Brain Imaging and Behavior (2020) 14:2817–2830

https://doi.org/10.1007/s11682-019-00187-4

REVIEW ARTICLE

The superior longitudinal fascicle: reconsidering the fronto-parietal

neural network based on anatomy and function
Riho Nakajima 1 & Masashi Kinoshita 2 & Harumichi Shinohara 3 & Mitsutoshi Nakada 2

Published online: 29 August 2019

# Springer Science+Business Media, LLC, part of Springer Nature 2019

Abstract
Due primarily to the extensive disposition of fibers and secondarily to the methodological preferences of researchers, the superior
longitudinal fasciculus (SLF) subdivisions have multiple names, complicating SLF research. Here, we collected and reassessed
existing knowledge regarding the SLF, which we used to propose a four-term classification of the SLF based mainly on function:
dorsal SLF, ventral SLF, posterior SLF, and arcuate fasciculus (AF); these correspond to the traditional SLF II, SLF III or anterior
AF, temporoparietal segment of the SLF or posterior AF, and AF or AF long segment, respectively. Each segment has a distinct
functional role. The dorsal SLF is involved in visuospatial attention and motor control, while the ventral SLF is associated with
language-related networks, auditory comprehension, and articulatory processing in the left hemisphere. The posterior SLF is
involved in language-related processing, including auditory comprehension, reading, and lexical access, while the AF is asso-
ciated with language-related activities, such as phonological processing; the right AF plays a role in social cognition and
visuospatial attention. This simple proposed classification permits a better understanding of the SLF and may comprise a
convenient classification for use in research and clinical practice relating to brain function.

Keywords Superior longitudinal fascicles . Arcuate fascicles . Anatomy . Imaging analysis . Neuropsychological function

Introduction
The superior longitudinal fasciculus (SLF) is an extensive
white-matter tract that mainly communicates between frontal
and parietal lobes, and provides partial communication with
the temporal lobe. It interconnects nearly all cortical areas of
the lateral cerebral hemisphere. The frontoparietal network is
Electronic supplementary material The online version of this article
(https://doi.org/10.1007/s11682-019-00187-4) contains supplementary
material, which is available to authorized users.
* Mitsutoshi Nakada
mnakada@med.kanazawa-u.ac.jp
1 and long. Additionally, in 2005–2006, Schmahmann and
Department of Occupational therapy, Faculty of Health Sciences,
Institute of Medical, Pharmaceutical and Health Sciences, Kanazawa
University, Kanazawa, Japan
2
Department of Neurosurgery, Faculty of Medicine, Institute of
Medical, Pharmaceutical and Health Sciences, Kanazawa University,
13-1 Takara-machi, Kanazawa, Ishikawa 920-8641, Japan
3
Department of Functional Anatomy, Kanazawa University,
Kanazawa, Japan
involved in various functions that are essential for daily living,
such as speaking, tool use, perspective taking, and empathy.
The existence of the SLF has been known for more than
100 years, and the arcuate fasciculus (AF) has been regarded
as part of the SLF. Since the 1980s and the methodological
development of neuroimaging analysis, a number of attempts
have been made to classify the SLF (details are shown in A1).
In Petrides and Pandya 1984, Petrides and Pandya (1984)
demonstrated that the frontoparietal network could be divided
into three segments, defined as SLF I, II, and III. Consistent
with previous systems, they considered the AF to be part of
the SLF and defined it as the fourth segment of the SLF. In
2005, using diffusion tensor imaging (DTI), Catani et al.
(2005) designated the classical AF as the SLF/AF (AF, name-
ly SLF) and divided it into three segments: anterior, posterior,
Pandya (2006) and Makris et al. (2005) investigated the struc-
tural connectivity of the SLF in human and monkey brains,
using the SLF I–III classification. Subsequently, a growing
number of studies on structural brain networks have been
conducted using neuroimaging techniques and fiber dissec-
tion. However, the classification of the SLF has become con-
fusing, due to the presence of multiple classification systems,
2818
each of which could be employed based upon the preferences
of the researchers performing a given study.
Over the past decade, multiple studies, using magnetic res-
onance images (MRI), functional MRI (fMRI), resting-state
fMRI, or direct electrical stimulation during awake surgery,
have indicated the involvement of the human SLF in several
different functions. For instance, the SLF is involved in lan-
guage, visuospatial cognition, attention, working memory,
and mentalizing; different segments of the SLF relate to these
functions depending on the cortices to which they communi-
cate (Duffau et al. 2002; Catani et al. 2005; Makris et al. 2005;
Curtis 2006; Thiebaut de Schotten et al. 2011a; Herbet et al.
2014). Regarding pathological conditions, the structural con-
nectivity of the SLF in patients with obsessive-compulsive
disorder, mild cognitive impairment, and developmental dis-
orders, including attention deficit hyperactivity disorder and
autism, differs from that of normal controls (Blanken et al.
2017; Gan et al. 2017; Gehricke et al. 2017; Liu et al. 2017).
In this review, we strive toward two goals. First, we revisit
existing variations in labeling of SLF segments. Second, we
propose a simple possible classification of the SLF, mainly
based on the functions of its component tracts. Our classifica-
tion is advantageous in that it focuses on function rather than
anatomical characteristics. It will be useful to researchers and
clinicians engaged in work pertaining to brain function, such
as in the fields of neuroscience, neurology, neurosurgery, and
neuropsychology.
Historical classification of the SLF
SLFs I, II, and III
In the early 1800s, Reil (1809, 1812) revealed the existence of
white matter bundles that communicate with the temporal,
parietal, and frontal lobes. Following those reports, Mayo
(1827), Burdach (1822), and Dejerine (1895) confirmed
Reil’s findings (1809, 1812) through more detailed anatomical
studies. At that time, the terms “SLF” and “AF” were used
interchangeably by Dejerine (1895) and their involvement in
language processing was suggested. In the same period,
Wernicke (1874) found a direct connection involving the lan-
guage network between Broca’s and Wernicke’s areas. Since
the 1980s, some researchers have investigated the structural
connectivity of the SLF because recent developments in neu-
roimaging have enabled the division of the SLF into several
segments. Petrides and Pandya (1984) provided the original
description of the anatomy of the SLF, using autoradiography
on the monkey brain; they divided it into SLFs I, II, III, and
the AF (regarded as the fourth component of the SLF). Their
main findings were as follows: SLF I, or the dorsal compo-
nent, originates from the medial and dorsal parietal cortex and
terminates in the dorsal and medial parts of the frontal lobe.
Brain Imaging and Behavior (2020) 14:2817–2830
SLF II, a major component of the SLF, derives from the caudal
inferior parietal region, corresponding to the angular gyrus
(AG) in humans, and terminates within the dorsolateral frontal
region. SLF III, or the ventral component, originates from the
supramarginal gyrus (SMG) and terminates predominantly in
the ventral premotor and prefrontal areas. The AF, or fourth
division of the SLF, originates in the caudal part of the supe-
rior temporal gyrus (STG) and runs contiguously with SLF II,
above the Sylvian fissure and insula. In 2006, a more anatom-
ically detailed study of the rhesus monkey was reported by
Schmahmann and Pandya (2006), popularizing the SLF I–III
classification. The first human study employing the SLF I–III
classification was conducted by Makris et al. (2005) in 2005,
using DTI; they reported that the anatomical locations of hu-
man SLFs I–III was nearly consistent with that of the monkey.
Notably, they discriminated between SLFs II and III of the
SLF anterior segment, which links the caudal portion of the
STG and the inferior frontal gyrus (IFG). Subsequently, more
advanced neuroimaging methods have been used instead of
DTI. Thiebaut de Schotten and colleagues (Thiebaut de
Schotten et al. 2012) investigated the anatomical and structur-
al differences among SLFs I, II, III, and the AF between hu-
man and monkey brains using spherical deconvolution (SD)
imaging; the structure of the human SLF, but not the AF, was
reportedly similar to that of the monkey. Specifically, projec-
tions of the AF to the middle and inferior temporal gyrus
absent in the monkey brain. Similarly, some studies also have
found reported the presence of the SLF in animal models
using autography, diffusion spectrum imaging (DSI), and
DTI; most of these studies used the classifications of SLFs I,
II, III, and the AF (Kawamura and Naito 1984; Schmahmann
and Pandya 2006; Schmahmann et al. 2007; Thiebaut de
Schotten et al. 2012; Hecht et al. 2015). A study (Hecht
et al. 2015) that compared the structures of SLFs between
chimpanzees and humans showed findings that were partially
different from those of Thiebaut de Schotten and colleagues
(Thiebaut de Schotten et al. 2012); moreover, it reported that
SLFs II and III had different connectivities between species.
These differences may be related to uniquely human abilities:
the AF plays a critical role in a language-related network, and
SLFs II and III to spatial action, tool use, and social learning.
Many of the studies employing the SLF I–III classification
have been neuroimaging studies, but the existence of anatomy
corresponding to this classification had not been proven by
postmortem fiber dissection until very recently. The connec-
tivities of SLFs I, II, and III were investigated by Wang et al.
(2016) using DSI, which employed fiber tracking on a 30-
subject DSI template (Carnegie Mellon University-30;
CMU-30) and a 488-subject template from the Human
Connectome Project (HCP-488). Notably, the DTI technique
frequently finds false negatives at points of crossing, kissing,
and twisting fibers; the tensor models do not perform well.
DSI comprises an alternative method to address this problem.
Brain Imaging and Behavior (2020) 14:2817–2830
Importantly, the study by Wang and colleagues (Wang et al.
2016) demonstrated that SLFs II and III could be identified as
isolated fiber bundles. SLF I, however, could not be located as
a separate structure, and the authors concluded that it would be
preferable to designate SLF I as a component of the cingulum,
from both anatomical and functional perspectives.
Conversely, Yagmurlu and colleagues (Yagmurlu et al.
2016) first provided evidence for the existence of SLF I using
fiber dissection with 25 formalin-fixed brains and three adult
cadaveric heads; this finding was supported by the results of a
very recent fiber dissection study (Komaitis et al. 2019),
which revealed that SLF I was a long association fiber tract
located medially to the callosal fibers and above the cingulum
tract, connecting the precuneus and cingulate cortex.
Anterior, posterior, and long segments of the SLF
Another major classification was provided by Catani and col-
leagues (Catani et al. 2005) in 2005: the SLF/AF anterior
segment, posterior segment, and long segment (the classical
AF). They classified the fiber bundle traditionally known as
the AF into three segments. In general, the anterior segment
and horizontal segment are considered to be synonymous; the
posterior segment and vertical or temporoparietal segment
(SLF-tp) are also considered to be synonymous (Makris
et al. 2005; Martino et al. 2011; Kamali et al. 2014b; Vassal
et al. 2016). Catani et al. (2005) described the structural con-
nectivity of the SLF as follows: the anterior segment links the
parietal and frontal lobes, mainly the SMG to the middle fron-
tal gyrus (MFG) and IFG, including the pars opercularis and
ventral premotor cortex (vPMC), and runs dorsal to the exter-
nal capsule and insular cortex. The posterior segment connects
the temporal and parietal lobes, mostly the posterior and mid-
dle thirds of the middle temporal gyrus (MTG) and STG, to
the posteroventral portion of the inferior parietal lobe (IPL)
and superior parietal lobe (SPL). Moreover, the long segment,
which corresponds to the classical AF, connects the frontal
and temporal lobes, primarily the IFG and MFG including
the vPMC to the superior, middle, and inferior temporal
gyri, and is located in the most medial part of the SLF. In
contrast to previous studies, Catani et al. (2005) demonstrated
that the AF did not communicate between Broca’s and
Wernicke’s areas. Since then, the findings of several studies
have supported the existence of the SLF as classified by
Catani et al. (2005), using neuroimaging techniques such as
DTI and DSI, as well as fiber dissection (Fernandez-Miranda
et al. 2008a; Lawes et al. 2008; Martino et al. 2011; Martino
et al. 2013; De Benedictis et al. 2014).
Recently, some researchers classified the SLF long seg-
ment into ventral and dorsal components of the AF using
DTI or fiber dissection (Yagmurlu et al. 2016; Fernandez-
Miranda et al. 2015). These studies identified the following
trajectories: the ventral AF originated in the middle and
2819
posterior parts of the STG and middle part of the MTG, then
passed the SMG. The dorsal AF originated in the posterior
part of the MTG and inferior temporal gyrus (ITG) and passed
the AG. Subsequently, both segments terminated in the frontal
lobe. Very recently, a combined study of DSI and fiber dissec-
tion suggested that the AF posterior segment could be divided
into two subdivisions, namely dorsal and ventral segments,
and proposed re-naming this segment as the “temporo-parietal
aslant tract” (Panesar et al. 2019). Additionally, recent DTI
studies have indicated that the posterior segment could be
divided into two further segments, namely the SLF/AF
temporo-parietal superior parietal lobe (TP-SPL) and
temporo-parietal inferior parietal lobe (TP-IPL). The TP-IPL
segment corresponds to the traditional posterior segment; it
originates in the caudal and posterior aspects of the STG and
MTG and terminates in the IPL (Kamali et al. 2014b). The
SLF/AF TP-SPL runs from the STG and MTG to the SPL
(Brodmann area [BA] 7) (Kamali et al. 2014a). Similarly,
using DSI and fiber dissection, Wu et al. (2016) also demon-
strated that the posterior segment extended to a broader range:
it originated from the MTG and ITG and terminated in the
SMG.
Multiple definitions and labeling of the SLF
Today, the classification of the SLF is confusing, although
some researchers have summarized the anatomical location
of the SLF based on language-related function (Bernal and
Altman 2010; Dick and Tremblay 2012). The confusion prob-
ably arises because readers of the literature are required to
consider the particular classification system used in each pa-
per. Moreover, the term “SLF” sometimes signifies only one
segment of the SLF; at other times, it represents the entire
frontoparietal network. We suppose that most classification
differences among recent studies are simply due to differences
in labeling procedures. In addition to these differences in lan-
guage, the following reasons may cause confusion.
The variably defined AF: as mentioned above, Catani et al.
(2005) defined the traditional AF as “the AF, namely SLF
(AF/SLF),” and classified it into three segments (anterior, pos-
terior, and long). Since then, two context-dependent defini-
tions of the AF have been used: 1) “the AF” as a synonym
of “the SLF,” in which the AF is classified into the SLF/AF
anterior, posterior, and long segments; and 2) “the AF” as a
synonym for “the AF long segment” or classical AF, while the
term “the SLF” is used to describe the frontoparietal connec-
tion. Thus, when the AF is referred to without clarification
(e.g., only “the AF”), the reader must determine which of
the two definitions was intended, and often this is not trivial.
In recent literature, “the AF” without clarification seems most
often to refer to the classical AF or the long segment AF/SLF
(Vassal et al. 2016; Petrides and Pandya 1984; Thiebaut de
Schotten et al. 2012; Schmahmann and Pandya 2006; Bernal
2820
and Altman 2010). Additionally, when “the AF” without clar-
ification is used, the term “SLF” often represents the
frontoparietal network (Petrides and Pandya 1984; Thiebaut
de Schotten et al. 2012; Schmahmann and Pandya 2006).
Lack of equivalence among classifications: most studies
define the anterior segment as equal to SLF III
(Makris et al. 2005; X. Wang et al. 2016). However,
some researchers regard the posterior segment as equiv-
alent to SLF III (De Benedictis et al. 2014). Anatomically,
SLF II links the IPL to the middle and superior frontal gyri
(SFG and MFG); it runs medial and dorsal to the area
of SLF III. In contrast, SLF III and the SLF/AF anterior
segment originate from the IPL and posterior part of the
STG and terminate in the MFG and IFG. Furthermore,
the posterior segment connects the STG and MTG to the IPL.
It is therefore preferable to regard the anterior segment as
equal to SLF III.
Disagreement in terms of the existence of SLF I: in the past,
the existence of SLF I has been demonstrated using neuroim-
aging, including autoradiography, DTI, DSI, and SD (Petrides
and Pandya 1984; Makris et al. 2005; Jang and Hong 2012;
Thiebaut de Schotten et al. 2012; Thiebaut de Schotten et al.
2011a); however, fiber dissection has not yet provided direct
evidence of its existence. Most recently, two fiber dissection
studies by Yagmurlu et al. (2016) and Komaitis et al. (2019)
successfully identified SLF I as an isolated long association
fiber. In contrast, a fiber dissection study conducted by Wang
and colleagues (Wang et al. 2016) indicated that SLF I could
not be found as an isolated fiber; it was instead found as a
component of the cingulum. Notably, a number of fiber dis-
section studies have reported the existence of all of the SLFs,
including SLFs II and III and the SLF/AF anterior, posterior,
and long segments, with the exception of SLF I (Lawes et al.
2008; Martino et al. 2011; Martino et al. 2013; De Benedictis
et al. 2014; Gungor et al. 2017). However, sometimes it is
difficult to distinguish SLF II from SLF III (equal to the ante-
rior segment) at the horizontal section (Makris et al. 2005).
From a functional perspective, SLF I and the cingulum
are associated with similar functions, as both fibers play
major roles in the attentional network. The cingulum is
involved in attentional network functions including inhi-
bition, working memory, and executive function.
Similarly, SLF I is mainly involved in attentional functions
such as top-down attention, voluntary orientation of attention,
and working memory.
As described above, there are presently two opposite
views: existence vs. non-existence of SLF I. The SLF/AF
was initially defined as a group of lateral fibers located around
the Sylvian fissure, running deeply into the white matter of the
temporal, parietal, and frontal regions. This view is also sup-
ported by recent studies, which indicated that SLF I is located
medially and connects functionally with the cingulate cortex
(Yagmurlu et al. 2016). Hence, we used the original definition
Brain Imaging and Behavior (2020) 14:2817–2830
of the SLF/AF, in which lateral tracts are regarded as the SLF/
AF, and the medial tract (SLF I) is not included in the SLF/AF.
Function of the SLF in each subdivision
As shown above, although there are various systems of label-
ing, we assembled the functions of the SLF in each segment,
and classified them into four subdivisions. Each of the SLF
subdivisions has a unique function, as evidenced by a growing
number of studies. In this section, we review this functional
segregation of the SLF.
SLF II and its functions
SLF II is regarded as the dorsal component of the SLF. It is a
major domain of the dorsal pathway of attention, with a prin-
cipal function involving visuospatial awareness, especially in
the non-dominant cerebral hemisphere (Urbanski et al. 2008;
Nakajima et al. 2017; Shulman et al. 2010; Corbetta and
Shulman 2002). Clinically, disturbance of visuospatial aware-
ness is regarded as unilateral spatial neglect, which is causally
related to damage of the frontoparietal network or inferior
fronto-occipital fasciculus; it induces attentional decline in
the left space, regardless of the existence of a visual-field
disorder (Vallar et al. 2014; Thiebaut de Schotten et al.
2014; Bartolomeo et al. 2007). Additionally, SLF II is broadly
involved in motor control; together with the ventral pathway,
it is involved in visuomotor processing, coordinating such
behaviors as eye movement paired with the grasping of an
object (Berman et al. 1999; Buch et al. 2012; Budisavljevic
et al. 2017; Montemurro et al. 2016). The typical symptom
induced by damage in SLF II is optic ataxia (Karnath and
Perenin 2005). In addition, the dorsal pathway is in-
volved in tool use, especially the understanding of
movement during tool use; therefore, disconnection of
the left SLF II induces apraxia, a disorder of tool use
(Timpert et al. 2015; Vry et al. 2015; Hoeren et al.
2013). SLF II is also involved in executive function,
including working memory. Typically, right-hand con-
nectivity is related to spatial working memory (Kinoshita
et al. 2016b; Curtis 2006), while left-hand connectivity is re-
lated to verbal working memory (Duffau et al. 2014;
Karlsgodt et al. 2008; Volle et al. 2008). Additionally, the
dominant SLF II ends in Exner’s area, which has a crucial role
in handwriting; disruption of the dorsal pathway may lead to
agraphia (Klein et al. 2016).
SLF III or anterior AF and its functions
The SLF III, which is a ventral segment of the SLF, corre-
sponds to the AF anterior or horizontal segment. The most
important function of the left ventral segment is related to a
Brain Imaging and Behavior (2020) 14:2817–2830
language network, involving articulatory processing and au-
ditory language comprehension (Kellmeyer et al. 2013; Saur
et al. 2010; Duffau 2008; Goranskaya et al. 2016). The in-
volvement of SLF III in language function can be confirmed
by direct electrical stimulation during awake craniotomy be-
cause stimulation of SLF III generates language disturbances
(Martino et al. 2011; Maldonado et al. 2011b; van Geemen
et al. 2014). Involvement in this language network is the most
remarkable functional difference between the ventral and
dorsal pathways. Recently, Papagno et al. (2017) revealed that
the left anterior segment of the AF was related to verbal mem-
ory, especially the order information of memory, by direct
electrical stimulation during awake surgery. In addition, SLF
III is associated with motor control, specifically motor plan-
ning, motor imagery, and recognition of postural changes in-
cluding kinesthetic emulations, in combination with the dorsal
pathway (Buch et al. 2012; Budisavljevic et al. 2017; Vaessen
et al. 2016; Johnson-Frey et al. 2005). The ventral pathway is
a component of the attentional network and is involved in
attentional function or alertness during unconsciousness, spe-
cifically bottom-up attention (Shulman et al. 2009; Martin-
Signes et al. 2019; Klarborg et al. 2013). The right SLF III
has lateralized involvement in spatial awareness, in combina-
tion with the SLF II or inferior fronto-occipital fasciculus;
therefore, disconnection of SLF III may lead to spatial neglect
(Doricchi et al. 2008; Toba et al. 2018; Thiebaut de Schotten
et al. 2005; Bartolomeo et al. 2007). A recent study proposed
that the right SLF III may process information associated with
the bodily self, as it plays a role in both self-face recognition
and proprioceptive illusion (Morita et al. 2017). Additionally,
regarding the mirror neuron system, SLF III is involved with
social cognition, which comprises prediction of behavior or
Fig. 1 Possible four-term clas-
sification of the SLF and its
anatomy. Number tags indicate
Brodmann areas. Although this
figure shows only the origin and
terminus of each fiber, some
fibers end in the middle of the
gyrus. Cyan, dorsal SLF;
magenta, ventral SLF; blue,
posterior SLF; green, AF. AF:
arcuate fasciculus, SLF: superior
longitudinal fasciculus
2821
emotional state in another person, based on external informa-
tion (Im et al. 2018; Nakajima et al. 2018a; Y. Wang et al.
2018).
SLF-tp or posterior AF and its functions
The traditional classification systems consider the SLF-tp, AF
posterior segment, and vertical segment of the SLF to be the
same tract. The posterior SLF of the dominant cerebral
hemisphere is primarily associated with a language-
related network. The posterior pathway plays an impor-
tant role in auditory comprehension, and emphasis has
been placed on the intraoperative assessment and pres-
ervation of its function during brain surgery (Catani et al.
2005; Fernandez-Miranda et al. 2008b; Maldonado et al.
2011b; Martino et al. 2011). Additionally, together with the
ventral SLF, the posterior pathway is involved in the lexical
retrieval stage, which comprises the process of retrieving the
corresponding lexical unit and phonological word form
(Herbet et al. 2016; Levelt et al. 1999). Disruption of the
posterior SLF therefore generates anomia (Sarubbo et al.
2015). Moreover, because the posterior AF is involved in an
interactive feedback system between visual and nonvisual in-
formation, damage to the posterior SLF may induce alexia
(Zemmoura et al. 2015).
Conversely, little is known regarding the functional role of
the non-dominant posterior SLF. Galantucci et al. (2011) pre-
sumed that the right posterior segment might relate to
logopedic aphasia together with the left SLF, but its functional
role remains unclear. Although the evidence is thus far insuf-
ficient, some studies have suggested that the right posterior
AF may be involved in auditory spatial attention (Barrick et al.
2822 Brain Imaging and Behavior (2020) 14:2817–2830

2007), since the posterior segment links the primary auditory
cortex of the STG and the parietal lobe, which functions
in auditory spatial attention (Griffiths et al. 1998, 2000).
In addition, this pathway may be involved in visuospa-
tial cognition because the posterior pathway connects
the visuospatial-related cortices, including the right pa-
rietal and temporal cortices. Clinically, damage in the
right temporal region induces the allocentric (object-
based) aspect of spatial neglect, while damage in the
right parietal region causes the visuospatial aspect of
spatial neglect (Verdon et al. 2010; Corbetta and
Shulman 2011; Karnath et al. 2011; Karnath and
Rorden 2012). A recent report indicated that the poste-
rior SLF was related to perceptual spatial neglect; this
could be assessed by a line bisection test or text reading
(Vaessen et al. 2016).
AF or AF long segment and its functions
The traditional classification systems consider the AF and AF
long segment to be the same tract; it is sometimes regarded as
the long segment of the perisylvian SLF. The AF communi-
cates with the auditory area and the posterior portion of the
MFG and IFG; it plays a major role in the processing of spatial
attributes of acoustic stimuli and auditory-related processing,
the so-called phonological stream (Duffau 2008; Axer et al.
2013; Duffau et al. 2014). In general, disruption of the left AF
induces semantic variant aphasia and disconnection of the AF
causes logopedic aphasia (Galantucci et al. 2011).
Furthermore, when the AF is stimulated directly during awake
craniotomy, phonological disorder is elicited, which enables
identification of the fiber (Sarubbo et al. 2015; Maldonado
et al. 2011a, 2011b). Additionally, the dominant AF may play

Fig. 2 Differences between the

existing major classifications
and our proposed classification.
The upper column (a) shows the
existing major classifications of
the SLF. Notably, the existence of
SLF I is currently a matter of
debate. In the current review, we
used the original definition of the
SLF/AF, namely lateral tracts are
regarded as the SLF/AF; a medial
tract was not included in the
SLF/AF. The lower column (b)
shows our proposed classification
system. We collected and
reassessed the existing classifica-
tion system and hereby propose a
simple four-term classification of
the SLF: the dorsal SLF (cyan),
ventral SLF (magenta), posterior
SLF (blue), and AF (green). AF:
arcuate fasciculus, SLF: superior
longitudinal fasciculus, SLF-tp:
temporoparietal segment
Brain Imaging and Behavior (2020) 14:2817–2830 2823

Fig. 3 Tractography of the SLF using DSI. Tractography of right (a)
and left (b) SLFs classified into four segments. Tractography was
obtained by DSI using the HCP-842 atlas (1-mm template). We can
identify the lateralization as previously reported, with the strongest later-
alization observed in the AF (green, right < left). Regions of interest were
placed bilaterally as follows; for the dorsal SLF, namely SLF II, ROIs
were placed at the AG, posterior SFG, and MFG; for the ventral SLF,
namely SLF III or anterior AF, ROIs were placed at the SMG, MFG, and
IFG; for the posterior SLF, namely the SLF-tp or posterior AF, ROIs were
placed at the posterior MTG, STG, and IPL; for the AF, namely the
classical AF or AF long segment, ROIs were placed at the posterior
ITG, MTG, and posterior IFG and MFG. B, back; F, front; L, lateral.
Cyan, dorsal SLF; magenta, ventral SLF; blue, posterior SLF; green,
AF. AF: arcuate fasciculus, DSI: diffusion spectrum imaging, SLF: supe-
rior longitudinal fasciculus

a role in verbal working memory (Meyer et al. 2014), and especially low-level mentalizing processes; it enables under-
verbal fluency (Li et al. 2017). Beyond the language-related standing of emotional states in other people based on their
network, the non-dominant AF is engaged in social cognition, facial expression, eye gaze, and other cues (Herbet et al.

Fig. 4 DTI tractography of the

SLF. An illustrative case of DTI
tractography of the right and left
SLFs in a healthy adult man is
shown. The four segments of the
SLFs were clearly identified in
this case. However, as has been
shown in prior studies,
assessments with DTI may yield
false negatives, as the tensor
model does not perform well at
points of crossing, kissing, and
twisting fibers. Cyan, dorsal SLF;
magenta, ventral SLF; blue,
posterior SLF; green, AF. AF:
arcuate fasciculus, DTI: diffusion
tensor imaging, SLF: superior
longitudinal fasciculus
2824
2014; Nakajima et al. 2018b). Moreover, the right AF is in-
volved in the attentional network, including visual search;
specifically, it plays a role in more complicated visual
search tasks, such as those where a participant must
locate the target in the presence of interference
(Hattori et al. 2018; Suchan et al. 2014). Furthermore,
the right AF is associated with prosodic processing and
may be a component of a language network (Glasser and
Rilling 2008). Interestingly, abnormal neurodevelopment of
the right AF leads to amusia, which is disrupted music pitch,
Fig. 5 SLF fiber dissection. The findings of fiber dissection of the brain
of a female donor, aged 88 years, are shown as a representative case. We
employed the medial approach to expose the SLF in this study. Yellow ball,
triangular part of the frontal operculum; white ball, central sulcus; red ball,
opercular part of the frontal operculum; blue ball, transverse temporal. a
Anatomical location of the lateral aspect of the left hemisphere after
removal of the insula. b-1 The SLF in the superficial (= medial) white
matter of the right hemisphere. Note that the insula has been removed.
Note also that some long, arcuate fibers originate from Heschl’s gyrus,
wind spirally and counterclockwise in the inferior parietal area, and reach
the frontal operculum. The arcuate fibers also include those from the
inferior temporal gyrus. Posterior SLF fibers are often observed in this
layer. c-1 The SLF in the superficial (= medial) white matter of the left
hemisphere. The long arcuate fibers, which have a fishing-hook shape, run
along the superior peri-insular sulcus. Some short fibers of the SLF
Brain Imaging and Behavior (2020) 14:2817–2830
while abnormality of the left AF induces speech tone deficit
(Chen et al. 2018).
Proposed simple classification of the SLF
Based on the presented information, we propose a simple
classification system of the SLF to be used as an alternative
to the classical terminology: the dorsal SLF, ventral SLF, pos-
terior SLF, and the AF (Fig. 1). Our classification enables
terminate near the precentral sulcus. The white ball indicates the central
sulcus. b-2, c-2 The segregated SLF in the deep (= lateral) white matter
layer. As the dissection advances into the deep white-matter layer, the long
fibers of the SLF terminate in the supramarginal and angular gyri (short
arrow); the SLF thus segregates into the anterior and posterior SLF. The
anterior SLF is further divided into the dorsal and ventral SLF (b-2). The
posterior SLF is present in the deep layer (c-2). AF, arcuate fasciculus; AG,
angular gyrus; CS, central sulcus; d-SLF, dorsal superior longitudinal fas-
ciculus (SLF); IFG, inferior frontal gyrus; ITG, inferior temporal gyrus;
MFG, middle frontal gyrus; MTG, middle temporal gyrus; OFC, orbito-
frontal cortex; Op, opercular part of the frontal operculum; p-SLF, posterior
SLF; post-CG, post central gyrus; pre-CG, pre central gyrus; SFG, superior
frontal gyrus; SMG, super marginal gyrus; TP, temporal pole; Tri, triangular
part of frontal operculum; vPMC, ventral premotor cortex; v-SLF, ventral
SLF
Brain Imaging and Behavior (2020) 14:2817–2830 2825

utilization of several traditional classifications of the SLF and
AF, with concurrent acknowledgment of the functional role of
each segment. The differences between the existing major
classifications and our proposed classification are summarized
in Fig. 2. Note that the terms “ventral SLF” and “dorsal SLF”
have been previously used by some researchers (Shulman
et al. 2009; X. Wang et al. 2016). The dorsal segment of the
SLF originates from the IPL (e.g., the AG) and terminates in
the SFG and MFG (BAs 6, 8, and 46); it corresponds to the
classical SLF II. The ventral segment, namely the classical
SLF III or AF anterior segment, communicates the IPL (e.g.,
the SMG) and the MFG and IFG, including the vPMC (BAs 6
and 44). The posterior segment, alternatively known as the
traditional posterior segment or SLF-tp, begins principally in
the posterior parts of the MTG and STG and terminates in the
IPL and SPL. Recent findings suggest that more extensive
regions may be governed by the posterior segment: a portion
of the posterior segment arises from the ITG and terminates in
the SPL (Barrick et al. 2007; Kamali et al. 2014b; Wu et al.
2016). The AF, alternatively known as the classical AF or AF
long segment, originates from the ITG, MTG, and STG before
curving around the insula and caudal Sylvian fissure, then
terminating in the posterior IFG and MFG, including the
vPMC. These four subdivisions have been validated by both
fiber dissection and neuroimaging studies in humans
(Thiebaut de Schotten et al. 2012; Martino et al. 2013;
Kamali et al. 2014a; Panesar et al. 2019), although there were
inter-subject variances (Catani et al. 2005; X. Wang et al.
2016, See also A 2).
We intend to use the following three-images to show that
these subdivisions of the SLF, based on previous findings, can
be clearly observed by neuroimaging methods and fiber dis-
section. Figure 3 shows the tractography of ambilateral SLFs
classified into four segments using DSI Studio (http://dsi-
studio.labsolver.org). We used the HCP-842 atlas (1-mm tem-
plate), which was constructed using the diffusion MRI data of
842 subjects from the Human Connectome Project. The dif-
fusion data were reconstructed in the Montreal Neurological
Institute space using q-space diffeomorphic reconstruction.
For the SLF reconstruction, regions of interest (ROIs) were
placed bilaterally at appropriate locations as previously de-
scribed (Kamali et al. 2014a; Thiebaut de Schotten et al.
2012; Martino et al. 2013; X. Wang et al. 2016). To maximize
the detection of the studied tracts, the following tracking pa-
rameters were applied: an angular threshold of 60, step size of
0.5 mm, smoothing of 0, and length constraint of 0 to 500 mm.

Fig. 6 Possible classification

system and key function of each
segment. The structural
connections and central functions
of each segment are summarized.
Notably, depending on the
function, there exists obvious
functional lateralization within
each segment. Although some
fibers originate from the temporal
pole and terminate in the
precuneus, most fibers run as
indicated by the blue line. Cyan,
dorsal SLF; magenta, ventral
SLF; blue, posterior SLF; green,
AF. AG, angular gyrus; B,
bilateral; IFG, inferior frontal
gyrus; IPL, inferior parietal lobe;
ITG, inferior temporal gyrus; L,
left; MFG, middle frontal gyrus;
MTG, middle temporal gyrus; p-,
posterior; R, right; SFG, superior
frontal gyrus; SLF, superior
longitudinal fasciculus; SMG,
super marginal gyrus; STG,
superior temporal gyrus
2826
All fibers terminating outside the trajectory of the involved
tracts were deleted (A3; Supplementary video data). We used
DTI, another method, to show the four segments of the SLF
(Fig. 4). Structural MR images were acquired using conven-
tional, high-resolution, three-dimensional, T1-weighted se-
quences on a 3.0 Tesla MRI scanner (Signa Excite HDx 3.
0 T, General Electric Medical Systems). A series of diffusion-
weighted axial images was obtained, with (b-value =
1000 s/mm 2 ) and without (b-value = 0) a diffusion-
sensitizing gradient along 30 directions. The remaining
diffusion parameters were as follows: repetition time =
14,000 ms, echo time = 69.6 ms, number of excitations =
2, number of axial slices = 60, slice thickness = 2.5 mm
with no interslice gap, field of view = 220 × 220 mm,
and matrix = 88 × 88, resulting in an effective resolution
of 2.5 mm3 isotropic voxels. ROIs targeting each white
matter tract, including the SLF, were selected manually
as previously described (Kinoshita et al. 2016a). All
three-dimensional tracts were reconstructed with fiber
propagation at a fractional anisotropy threshold of <0.
18. Mean FA values (ranges) of each tract were as fol-
lows: left dorsal SLF = 0.39 (0.18–0.81), left ventral
SLF = 0.39 (0.18–0.68), left posterior SLF = 0.39 (0.18–
0.65), left AF = 0.42 (0.18–0.65); right dorsal SLF = 0.
39 (0.18–0.75), right ventral SLF = 0.46 (0.18–0.75),
right posterior SLF = 0.41 (0.41–0.76), and right AF =
0.45 (0.18–0.75).
Additionally, we regard the results of fiber dissection as the
warrant for a new nomenclature of the SLF subdivisions.
There are advantages and limitations in both fiber dis-
section and neuroimaging techniques, including the DSI
and DTI methods (Dell'Acqua and Catani 2012). Since
both of them redeem limitation of the other, we used
fiber dissection. Here, we show an illustrative case of
the use of fiber dissection (Fig. 5). The brain was main-
tained and used in accordance with the guidelines of the
Internal Review Board of Kanazawa University (Nos.
2348 and 2957). As shown in Fig. 5, the SLFs had four
clear subdivisions.
Our current proposed classification is mainly based on the
functions outlined below. Hence, the classification is compre-
hensible and easy for clinicians to use when considering in-
terrelations between white matter damage and functional def-
icits. We summarized the functions of each SLF subdivision in
Fig. 6. Moreover, the relationships between function and each
SLF segment correspond well to the results of direct electrical-
stimulation studies during awake surgery (Kinoshita et al.
2016b; Martino et al. 2011; Sarubbo et al. 2015). Awake sur-
gery is a brain surgery method that is performed under wake-
fulness to preserve brain function, such as language and sen-
sorimotor function. Preservation of function can be confirmed
because localization in the brain is assessed through direct
electrical stimulation of cortices or white matter tracts and
Brain Imaging and Behavior (2020) 14:2817–2830
evaluation with appropriate brain function tests. Hence, these
findings can help locate SLFs effectively during awake sur-
gery, through symptoms elicited by direct electrical
stimulation.
Lateralization
Finally, it is important to discuss lateralization when consider-
ing the human white-matter network. It is common knowledge
that lateralization exists in the SLF and is related to cerebral
function in most cases. For instance, the frontoparietal pathway
(dorsal and ventral pathways) is lateralized in the right cerebral
hemisphere and is associated with visuospatial processing
(Makris et al. 2005; Thiebaut de Schotten et al. 2011b).
However, regarding the ventral pathway, one study reported
lateralization in the left hemisphere (Barrick et al. 2007).
Although little is known about the actual functional role of
the posterior segment, lateralization exists in the right hemi-
sphere and may be associated with auditory spatial attention
(Barrick et al. 2007). The strongest lateralization is observed
anatomically and functionally in the AF, which may make hu-
man brain unique; this lateralization is specifically present in
the dominant cerebral hemisphere (Barrick et al. 2007; Parker
et al. 2005; Eichert et al. 2018; Eluvathingal et al. 2007;
Thiebaut de Schotten et al. 2011b; Upadhyay et al. 2008).
Previous studies revealed that this lateralization is related to
language function, including reading and cognitive function
(Barrick et al. 2007; Lebel and Beaulieu 2009; Thiebaut de
Schotten et al. 2011b). Notably, this lateralization does not al-
ways exist and may differ depending on handedness or sex
(Hagmann et al. 2006; Herve et al. 2006; Herve et al. 2009;
Lebel and Beaulieu 2009).
Conclusions
This review assembled and revisited previous knowledge with
respect to the SLF and proposed an alternative to the tradition-
al classification systems: a simple classification including the
dorsal SLF, ventral SLF, posterior SLF, and the AF. This mod-
el fits well with the anatomical structure and functional segre-
gation of the SLF and is supported by neuroimaging, fiber
dissection, and neuropsychological findings. Our proposal of
a four-term classification of the SLF will be a convenient and
useful tool for disciplines involving brain function, such as
neuroscience, neurology, and neurosurgery. However, some
functions of the SLF, and the interactions of each segment,
remain unclear. Further research is thus required before a com-
prehensive description of this extensive white matter network
is possible.
Funding This research did not receive any specific grant from funding
agencies in the public, commercial, or not-for-profit sectors.
Brain Imaging and Behavior (2020) 14:2817–2830
Compliance with ethical standards
Conflict of interest The authors report no conflict of interest concerning
the materials or methods used in this study or the findings specified in this
paper.
Ethical approval All procedures performed in studies involving human
participants were in accordance with the ethical standards of the
institutional and/or national research committee and with the 1964
Helsinki declaration and its later amendments or comparable eth-
ical standards.
Informed consent Informed consent was obtained from the families of
all individual participants included in the studies reviewed in this paper.
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