fractionation also argues against a monolithic view of the system as a single “amodal” store, which is not specific for the
different sensory modalities. Additional support to this input locus of the system comes from the observation, mentioned earlier,
that in some of these patients the production of articulated speech is entirely preserved (Shallice and Butterworth, 1977; Vallar and
Baddeley, 1984a).
Patients with defective PhSTM may show unimpaired long-term verbal (eg, paired-associate learning, Fig. 9; learning of
a short story) and visuo-spatial (eg, visual maze learning) memory. This observation supports the notion of a complete inde-
pendence of STM and LTM systems, and is incompatible with a serial organization, in which defective STM entails an LTM
impairment (see Fig. 1), suggesting instead a parallel architecture: After early perceptual analysis, information may enter

AUDITORY SPAN DIGIT SPAN

6
DIGITS DIGITS

5 LETTERS

4 WORDS

Figure 8 Auditory and visual span in patients with defective auditory-verbal (phonological) STM. Repetition span for all types of auditory-verbal
material (digits, letters, words) is defective, while level of performance is higher with visual presentation. In healthy participants, by contrast, a minor
advantage of the auditory input modality is typically found (“modality effect”). The histograms indicate average spans, the vertical bars the ranges of
the patients’ span performances, the horizontal dashed line the cut-off for auditory digit span. Source: Meta-analytic data from Vallar, G., Papagno,
C., 2002. Neuropsychological impairments of verbal short-term memory. In: Baddeley, A., Wilson, B., Kopelman, M. (Eds.), Handbook of Memory
Disorders. Wiley, Chichester, England, pp. 249–270.
10 Short-Term Memory

Figure 9 Long-term verbal learning in a patient with defective PhSTM. Patient PV (see caption to Fig. 3) shows unimpaired paired-associate
learning of Italian words (C: control participants). In a paired-associate learning paradigm, participants are presented with an auditory list of pairs of
stimuli (eg, unrelated words, such as “dog”d“pint,” “hammer”d“truth”). Immediately after presentation, or after a delay, participants are presented
with the first member of each pair, in an order different from presentation, and are required to provide the second member (eg, “hammer”. /
“truth,” “dog”. / “pint”). Source: Redrawn from Baddeley, A.D., Papagno, C., Vallar, G., 1988. When long-term learning depends on short-term
storage. J. Mem. Lang. 27, 586–595.

STM or LTM, either of which may be selectively disrupted by brain damage. The learning abilities of these patients are however
dramatically impaired when the phonological material to be learned does not possess pre-existing lexical-semantic represen-
tations in LTM. This is the case of pronounceable meaningless sequences of letters: nonwords, or words in a language unknown
to the participant (eg, “svieti,” “tevome,” Russian words, transliterated into Italian, in which language they are nonwords, to be
learned by an Italian participant) (Fig. 10). This deficit of phonological learning indicates that, within a specific representa-
tional domain, temporary storage in STM is necessary for a stable long-term learning and retention, as predicted by the serial
model illustrated in Fig. 1.

Impairment of Articulatory Rehearsal

The process of ArReh has been traditionally described as an activity, which, through rote repetition, refreshes the STM trace, prevent-
ing its decay. ArReh may be conceived in terms of the recoding of the memory trace from input (“auditory-verbal”) to output-related
(“articulatory”) representations and vice-versa (see Fig. 6). More specifically, rehearsal of verbal material has long been regarded as
an “articulatory” activity (“inner” or “internal” speech), which involves output-related verbal processes, including the motor
programming of speech production in a “Phonological Assembly System” or “Phonological Output Buffer” (see Fig. 6), the actual
articulation of the material to be rehearsed, making use of the peripheral musculature (“subvocal” rehearsal), or both. Anarthric
patients, who, due to congenital disorders (cerebral palsy, Bishop and Robson, 1989), or brain damage acquired in adult age
(Schnakers et al., 2008; Vallar and Cappa, 1987), are unable to utter any articulated speech sound, may however show an entirely
preserved PhSTM, including ArReh, as revealed by normal span, similarity and length effects. This suggests that the ArReh process is
“central,” and does not require the activity of the peripheral musculature. Brain-damaged patients with a selective impairment of
ArReh show a defective immediate verbal span, as patients with damage to the PhSTS, because both components of PhSTM
contribute to immediate retention. Damage confined to ArReh (and not involving the PhSTS) disrupts however the ability to
perform “phonological judgments,” such as about stress position on the word and initial sound. This result, together with the above
mentioned interference effects of articulatory suppression found in healthy participants, suggests that some phonological processes,
such as segmentation and deletion, are based on output phonological codes. However, a defective ArReh leaves largely unimpaired
the recency effect in immediate free recall of auditory lists of words (patient TO, Vallar et al., 1997). Conversely, the recency effect
with auditorily presented lists of words is grossly reduced or absent in patients with damage to the PhSTS (Fig. 4). This dissociation
 Short-Term Memory 11

Figure 10 Long-term phonological learning in a patient with defective PhSTM. Patient PV (see caption to Fig. 3) is entirely unable to learn
pronounceable nonwords (Russian words transliterated into Italian, eg,: “orange”d“apielsin,” “bear”d“miedvied”) in a paired-associate word-
nonword paradigm (see caption to Fig. 9) (C: unimpaired control participants). Source: Redrawn from Baddeley, A.D., Papagno, C., Vallar, G., 1988.
When long-term learning depends on short-term storage. J. Mem. Lang. 27, 586–595.

elucidates the mechanisms underlying the recency effect in immediate free recall of lists of auditory stimuli, as based on the capacity
of the PhSTS.
Articulatory suppression, which interferes with ArReh, does not further impair serial recall performance of auditory stimuli by
patients with a selective deficit of this component of PhSTM (patient TO, Vallar et al., 1997). Conversely, suppression reduces audi-
tory serial recall of patients with a selective deficit of the PhSTS, with ArReh being spared (patient LA, Vallar et al., 1997).
In accord with the main role of phonological representations in verbal STM, patients with Semantic Dementia have a largely
preserved level of performance in STM tasks, such as digit span, and show the effects of phonological similarity and word length
(Jefferies et al., 2005). This indicates a basically normal function of the PhSTS-ArReh system, although, support from Semantic
Memory to performance in STM tasks is reduced (Jefferies et al., 2011; Papagno et al., 2013).

Impairment of Phonological Recoding

Phonological recoding (PhRec), though not a “storage” component per se, participates in the operation of the PhSTS-ArReh system,
by converting visual, orthographic, stimuli into a phonological output format (a planned articulatory representation in the
phonological output buffer, see Fig. 6), which allows visually presented verbal material to gain access to ArReh. Patients with
a selective deficit of PhRec, and sparing of the PhSTS-ArReh system, present with a specific pattern of impaired, and preserved
performances:
a. immediate auditory-verbal span is preserved;
b. the phonological similarity and word length effects are present with auditory input, suggesting preserved capacity of the PhSTS
and operation of ArReh;
c. with visual presentation, patients do not exhibit the phonological similarity and word length effects in serial span, indexing no
access of visual-verbal material to PhSTM;
d. the patients’ ability to make phonological judgments is defective.
This pattern suggests that written material fails to enter a preserved ArReh system (as indexed by its normal operation with audi-
tory stimuli), but unavailable to written stimuli, due to the deficit of phonological recoding. Accordingly, unattended speech inter-
ference does not impair visual span performance of patients with defective PhRec; visual span may even improve in such patients,
with an increase of visual confusion errors. A number of single case studies of anarthric patients with a selective impairment of
PhRec have been reported: FC (Cubelli and Nichelli, 1992), MDC (Vallar and Cappa, 1987), and AMB (Papagno et al., 2008).
12 Short-Term Memory

Table 1 Patterns of preserved/defective performance and present/absent effects found following damage to the PhSTS, and to the ArReh
components of PhSTM, and to PhRec

Task
Span PhSE WLE RE
Damaged component A V A V A V PhJ-V A V

PhSTM
PhSTS XX (O) (O/X) X X X O X (O)
ArReh X (O) O X X X X (O) (O)
PhRec O (O) O X O X X # #

A/V, Auditory/visual input; PhS/WL/E, phonological similarity/word length effect; PhJ-V, phonological judgments on visual stimuli; RE, recency effect; O, performance preserved/effect
present; X, performance defective/effect absent; XX, maximal deficit; (), performance level reduced as compared with healthy participants; #, not assessed.

The patterns of defective and impaired performance of patients with selective deficits of the PhSTS, of ArReh, and of PhRec are
summarized in Table 1.

Impairments of Spatial and Visual STM

Two main patterns of STM impairment have been described in brain-injured patients: for spatial locations, and for visual patterns.
The relationships of these disorders with visuo-spatial learning have been also investigated. These deficits have been explored less
extensively than those of PhSTM, and less patients have been described. In addition, the distinction between visual and spatial
component processes in memory is intrinsically blurred, because visual objects have spatial properties, such as their location in
egocentric space, and the spatial relationships between their component parts. Finally, also the distinction between storage and
rehearsal components is less definite.

Impairments of STM for Spatial Locations

These patients show a defective immediate retention of short sequences of spatial locations, as assessed by the Block Tapping Task
(Corsi’s test, a spatial analogue of digit span, see Fig. 11A), with verbal memory being unaffected, in both its STM and LTM
components (Fig. 11B). As for the deficits of PhSTM, the impaired retention of spatial locations cannot be accounted for in terms
of visuo-perceptual or response-related disorders (eg, unilateral spatial neglect, misreaching, see De Renzi et al., 1977; De Renzi
and Nichelli, 1975). These patients may show no evidence of topographical disorientation or impairment in long-term learning of

(A) (B) VISUO-SPATIAL

SERIAL SHORT-TERM MEMORY
SPAN
AUDITORY
8 DIGIT SPAN
8 7 SPATIAL
7 SPAN
6
4 5
5 9 4

3
1 6 2

1
2
3 0
PATIENT 1 PATIENT 2

Figure 11 Impairments of STM for spatial locations. (A) The board of the Block Tapping Test. The patient’s task is to reproduce the sequence of
spatial positions pointed to by the examiner. As for verbal span, sequences of increasing length are presented. In the actual board, numbers are on
the examiner’s side. (B) Patient #1 shows a defective visuo-spatial span, associated with a normal auditory digit span; patient #2 the opposite pattern
of performance. The dashed line indicates the cut-off for digit span, the dotted line for visuo-spatial span. This double dissociation illustrates the
independence of STM for spatial locations from PhSTM. Source: Data from De Renzi, E., Nichelli, P., 1975. Verbal and non-verbal short-term
memory impairment following hemispheric damage. Cortex 11, 341–354.
 Short-Term Memory 13

visuo-spatial information, such as the path of a visual maze. These findings suggest a complete independence of STM and LTM
spatial systems concerned with spatial locations.
Suggestions have been repeatedly made (see eg, Malhotra et al., 2004; Wansard et al., 2014) that deficits of spatial STM, or, more
widely, WM (Striemer et al., 2013; Van Vleet and DeGutis, 2013) may be a component of the syndrome of left spatial unilateral
neglect (review in Vallar and Bolognini, 2014). However, as originally reported by De Renzi et al. (1977, 1975), deficits of spatial
STM may occur in patients without spatial neglect, and, vice versa, patients with spatial neglect may show no impairments of spatial
STM (see eg, Wansard et al., 2015). Also patients with a spatial neglect confined to mental images may show no impairment of
spatial STM span (patient LG, see Piccardi et al., 2008). This pattern of selective deficits conjures up a double dissociation between
impairments of spatial STM, and the core attentional and representational deficit featuring left spatial neglect (Vallar and Bolognini,
2014).
Patients with neurodegenerative disorders such as Alzheimer’s disease show impairments of spatial span as assessed by the Block
Tapping Task, which requires the retention of sequences of targeted movements; these deficits may occur in the absence of impair-
ment of STM for visuo-spatial patterns (Grossi et al., 1993).

Impairments of STM for Visual Patterns

Patients with a selective deficit of visual STM are impaired in pattern span, spatial span being preserved (patient LG, see Wilson
et al., 1999). The disorder of patients with defective visual imagery, as assessed, for instance, by tasks requiring color, size, and
shape comparisons (patient LH, see Farah et al., 1988), has been also interpreted in terms of a defective visual STM/WM (Della
Sala and Logie, 2002). Deficits of STM for visual patterns in patients with Alzheimer’s disease may occur with no impairment of
immediate retention for sequences of spatial locations, as assessed by Corsi’s test (Grossi et al., 1993).
Deficit of visuo-spatial STM may selectively disrupt long-term learning of unfamiliar non-verbal material (ie, new visual infor-
mation), as assessed by recognition memory for unfamiliar faces and objects (patient ELD, see Hanley et al., 1990). This extends
to the visuo-spatial domain the conclusion that long-term acquisition requires STM, when the material is not familiar, such as
new vocabulary or visual patterns.

Neurological Architecture

The investigation of the neural correlates of STM systems takes advantage of three main sources of evidence:
a. the traditional anatomo-clinical correlations between the site and the size of the cerebral lesion and the behavioral impairment;
b. the functional neuroimaging methods, involving the measurement of regional cerebral activity in neurologically unimpaired
participants engaged in behavioral tasks probing the function of the component under investigation;
c. the brain stimulation methods, involving the measurement of behavioral performance under condition of interference with, or
increase of, the activity of cerebral areas and networks involved in the function of interest.

Phonological STM
Correlations performed in patients with defective auditory-verbal span suggest an association between damage to a number of areas
in the left cerebral hemisphere, located around the sylvian fissure, and the dysfunction of PhSTM. The neural correlates of the PhSTS-
ArReh system are shown in Fig. 12. The association posterior parietal cortex (PPC) in the left hemisphere [supramarginal gyrus of
the Inferior Parietal Lobule (IPL), Brodmann’s area, BA 40, at the temporo-parietal junction (TPJ)] represents the main neural
correlate of the STS component of PhSTM. The frontal ventral premotor regions in the left hemisphere (BA 44, BA 6), and other
structures, such as the insula, constitute the major neural correlates of the ArReh component (reviews in Buchsbaum and
D’Esposito, 2008; Vallar and Papagno, 2002). Damage to the right hemisphere does not affect immediate verbal memory,
suggesting a marked left-sided lateralization of the system.
Cerebellar lesions impair immediate retention of verbal material, with disruptive effects, that may particularly affect the ArReh
process (Silveri et al., 1998), or involve more extensively the whole PhSTM system (Ravizza et al., 2006); in any case, the STM deficit
is milder than that caused by left hemispheric cortical lesions (see Fig. 12), and does not extent to visuo-spatial STM. In the light of
the predominantly cerebellar-cerebral contralateral connections suggestions have been made that the right cerebellar hemisphere
may be more involved in language (Jansen et al., 2005) and, specifically, in verbal STM (Silveri et al., 1998; Tomlinson et al.,
2014) functions.
Studies in healthy participants using functional neuroimaging methods concur with this pathological evidence, to suggest
a left-hemisphere-based network (Buchsbaum and D’Esposito, 2008; Jonides et al., 2008; Paulesu et al., 1996). The STS compo-
nent of PhSTM appears associated with activation in the left IPL (supramarginal gyrus, BA 40) at the temporo-parietal junction, the
ArReh component with activation in the left frontal premotor regions (Broca’s area, BA 44; premotor area, BA 6; supplementary
motor area), and in the left insula. In these studies, in line with the behavioral evidence from healthy participants and brain-
damaged patients, an immediate verbal span task activates both the IPL at the TPJ (PhSTS), and the premotor cortex (ArReh),
in the left hemisphere. Conversely, a phonological judgment task selectively activates the left premotor regions, but not the supra-
marginal gyrus.
14 Short-Term Memory

Figure 12 STM systems in the brain: neural correlates. A simplified and schematic summary of the main neural correlates of STM systems in the
two cerebral hemispheres, and in the cerebellum (box). M1, primary motor cortex; S1, primary sensory cortex; CS, central sulcus; IPS, intraparietal
sulcus; STS, superior termporal sulcus; SMG, supramarginal gyrus; AG, angular gyrus; STG, superior temporal gyrus; TPJ, temporo-parietal junction.
(1) The verbal (phonological) STM system (PhSTS and ArReh) is based on neural networks markedly lateralized to the left cerebral hemisphere. The
main neural correlates of the PhSTS component of PhSTM include the left IPL (SMG, BA 40), at the TPJ. The ArReh process’ main neural bases
include Broca’s area (BA 44), the premotor area (BA 6), and the supplementary motor area in the left hemisphere. These two neural STM systems are
likely to be connected through the arcuate fasciculus, the superior longitudinal fasciculus (SLF), and white matter fiber tracts in the insular region.
The PhRec process (“orthographic-to-phonological recoding,” grapheme-to-phoneme conversion) involves activity in the left motor cortex. For the
cerebellum, the dotted lines on the left side denote a possible less definite role. (2) Spatial STM is based on networks including the occipital visual
association cortices, bilaterally, the PPC, the dorsal premotor cortex, and the dorso-lateral prefrontal cortex, with a lateralization to the right cerebral
hemisphere. (3) Visual STM is based on neural networks including the occipital visual association cortices, with a possible greater role of the left
hemisphere, the PPC, the frontal (premotor and prefrontal), and temporal regions, more ventrally with respect to the dorsal regions involved in STM
for spatial locations. Sources: modified from Petrides, M., Tomaiuolo, F., Yeterian, E.H., Pandya, D.N., 2012. The prefrontal cortex: Comparative
architectonic organization in the human and the macaque monkey brains. Cortex 48, 46–57; Kelly, C., Uddin, L.Q., Shehzad, Z., Margulies, D.S.,
Castellanos, F.X., Milham, M.P., Petrides, M., 2010. Broca’s region: linking human brain functional connectivity data and non-human primate tracing
anatomy studies. Europ J. Neurosci. 32, 383–398; Bonnì, S., Perri, R., Fadda, L., Tomaiuolo, F., Koch, G., Caltagirone, C., Carlesimo, G.A., 2014.
Selective deficit of spatial short-term memory: role of storage and rehearsal mechanisms. Cortex 59, 22–32; Courtney, S.M., 2004. Attention and
cognitive control as emergent properties of information representation in working memory. Cogn. Affect. Behav. Neurosci. 4, 501–516; Fletcher, P.C.,
Henson, R.N., 2001. Frontal lobes and human memory: insights from functional neuroimaging. Brain 124, 849–881; Jonides, J., Lewis, R.L.,
Nee, D.E., Lustig, C.A., Berman, M.G., Moore, K.S., 2008. The mind and brain of short-term memory. Annu. Rev. Psychol. 59, 193–224; Papagno,
C., Lucchelli, F., Vallar, G., 2008. Phonological recoding, visual short-term store and the effect of unattended speech: evidence from a case of slowly
progressive anarthria. Cortex 44, 312–324; Vallar, G., 2006. Memory systems: the case of phonological short-term memory. A festschrift for Cogni-
tive Neuropsychology. Cogn. Neuropsychol. 23, 135–155; Zimmer, H.D., 2008. Visual and spatial working memory: from boxes to networks. Neuro-
sci. Biobehav. Rev. 32, 1373–1395.

Interference by repetitive rTMS with neural activity in the left IPL (BA 40), and in BA 44 of the left hemisphere disrupts imme-
diate memory span (Romero et al., 2006); interference with neural activity in the left IPL (including BA 40) reduces the recency effect
in immediate free recall of auditory-verbal stimuli (Innocenti et al., 2013).
These findings qualify ArReh as a process which makes use of components (see Fig. 6) also concerned with the planning (ie,
programming in the left premotor cortex) of articulated speech. Finally, seen in this perspective, PhSTM may be regarded as
a component part of the language system. In line with the neuropsychological findings in brain-damaged patients that a dysfunction
of PhSTM disrupts the acquisition of new words (ie, nonwords to the patient, Baddeley et al., 1988), when healthy participants learn
nonwords, structures implicated in PhSTM (left BA 44, Broca’s area, the left TPJ) are activated (Paulesu et al., 2009).
When participants are engaged in tasks that require more “executive” processes, in addition to “storage” (eg, free recall, match-
ing a target one-, two-, or three-back in a sequence), the pattern of activation becomes more anterior, extending to the dorsolat-
eral prefrontal cortex (ie, BA 8, 45, 46), and bilateral (Fletcher and Henson, 2001; Smith and Jonides, 1999; Wager and Smith,
2003).
As often found in neuroimaging studies in healthy participants, the network activated by an immediate verbal retention task
tapping PhSTM is more extensive and bilateral, compared to the anatomo-clinical observations in brain-injured patients. In the
study by Paulesu et al. (1996), the activation extends to Wernicke’s area (BA 22) in the left hemisphere, and to a number of
regions in the right hemisphere homologous to the ones active in the left hemisphere (inferior frontal gyrus: Broca’s area, BA
44, and BA 6; supramarginal gyrus, BA 40; superior temporal gyrus, BA 22; insula). Similarly, a phonological judgment task acti-
vates in unimpaired participants also Wernicke’s area in the left hemisphere, and the insula in the right hemisphere. In a similar
vein, Romero et al. (2006) found that rTMS delivered not only to BA 44, but also to BA 40, impairs the ability of unimpaired
 Short-Term Memory 15

participants to perform phonological judgments on written words. Differences between evidence from lesion studies in brain-
damaged patients, on the one hand, and functional neuroimaging activation and interference by NIBS, on the other hand,
may be related to the redundancy of any neural system supporting a given function. Though the active network is bilateral,
its major components are however lateralized to the left hemisphere, where the regions mainly contributing to the execution
of the task of interest (in the present case, immediate memory span) are localized. Accordingly, only damage to these mainly
involved regions typically brings about clinically relevant behavioral consequences in brain-damaged patients. As for paradigms
using interference by NIBS in healthy participants, it should be noted that the behavioral effects are comparatively minor, as
compared with those of a cerebral lesion, not clinically apparent, and are shown only through sensitive paradigms, such as
response latencies, as in the study by Romero et al. (2006). In conclusion, while the network involved in a given function of
interest may be extensive, as revealed by functional neuroimaging and NIBS, brain damage elucidates the more relevant regions
and connections supporting the function of interest.
This approach implies that the complete understanding of the neural correlates of cognitive functions requires evidence from
lesion studies in brain-injured patients, neuroimaging activation, and NIBS experiments in unimpaired participants, reviving the
time-honored concept of “converging operations” (Garner et al., 1956), where “inferences (are) built upon many intersecting
and overlapping observations” (Hicks, 1971).

Visual and Spatial STM

STM for spatial locations, as assessed by tasks requiring the reproduction of sequences of positions in near extra-personal space,
is disrupted by damage to the posterior (parieto-occipital) regions of both the left and the right hemisphere, but the role of
right hemisphere damage appears to be more relevant (De Renzi et al., 1977; De Renzi and Nichelli, 1975; van Asselen
et al., 2006). These early suggestions have been confirmed and specified by whole brain voxel-based morphometry and
tract-wise lesion deficits analyses: bilateral occipital damage (middle occipital gyrus), and damage to the right IPL and the
TPJ, with disconnection of the right parieto-temporal segment of arcuate fasciculus, are associated with low spatial memory
span (Chechlacz et al., 2014). Studies in individual right-brain-damaged patients suggest that a network including the dorso-
medial posterior parietal cortex (BA 7), and the dorsal premotor cortex (BA 6) is involved in STM for spatial locations
(assessed, for instance, by Corsi’s test, see Fig. 11A), with verbal and visual STM being spared (Carlesimo et al., 2001). Damage
to the premotor, and prefrontal dorsomedial regions of the right hemisphere may specifically disrupt the rehearsal of spatial
information, causing disproportionate short-term forgetting in the range of seconds (Bonnì et al., 2014). Damage to the right
hemisphere may also affect STM for unfamiliar faces (Hanley et al., 1990). Conversely, PhSTM is not impaired by lesions in the
right hemisphere.
Some deficits of visual STM are associated with damage to the posterior (parieto-occipital) regions of the left hemisphere. One
such impairment concerns the immediate retention of sequences of visual stimuli, such as straight or curved lines (Warrington and
Rabin, 1971). This deficit may occur in the presence of a normal auditory-verbal span.
In sum, lesion studies in brain-damaged patients provide support to the distinction between STM for spatial locations and for
visual patterns, the former being more closely associated with right hemisphere damage, the latter with left hemisphere damage. The
hemispheric lateralization of neural networks concerned with visuo-spatial STM, particularly for the visual component, is less
pronounced than that of the networks supporting PhSTM.
The anatomical differences between primarily spatial (location, “where”) and visual (recognition “what”) STM components
concern both the relevant cerebral areas and their prevailing lateralization. When healthy participants are engaged in tasks
involving STM for spatial location the activated areas include the occipital extra-striate (BA 19), the posterior parietal, dorso-
lateral premotor (BA 6) and prefrontal cortices. Short-term recognition memory for visual patterns (eg, faces, designs) is asso-
ciated with activation in a more ventral network including the occipital and temporal (BA 37) regions, the posterior-inferior
parietal region (Todd and Marois, 2004), and the prefrontal cortex. Within the prefrontal cortex (BA 45, 46, 47, 8, 9), more
dorsolateral areas have been associated with STM for spatial location, more ventral areas with STM for patterns. Activation
has been frequently found to be bilateral, but there is evidence suggesting more sustained activity in the right hemisphere
when the task assesses recognition memory for spatial location, in the left hemisphere when the task probes recognition
memory for visual patterns.
These non-overlapping patterns of activation indicate an association between the “dorsal visual stream” and STM for spatial loca-
tions, and between the “ventral visual stream” and recognition STM for visual patterns. These findings also suggest the existence of
some hemispheric asymmetry, with right-sided areas being more concerned with the spatial (location) aspects of short-term reten-
tion, left-sided areas with the visual (pattern recognition) aspects. The relative contribution of different cerebral regions of the two
sides of the brain may also be related to the format of the cerebral representation involved. For instance, a left lateralization for
visual object STM may reflect a more symbolic or verbal encoding, a right-sided lateralization a more image-based and spatial
encoding.
Finally, the different cerebral areas participating in the network may provide distinct contributions to the short-term retention
process. The occipital and temporal regions may mainly participate in perceptual analysis, the PPC in computing the coordinates of
the stimulus and in providing the capacity for STS, the premotor and prefrontal cortices in the rehearsal and retention processes
(Courtney, 2004; Zimmer, 2008).
16 Short-Term Memory

Short-Term Memory and Cognition

In the preceding sections behavioral and anatomical evidence for the existence of a number of discrete systems concerned with
short-term retention has been considered. In this section, STM systems are seen from the perspective of their relationships with other
components of the cognitive system, and their role in cognitive activity. A general starting question may be the following. Is there
a use for a system providing the temporary retention of a limited amount of stimuli, in addition to infrequent situations, namely:
a friend says us once an unfamiliar eight-digit number, which we have to dial on a mobile telephone we left in a building placed on
the other side of a large street, and we have no paper and pencil to write it down?
Serial models of memory (see Fig. 1) imply that STM is a necessary stage for the stable acquisition of new information in LTM.
More specifically, there is definite evidence that PhSTM plays an important role in learning new vocabulary. In addition, this system
participates in the processes of speech comprehension and production. The evidence concerning visual and spatial STM is less
definite.

Long-Term Learning
The observation that patients with defective auditory-verbal span are also impaired in learning unfamiliar pronounceable letter
sequences raises the possibility that PhSTM may contribute to a relevant aspect of language development, the acquisition of
vocabulary (Fig. 10). Similarly, participants with a developmental deficit of phonological memory are impaired in vocabulary
acquisition and in nonword learning. An opposite pattern is provided by individual participants with a congenital cognitive
impairment, which selectively spares PhSTM: Acquisition of vocabulary, foreign languages and nonword learning are also
preserved (Vallar and Papagno, 1993). Converging evidence from different participants’ populations supports this view. First,
correlational studies in children show that the capacity of PhSTM is a main predictor of the subsequent acquisition of vocab-
ulary, both in the native and in a second language (Baddeley et al., 1998). Second, in unimpaired adult participants, the vari-
ables which disrupt immediate memory span (phonological similarity, item length, articulatory suppression) also impair the
acquisition of nonwords (Papagno and Vallar, 1992). Third, polyglot participants have a higher auditory-verbal span,
compared to non-polyglots, and a better ability to learn novel words (Fig. 13) This advantage of polyglots in nonword learning
is not related to differences in general intelligence, verbal or spatial LTM, * and is likely to reflect a greater capacity and superior
function of PhSTM (Papagno and Vallar, 1995).

(A) (B)
SPAN
9 AUDITORY DIGIT SPAN CORRECT RECALL

SPATIAL POSITION SPAN

8 8

7 7

6 6

5 5

4 4

3 P-WORDS
3
P-NONWORDS
2 2
NP-WORDS
1 1 NP-NONWORDS

0 0
P NP 1 2 3 4 5 6 7 8 9 10
TRIALS

Figure 13 Phonological STM and learning of nonwords by polyglot and non-polyglot participants. (A) Polyglot (P) participants have a higher
average immediate auditory-verbal digit span (*), while their average spatial span is comparable to that of non-polyglot (NP) participants. (B) In
a word-nonword paired-associate learning task, P participants show a better learning of pronounceable strings of letters, not corresponding to any
lexical item known to them. The P participants’ superior learning of new phonological information does not reflect higher learning skills in general, as
shown by their word learning performance, which is comparable to that of NP participants. Similarly, the comparable performance of the two groups
in spatial span indicates that the short-term memory advantage of P participants is confined to the phonological system. Source: Redrawn from
Papagno, C., Vallar, G., 1995. Verbal short-term memory and vocabulary learning in polyglots. Q. J. Exp. Psychol. 48A, 98–107.
 Short-Term Memory 17

In line with these observations, anodal excitatory tDCS to the left temporo-parietal cortex enhances immediate serial recall of
nonwords (Savill et al., 2015), and the acquisition of new vocabulary (Flöel et al., 2008; Meinzer et al., 2014). This system may
be considered as a learning device for the acquisition of novel phonological representations and the building up of the phonological
lexicon. Some observations in brain-damaged patients suggest a similar role for visuo-spatial STM in the acquisition of new visual
information, such as unfamiliar faces and objects (Hanley et al., 1990, 1991).

Language Processing
Sentence Comprehension
The general idea that temporary retention in phonological memory may contribute to aspects of speech comprehension is far
from novel, dating back at least to the late 1970s (Clark and Clark, 1977; Just and Carpenter, 1992). WM resources are involved
in the comprehension of sentences (Martin, 2006), particularly for aspects of syntactic processing (Caplan and Waters, 1999).
Specifically, PhSTM may hold incoming auditory-verbal strings, while syntactic and lexical-semantic analyses are performed.
Patients with defective PhSTM show preserved comprehension of individual words, and of many sentential materials, and
a normal ability to decide whether or not sentences are grammatically correct. This may reflect on the one hand the operation
of on-line syntactic and lexical-semantic processes, heuristics and pragmatic, on the other hand the complete independence of
syntactic and lexical-semantic processes from PhSTM. Patients with a deficit of PhSTM are impaired, however, with “complex”
sentences. “Complexity” may refer to a number of non-mutually exclusive factors, with word order being a relevant and widely
investigated parameter. There patients exhibit a defective performance in the Token Test (De Renzi and Faglioni, 1978), partic-
ularly with long commands, in which word order is crucial (eg, “Touch the small green square and the large black circle”), and,
similarly, with sentences in which word order conveys information crucial for meaning, such as when a semantic anomaly is
introduced by a change in their linear arrangement (eg, “The world divides the equator into two hemispheres, the southern and
the northern,” Vallar and Baddeley, 1984b). Syntactically complex sentences, such as center-embedded subject and object
relative sentences (“The dog which chases the cat looks at grandpa,” and “The cat which the child caresses drinks milk”) are
difficult to be understood for these patients (Papagno et al., 2007). With complex sentential material of this sort, an adequate
interpretation may require backtracking to the verbatim (phonological) representation of the sentence, temporarily held in
PhSTM, through the process of ArReh. This may provide a “backup” or “mnemonic window” resource, for performing the
supplementary operations necessary for comprehension.

Speech Production
In spontaneous speech some errors involve the sequential misplacement of consonant or vowels (phonemic Spoonerism, eg,
“You have missed all my history lectures” / “You have hissed all my mistery lectures.” This type of errors shows that parts of
speech several words in length are planned in advance, and stored in a phonological format, before actual articulation. The
short-term retention component “Phonological Assembly System” or “Phonological Output Buffer” (Caramazza et al.,
1986; Shallice et al., 2000) where such errors are generated may be conceived as a working memory space, in which phono-
logical segments are temporarily stored prior to the application of various output processes, such as planning and editing of
the articulatory procedures needed to produce speech. This output retention system, in addition, participates in the process of
rehearsing material held in the PhSTS (see Fig. 6), and may be hence involved in the comprehension of complex sentences,
that need to be temporarily held in a phonological format, as discussed in the preceding section. Seen in this perspective,
ArReh is an activity, which makes use of components primarily concerned with a basic linguistic function: speech production.
Finally, the Phonological Output Buffer provides a WM space where operations such as segmentation and deletion may be
performed. These may support phonological skills involving a comparison between segments of pairs of stimuli, for example:
phonological judgments about stress assignment or similarity of the initial sound of words (Papagno et al., 2007; Vallar et al.,
1997).

Spatial Orientation and Navigation

Spatial orientation and navigation in three-dimensional space are complex skills, which require, among other things, keeping track
of the current position of the body, with reference to previous locations in the environment. A spatial STM system is likely to be
involved in the retention of the successive body displacements, occurring during navigation. Such a system is likely to integrate
multiple sensory inputs (visual, proprioceptive-somatosensory, vestibular, auditory), and to interact with relevant spatial informa-
tion, stored in LTM. These spatial memory systems are likely to make use of multiple reference frames along the egocentric-
allocentric dimension (Wolbers and Hegarty, 2010), and their impairment may contribute to topographical disorientation (Aguirre
and D’Esposito, 1999; Brunsdon et al., 2007).

Conclusions

Behavioral observations and neuroanatomical evidence from healthy participants and brain-injured patients concur to suggest that
STM should be conceived as a multiple-component system with specific functional properties and discrete neural correlates. Current
18 Short-Term Memory

evidence suggests the existence of phonological, spatial and visual STM systems, each including store and rehearsal components.
These systems secure retention of a limited amount of material in the time range of seconds, and contribute to relevant aspects
of cognition, such as long-term learning of new, unfamiliar material. Specifically, PhSTM is related to a number of aspects of
language: vocabulary acquisition, speech production, phonological judgments, and comprehension of complex sentences. Spatial
and visual STM may be involved in spatial orientation and navigation.

Acknowledgments

Supported in part by ATE grants from the University of Milan-Bicocca, and Ricerca Corrente from the Ministry of Health to the IRCCS Istituto
Auxologico Italiano.

References

Acheson, D.J., Postle, B.R., MacDonald, M.C., 2010. The interaction of concreteness and phonological similarity in verbal working memory. J. Exp. Psychol. Learn. Mem. Cogn. 36,
17–36.
Aguirre, G.K., D’Esposito, M., 1999. Topographical disorientation: a synthesis and taxonomy. Brain 122, 1613–1628.
Ardila, A., 2010. A review of conduction aphasia. Curr. Neurol. Neurosci. Rep. 10, 499–503.
van Asselen, M., Kessels, R.P.C., Neggers, S.F.W., Kappelle, L.J., Frijns, C.J.M., Postma, A., 2006. Brain areas involved in spatial working memory. Neuropsychologia 44,
1185–1194.
Atkinson, R.C., Shiffrin, R.M., 1971. The control of short-term memory. Sci. Am. 225, 82–90.
Baddeley, A.D., 1966. Short-term memory for word sequences as a function of acoustic, semantic and formal similarity. Q. J. Exp. Psychol. 18, 362–365.
Baddeley, A.D., 1976. The Psychology of Memory. Basic Books Inc., New York.
Baddeley, A.D., 2003. Working memory: looking back and looking forward. Nat. Rev. Neurosci. 4, 829–839.
Baddeley, A.D., Gathercole, S., Papagno, C., 1998. The phonological loop as a language learning device. Psychol. Rev. 105, 158–173.
Baddeley, A.D., Papagno, C., Vallar, G., 1988. When long-term learning depends on short-term storage. J. Mem. Lang. 27, 586–595.
Baddeley, A.D., Thomson, N., Buchanan, M., 1975. Word length and the structure of short-term memory. J. Verbal Learn. Verbal Behav. 14, 575–589.
Baddeley, A.D., Warrington, E.K., 1970. Amnesia and the distinction between long- and short-term memory. J. Verbal Learn. Verbal Behav. 9, 176–189.
Basso, A., Spinnler, H., Vallar, G., Zanobio, M.E., 1982. Left hemisphere damage and selective impairment of auditory verbal short-term memory. A case study. Neuropsychologia
20, 263–274.
Bishop, D.V.M., Robson, J., 1989. Unimpaired short-term memory and rhyme judgement in congenitally speechless individuals: implications for the notion of “articulatory coding”.
Q. J. Exp. Psychol. 41A, 123–140.
Bonnì, S., Perri, R., Fadda, L., Tomaiuolo, F., Koch, G., Caltagirone, C., Carlesimo, G.A., 2014. Selective deficit of spatial short-term memory: role of storage and rehearsal
mechanisms. Cortex 59, 22–32.
Brener, R., 1940. An experimental investigation of memory span. J. Exp. Psychol. 26, 467–482.
Brunsdon, R., Nickels, L., Coltheart, M., 2007. Topographical disorientation: towards an integrated framework for assessment. Neuropsychol. Rehabil. 17, 34–52.
Buchsbaum, B.R., Baldo, J., Okada, K., Berman, K.F., Dronkers, N., D’Esposito, M., Hickok, G., 2011. Conduction aphasia, sensory-motor integration, and phonological short-term
memory - an aggregate analysis of lesion and fMRI data. Brain Lang. 119, 119–128.
Buchsbaum, B.R., D’Esposito, M., 2008. The search for the phonological store: from loop to convolution. J. Cogn. Neurosci. 20, 762–778.
Burani, C., Vallar, G., Bottini, G., 1991. Articulatory coding and phonological judgements on written words and pictures: the role of the phonological output buffer. Eur. J. Cogn.
Psychol. 3, 379–398.
Campoy, G., Castellà, J., Provencio, V., Hitch, G.J., Baddeley, A.D., 2015. Automatic semantic encoding in verbal short-term memory: evidence from the concreteness effect. Q. J.
Exp. Psychol. 68, 759–778.
Caplan, D., Waters, G.S., 1999. Verbal working memory and sentence comprehension. Behav. Brain Sci. 22, 77–94 discussion 95–126.
Caramazza, A., Miceli, G., Villa, G., 1986. The role of the (output) phonological buffer in reading, writing, and repetition. Cogn. Neuropsychol. 3, 37–76.
Carlesimo, G.A., Perri, R., Turriziani, P., Tomaiuolo, F., Caltagirone, C., 2001. Remembering what but not where: independence of spatial and visual working memory in the human
brain. Cortex 37, 457–473.
Chechlacz, M., Rotshtein, P., Humphreys, G.W., 2014. Neuronal substrates of Corsi Block span: lesion symptom mapping analyses in relation to attentional competition and spatial
bias. Neuropsychologia 64, 240–251.
Clark, H.H., Clark, E., 1977. Psychology and Language. An Introduction to Psycholinguistics. Harcourt Brace Jovanovich, New York.
Corkin, S., 2002. What’s new with the amnesic patient H.M.? Nat. Rev. Neurosci. 3, 153–160.
Courtney, S.M., 2004. Attention and cognitive control as emergent properties of information representation in working memory. Cogn. Affect. Behav. Neurosci. 4, 501–516.
Craik, F.I.M., Watkins, M.J., 1973. The role of rehearsal in short-term memory. J. Verbal Learn. Verbal Behav. 12, 599–607.
Cubelli, R., Nichelli, P., 1992. Inner speech in anarthria: neuropsychological evidence of differential effects of cerebral lesions on subvocal articulation. J. Clin. Exp. Neuropsychol.
14, 499–517.
Dale, H.C., 1973. Short-term memory for visual information. Br. J. Psychol. 64, 1–8.
De Renzi, E., Faglioni, P., 1978. Normative data and screening power of a shortened version of the Token Test. Cortex 14, 41–49.
De Renzi, E., Faglioni, P., Previdi, P., 1977. Spatial memory and hemispheric locus of lesion. Cortex 13, 424–433.
De Renzi, E., Nichelli, P., 1975. Verbal and non-verbal short-term memory impairment following hemispheric damage. Cortex 11, 341–354.
Della Sala, S., Logie, R.H., 2002. Neuropsychological impairments of visual and spatial working memory. In: Baddeley, A., Wilson, B., Kopelman, M. (Eds.), Handbook of Memory
Disorders. Wiley, Chichester, England, pp. 271–292.
Drachman, D.A., Arbit, J., 1966. Memory and the hippocampal complex. Arch. Neurol. 15, 52–61.
Farah, M.J., Hammond, K.M., Levine, D.N., Calvanio, R., 1988. Visual and spatial mental imagery: dissociable systems of representation. Cogn. Psychol. 20, 439–462.
Fletcher, P.C., Henson, R.N., 2001. Frontal lobes and human memory: insights from functional neuroimaging. Brain 124, 849–881.
Flöel, A., Rösser, N., Michka, O., Knecht, S., Breitenstein, C., 2008. Noninvasive brain stimulation improves language learning. J. Cogn. Neurosci. 20, 1415–1422.
Garner, W.R., Hake, H.W., Eriksen, C.W., 1956. Operationism and the concept of perception. Psychol. Rev. 63, 149–159.
Glanzer, M., 1972. Storage mechanisms in recall. In: Bower, G.H. (Ed.), The Psychology of Learning and Motivation, Advances in Research and Theory, vol. 5. Academic Press,
New York, pp. 129–193.
Grossi, D., Becker, J.T., Smith, C., Trojano, L., 1993. Memory for visuospatial patterns in Alzheimer’s disease. Psychol. Med. 23, 65–70.
Hanley, J.R., Pearson, N.A., Young, A.W., 1990. Impaired memory for new visual forms. Brain 113, 1131–1148.
 Short-Term Memory 19

Hanley, J.R., Young, A.W., Pearson, N.A., 1991. Impairment of the visuo-spatial sketch pad. Q. J. Exp. Psychol. A Hum. Exp. Psychol. 43, 101–125.
Heyden, A., Sparr, G., Nielsen, M., Johansen, P. (Eds.), 2002. Computer Vision – ECCV 2002. In 7th European Conference on Computer Vision Copenhagen, Denmark, May 28–31,
2002 Proceedings, Part IV. Springer-Verlag, Berlin.
Hicks, R.G., 1971. Converging operations in the psychological experiment. Psychophysiology 8, 93–101.
Hulme, C., Newton, P., Cowan, N., Stuart, G., Brown, G., 1999. Think before you speak: pauses, memory search, and trace redintegration processes in verbal memory span. J. Exp.
Psychol. Learn. Mem. Cogn. 25, 447–463.
Hurlstone, M.J., Hitch, G.J., Baddeley, A.D., 2014. Memory for serial order across domains: an overview of the literature and directions for future research. Psychol. Bull. 140,
339–373.
Innocenti, I., Cappa, S.F., Feurra, M., Giovannelli, F., Santarnecchi, E., Bianco, G., Cincotta, M., Rossi, S., 2013. TMS interference with primacy and recency mechanisms reveals
bimodal episodic encoding in the human brain. J. Cogn. Neurosci. 25, 109–116.
James, W., 1890. The Principles of Psychology. Henry Holt, New York.
Jansen, A., Flöel, A., Van Randenborgh, J., Konrad, C., Rotte, M., Förster, A.-F., Deppe, M., Knecht, S., 2005. Crossed cerebro-cerebellar language dominance. Hum. Brain Mapp.
24, 165–172.
Jefferies, E., Bott, S., Ehsan, S., Lambon Ralph, M.A., 2011. Phonological learning in semantic dementia. Neuropsychologia 49, 1208–1218.
Jefferies, E., Jones, R.W., Bateman, D., Lambon Ralph, M.A., 2005. A semantic contribution to nonword recall? Evidence for intact phonological processes in semantic dementia.
Cogn. Neuropsychol. 22, 183–212.
Jonides, J., Lewis, R.L., Nee, D.E., Lustig, C.A., Berman, M.G., Moore, K.S., 2008. The mind and brain of short-term memory. Annu. Rev. Psychol. 59, 193–224.
Just, M.A., Carpenter, P.A., 1992. A capacity theory of comprehension: individual differences in working memory. Psychol. Rev. 99, 122–149.
Kelly, C., Uddin, L.Q., Shehzad, Z., Margulies, D.S., Castellanos, F.X., Milham, M.P., Petrides, M., 2010. Broca’s region: linking human brain functional connectivity data and non-
human primate tracing anatomy studies. Europ. J. Neurosci. 32, 383–398.
Lambon Ralph, M.A., 2013. Neurocognitive insights on conceptual knowledge and its breakdown. Philos. Trans. R. Soc. Lond. B: Biol. Sci. 369, 20120392.
Locke, J., 1700. An Essay Concerning Human Understanding. Clarendon Press, Oxford.
Logie, R.H., Della Sala, S., 2005. Disorders of visuospatial working memory. In: Logie, R.H., Della Sala, S. (Eds.), The Cambridge Handbook of Visuospatial Thinking. Cambridge
University Press, Cambridge, UK, pp. 81–120.
Malhotra, P., Mannan, S., Driver, J., Husain, M., 2004. Impaired spatial working memory: one component of the visual neglect syndrome? Cortex 40, 667–676.
Martin, R.C., 2006. The neuropsychology of sentence processing: where do we stand? Cogn. Neuropsychol. 23, 74–95.
Meinzer, M., Jähnigen, S., Copland, D.A., Darkow, R., Grittner, U., Avirame, K., Rodriguez, A.D., Lindenberg, R., Flöel, A., 2014. Transcranial direct current stimulation over multiple
days improves learning and maintenance of a novel vocabulary. Cortex 50, 137–147.
Melton, A., 1963. Implications of short-term memory for a general theory of memory. J. Verbal Learn. Verbal Behav. 2, 1–21.
Miller, G.A., 1956. The magical number seven plus or minus two: some limits on our capacity for processing information. Psychol. Rev. 63, 81–97.
Milner, B., Corkin, S., Teuber, H.-L., 1968. Further analysis of the hippocampal amnesic syndrome: 14-year follow-up study of H.M. Neuropsychologia 6, 215–234.
Moreton, B.J., Ward, G., 2010. Time scale similarity and long-term memory for autobiographical events. Psychon. Bull. Rev. 17, 510–515.
Papagno, C., Cecchetto, C., Reati, F., Bello, L., 2007. Processing of syntactically complex sentences relies on verbal short-term memory: evidence from a short-term memory
patient. Cogn. Neuropsychol. 24, 292–311.
Papagno, C., Lucchelli, F., Vallar, G., 2008. Phonological recoding, visual short-term store and the effect of unattended speech: evidence from a case of slowly progressive anarthria.
Cortex 44, 312–324.
Papagno, C., Vallar, G., 1992. Phonological short-term memory and the learning of novel words: the effect of phonological similarity and item length. Q. J. Exp. Psychol. 44A,
47–67.
Papagno, C., Vallar, G., 1995. Verbal short-term memory and vocabulary learning in polyglots. Q. J. Exp. Psychol. 48A, 98–107.
Papagno, C., Vernice, M., Cecchetto, C., 2013. Phonology without semantics? Good enough for verbal short-term memory. Evidence from a patient with semantic dementia. Cortex
49, 626–636.
Paulesu, E., Frith, U., Snowling, M., Gallagher, A., Morton, J., Frackowiak, R.S.J., Frith, C.D., 1996. Is developmental dyslexia a disconnection syndrome? Evidence from PET
scanning. Brain 119, 143–157.
Paulesu, E., Vallar, G., Berlingeri, M., Signorini, M., Vitali, P., Burani, C., Fazio, F., 2009. Supercalifragilisticexpialidocious: how the brain learns words never heard before.
NeuroImage 45, 1368–1377.
Penfield, W., Milner, B., 1958. Memory deficit produced by bilateral lesions in the hippocampal zone. Arch. Neurol. Psychiatry 79, 475–497.
Petrides, M., Tomaiuolo, F., Yeterian, E.H., Pandya, D.N., 2012. The prefrontal cortex: Comparative architectonic organization in the human and the macaque monkey brains. Cortex
48, 46–57.
Phillips, W.A., 1974. On the distinction between sensory storage and short-term memory. Percept. Psychophys. 16, 283–290.
Piccardi, L., Bianchini, F., Zompanti, L., Guariglia, C., 2008. Pure representational neglect and navigational deficits in a case with preserved visuo-spatial working memory.
Neurocase 14, 329–342.
Ravizza, S.M., McCormick, C.A., Schlerf, J.E., Justus, T., Ivry, R.B., Fiez, J.A., 2006. Cerebellar damage produces selective deficits in verbal working memory. Brain 129, 306–320.
Repovs, G., Baddeley, A., 2006. The multi-component model of working memory: explorations in experimental cognitive psychology. Neuroscience 139, 5–21.
Romero, L., Walsh, V., Papagno, C., 2006. The neural correlates of phonological short-term memory: a repetitive transcranial magnetic stimulation study. J. Cogn. Neurosci. 18,
1147–1155.
Roodenrys, S., Hulme, C., Lethbridge, A., Hinton, M., Nimmo, L.M., 2002. Word-frequency and phonological-neighborhood effects on verbal short-term memory. J. Exp. Psychol.
Learn. Mem. Cogn. 28, 1019–1034.
Ruchkin, D.S., Grafman, J., Cameron, K., Berndt, R.S., 2003. Working memory retention systems: a state of activated long-term memory. Behav. Brain Sci. 26, 709–769.
Salamé, P., Baddeley, A.D., 1982. Disruption of short-term memory by unattended speech: implications for the structure of working memory. J. Verbal Learn. Verbal Behav. 21,
150–164.
Savill, N., Ashton, J., Gugliuzza, J., Poole, C., Sim, Z., Ellis, A.W., Jefferies, E., 2015. tDCS to temporoparietal cortex during familiarisation enhances the subsequent phonological
coherence of nonwords in immediate serial recall. Cortex 63, 132–144.
Schnakers, C., Majerus, S., Goldman, S., Boly, M., Van Eeckhout, P., Gay, S., Pellas, F., Bartsch, V., Peigneux, P., Moonen, G., Laureys, S., 2008. Cognitive function in the locked-
in syndrome. J. Neurol. 255, 323–330.
Scoville, W.B., Milner, B., 1957. Loss of recent memory after bilateral hippocampal lesions. J. Neurol. Neurosurg. Psychiatry 20, 11–21.
Shallice, T., Butterworth, B., 1977. Short-term memory impairment and spontaneous speech. Neuropsychologia 15, 729–735.
Shallice, T., Rumiati, R.I., Zadini, A., 2000. The selective impairment of the phonological output buffer. Cogn. Neuropsychol. 17, 517–546.
Shallice, T., Vallar, G., 1990. The impairment of auditory-verbal short-term storage. In: Vallar, G., Shallice, T. (Eds.), Neuropsychological Impairments of Short-Term Memory.
Cambridge University Press, Cambridge, pp. 11–53.
Sidman, M., Stoddard, L.T., Mohr, J.P., 1968. Some additional quantitative observations of immediate memory in a patient with bilateral hippocampal lesions. Neuropsychologia 6,
245–254.
Silveri, M.C., Di Betta, A.M., Filippini, V., Leggio, M.G., Molinari, M., 1998. Verbal short-term store-rehearsal system and the cerebellum. Evidence from a patient with a right
cerebellar lesion. Brain 121, 2175–2187.
20 Short-Term Memory

Smith, E.E., Jonides, J., 1999. Storage and executive processes in the frontal lobes. Science 283, 1657–1661.
Striemer, C.L., Ferber, S., Danckert, J., 2013. Spatial working memory deficits represent a core challenge for rehabilitating neglect. Front. Hum. Neurosci. 7, 334.
Thorn, A., Page, M. (Eds.), 2009. Interactions Between Short-Term and Long-Term Memory in the Verbal Domain. Psychology Press, Hove.
Todd, J.J., Marois, R., 2004. Capacity limit of visual short-term memory in human posterior parietal cortex. Nature 428, 751–754.
Tomlinson, S.P., Davis, N.J., Morgan, H.M., Bracewell, R.M., 2014. Cerebellar contributions to verbal working memory. Cerebellum (London, England) 13, 354–361.
Tulving, E., 1972. Episodic and semantic memory. In: Tulving, E., Donaldson, W. (Eds.), Organization of Memory. Academic Press, New York, pp. 381–403.
Unsworth, N., Engle, R.W., 2007. On the division of short-term and working memory: an examination of simple and complex span and their relation to higher order abilities.
Psychol. Bull. 133, 1038–1066.
Vallar, G., 2006. Memory systems: the case of phonological short-term memory. A festschrift for Cognitive Neuropsychology. Cogn. Neuropsychol. 23, 135–155.
Vallar, G., Baddeley, A.D., 1982. Short-term forgetting and the articulatory loop. Q. J. Exp. Psychol. 34, 53–60.
Vallar, G., Baddeley, A.D., 1984a. Fractionation of working memory: neuropsychological evidence for a phonological short-term store. J. Verbal Learn. Verbal Behav. 23, 151–161.
Vallar, G., Baddeley, A.D., 1984b. Phonological short-term store, phonological processing and sentence comprehension. Cogn. Neuropsychol. 1, 121–141.
Vallar, G., Bolognini, N., 2014. Unilateral spatial neglect. In: Nobre, A.C., Kastner, S. (Eds.), Oxford Handbook of Attention. Oxford University Press, Oxford, pp. 972–1027.
Vallar, G., Cappa, S.F., 1987. Articulation and verbal short-term memory. Evidence from anarthria. Cogn. Neuropsychol. 4, 55–78.
Vallar, G., Di Betta, A.M., Silveri, M.C., 1997. The phonological short-term store-rehearsal system: patterns of impairment and neural correlates. Neuropsychologia 35, 795–812.
Vallar, G., Papagno, C., 1986. Phonological short-term store and the nature of the recency effect: evidence from neuropsychology. Brain Cogn. 5, 428–442.
Vallar, G., Papagno, C., 1993. Preserved vocabulary acquisition in Down’s syndrome: the role of phonological short-term memory. Cortex 29, 467–483.
Vallar, G., Papagno, C., 2002. Neuropsychological impairments of verbal short-term memory. In: Baddeley, A., Wilson, B., Kopelman, M. (Eds.), Handbook of Memory Disorders.
Wiley, Chichester, England, pp. 249–270.
Van Vleet, T.M., DeGutis, J.M., 2013. The nonspatial side of spatial neglect and related approaches to treatment. Prog. Brain Res. 207, 327–349.
Wager, T.D., Smith, E.E., 2003. Neuroimaging studies of working memory: a meta-analysis. Cogn. Affect. Behav. Neurosci. 3, 255–274.
Wansard, M., Bartolomeo, P., Bastin, C., Segovia, F., Gillet, S., Duret, C., Meulemans, T., 2015. Support for distinct subcomponents of spatial working memory: a double
dissociation between spatial-simultaneous and spatial-sequential performance in unilateral neglect. Cogn. Neuropsychol. 32, 14–28.
Wansard, M., Meulemans, T., Gillet, S., Segovia, F., Bastin, C., Toba, M., Bartolomeo, P., 2014. Visual neglect: is there a relationship between impaired spatial working memory
and re-cancellation? Exp. Brain Res. 232, 3333–3343.
Warrington, E.K., Rabin, P., 1971. Visual span of apprehension in patients with unilateral cerebral lesions. Q. J. Exp. Psychol. 23, 423–431.
Warrington, E.K., Shallice, T., 1969. The selective impairment of auditory-verbal short-term memory. Brain 92, 885–896.
Watkins, M.J., 1977. The intricacy of memory span. Mem. Cogn. 5, 529–534.
Wickelgren, W.A., 1968. Sparing of short-term memory in an amnesic patient: implications for strength theory of memory. Neuropsychologia 6, 235–244.
Wilson, B.A., Baddeley, A.D., Young, A.W., 1999. LE, a person who lost her “mind’s eye”. Neurocase 5, 119–127.
Wolbers, T., Hegarty, M., 2010. What determines our navigational abilities? Trends Cogn. Sci. 14, 138–146.
Zannino, G.D., Caltagirone, C., Carlesimo, G.A., 2015. The contribution of neurodegenerative diseases to the modelling of semantic memory: a new proposal and a review of the
literature. Neuropsychologia 75, 274–290.
Zimmer, H.D., 2008. Visual and spatial working memory: from boxes to networks. Neurosci. Biobehav. Rev. 32, 1373–1395.