Chapter 29
Beyond ERP and fMRI: Other
Imaging Techniques for
Studying Human Brain
Function
Gabriele Gratton and Monica Fabiani
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The past few decades have seen an explosion of psy-
chological research using noninvasive (or low-
invasivity) measures of human brain function. This
work is motivated by two main goals: Measures of
brain function may (a) provide additional informa-
tion, beyond that obtained from overt response vari-
ables, about intermediate states of the information
processing system and (b) illuminate some of the
physiological mechanisms that underlie the psycho-
logical phenomena of interest. The vast majority of
this work has employed two measures: event-related
brain potentials (ERPs; see Fabiani, Gratton, &
Federmeier, 2007) and functional magnetic reso-
nance imaging (fMRI; see Van Horn & Poldrack,
2009). Brain function, however, is a complex phe-
nomenon, and several other measures are currently
available that allow investigators to explore its vari-
ous facets. These measures complement ERPs and
fMRI, generating a more articulated view of this
enormously complex organ’s functions. In this chap-
ter, we review some of these methods, their ratio-
nale, the types of data they provide, and their
advantages and limitations. Specifically, we review
the following techniques: magnetoencephalography
(MEG); single-photon emission computerized
tomography (SPECT) and its higher resolution
counterpart, positron emission tomography (PET);
fast (i.e., the event-related-optical signal [EROS])
and slow (i.e., near-infrared spectroscopy [NIRS])
optical methods; and transcranial magnetic stimula-
tion (TMS). These techniques vary widely with
We acknowledge the support of National Institute of Mental Health Grant MH80182 to G. Gratton and of National Institute on Aging Grant
1RC1AG035927 to M. Fabiani.
DOI: 10.1037/13619-030
APA Handbook of Research Methods in Psychology: Vol. 1. Foundations, Planning, Measures, and Psychometrics, H. Cooper (Editor-in-Chief)
Copyright © 2012 by the American Psychological Association. All rights reserved.
respect to the physical mechanisms involved, the
types of data they can provide, and the subject pop-
ulations to and environments in which they can be
applied. They also do not exhaust all possible physi-
ological techniques that can be used to study human
brain function in a noninvasive manner, but they do
represent the most used complementary approaches
to ERPs and fMRI found in the literature in the past
decade or 2.
Review of Methods
Magnetoencephalography
MEG is a technique used to measure the tiny mag-
netic fields linked to the electrical currents associ-
ated with the depolarization and hyperpolarization
of neurons (for reviews see Hari, 1996; Hari,
Levänen, & Raij, 2000; Stufflebeam, Tanaka, & Ahl-
fors, 2009). To be measurable at the surface of the
head, the magnetic fields generated by individual
neurons must summate, which requires that individ-
ual fields be oriented in the same direction. This
means that only certain structures and neuronal
populations possessing a specific geometric organi-
zation may generate surface-recordable MEG (e.g.,
the cortex). Even so, the magnetic fields generated
by the brain’s electrical currents are extremely small
(on the order of 101–103 fT), much smaller than the
magnetic fields from various environmental sources,
such as the Earth’s magnetic field (108 fT). There-
fore their recording requires special instruments
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 Gratton and Fabiani
called superconducting quantum interference devices
(SQUIDs). In current MEG systems, up to 300 dif-
ferent SQUID detectors are used to generate maps of
the magnetic fields over the head. These devices
need to be kept at extremely low temperatures (a
few degrees Kelvin), which permit superconducting
phenomena to occur. Thus, the recording instru-
ment needs to be supercooled, making it bulky and
costly (around $1.5–2 million). In addition, special
methods are used to reduce the influence of envi-
ronmental magnetic sources on the measurements.
These include (a) shielding, which is expensive and
adds significantly to the cost of the machine, and (b)
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special circuit design, which makes MEG particu-
larly sensitive to locally generated compared with
distantly generated fields, measuring field gradients
rather than absolute magnitudes.
Although effective, these circuit designs restrict
MEG measures to activity in fairly superficial areas
of the brain. They also make MEG particularly sen-
sitive to currents flowing in a direction tangential to
the surface of the head compared with currents
flowing in a radial direction. As a consequence,
MEG measures are more sensitive to phenomena
occurring in sulci (which tend to be orthogonal to
the surface of the head, resulting in currents ori-
ented tangentially) than to those occurring in gyri
(which are parallel to the surface of the head, result-
ing in currents oriented orthogonally).
Notwithstanding its high cost, MEG is progres-
sively gaining acceptance from researchers and clini-
cians. Its ability to describe rapid brain events (on
the order of milliseconds) coupled with a spatial res-
olution on the order of a few centimeters make it
useful for a number of research areas. In particular,
MEG has been used extensively in studies of lan-
guage processing (Frye, Rezaie, & Papanicolaou,
2009)1 and to some degree also in studies of atten-
tion and memory (e.g., Hopf, Boehler, Schoenfeld,
Heinze, & Tsotsos, 2010; Stephane et al., 2010). A
particularly attractive feature of MEG is the absence
of invasivity, as no substance or energy is intro-
duced into the body for its measurement. Thus,
MEG can be used with many types of populations,
including children (e.g., Tesan, Johnson, Reid,
This includes the use of MEG to assess the lateralization of language in preparation for surgery, as an alternative to more invasive methods such as the
1
Wada test (e.g., Doss, Zhang, Risse, & Dickens, 2009).
568
Thornton, & Crain, 2010) and even fetuses (e.g.,
Sheridan, Matuz, Draganova, Eswaran, & Preissl,
2010). Specialized instruments, however, are
required for these populations. Furthermore,
because of its high sensitivity to electromagnetic
noise, MEG cannot be easily combined with other
methods for studying brain function (such as mag-
netic resonance [MR] methods).
Single-Photon Emission Computerized
Tomography and Positron Emission
Tomography
SPECT and PET are two related techniques used in
radiology and nuclear medicine. Both techniques
introduce a radioactive substance (radiotracer or
radioligand) into the body and obtain images of its
diffusion in the body by mapping the radioactivity it
generates. SPECT is based on gamma decay, a radio-
active decay characterized by the production of a
single gamma ray by a decaying isotope. The most
common isotope used is 99mTc, which has a half-life
of a couple of hours. PET instead is based on beta
decay, in which a radioactive isotope emits a
positron, which then reacts with electrons in the
medium annihilating and producing a pair of
high-energy gamma rays oriented 180 degrees from
each other. Several types of isotopes are used for
PET, depending on the application, including 11C
(half-life ≈ 20 min), 13N (half-life ≈10 min), 15O
(half-life ≈ 2 min), and 18F (half-life ≈ 110 min).
Because two gamma ray photons traveling in oppo-
site directions are produced, it is possible to deter-
mine a line where the annihilation occurred by
determining whether they arrived almost simultane-
ously (within a few nanoseconds) to two among a
circular array of detectors located around the head.
As the annihilation occurs within 1 cm to 2 cm from
where the radioactive decay happened, this provides
a way of mapping the decay process in the body, or
in its psychology applications, within the brain.
The great advantage of PET and SPECT is that
they can be used for a wide variety of purposes.
Radioligands can be incorporated in a number of
different substances, each of which can provide
interesting information about the physiology and

metabolism of the body. Of specific interest for psy-
chologists are two classes of applications: (a) analy-
sis of blood flow and metabolism in different regions
of the brain (e.g., Petersen, Fox, Posner, Mintun, &
Raichle, 1989) and (b) mapping of neurotransmitter
receptors or other biochemicals of interest within
the brain (e.g., Tateno, Kobayashi, & Saito, 2009).
Analysis of blood flow and metabolism is typi-
cally carried out using PET. Two methods have been
most extensively used: 15O incorporated in water
and 18F incorporated in fluorodeoxyglucose. The
first compound is used to study blood flow. Impor-
tantly for cognitive studies, 15O has a very short half-
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life, allowing for multiple measurements to be taken
within a session. This enables investigators to study
the difference in the amount of blood flowing
through a particular region during different task
conditions, through digital subtraction of different
maps obtained under each condition. This method-
ology, developed by the Washington University
group in the 1980s, has had a significant role in the
history of cognitive neuroscience (see Raichle,
1998). The spatial resolution of the images obtained
with this approach is about 1 cm to 2 cm. The tem-
poral resolution is of the order of minutes. Cur-
rently, however, this approach has been largely
supplanted by fMRI, which provides images with a
greater spatial and, importantly, temporal resolu-
tion. In addition, 15O PET is expensive, requiring a
cyclotron on site and rapid incorporation and deliv-
ery technology.
18F deoxyglucose PET is used to monitor glucose
consumption by tissue. Deoxyglucose is processed
only through the initial stages of oxidative metabo-
lism, and is then retained in the cells for an
extended period of time: Thus, the greater the glu-
cose consumption, the greater the accumulation of
deoxyglucose. Because the half-life of 18F is much
longer than that of 15O, only one measurement can
be carried out within a session, limiting its use in
human studies (a similar technique, autoradiogra-
phy, has been used extensively in animal studies)
because a control condition is not easy to imple-
ment. LaBerge and Buchsbaum (1990) used a
contralateral control approach to provide a reference
By appropriately selecting the pharmachological agent, it is possible to distinguish among different types of neurotransmitter receptors.
2
Beyond ERP and fMRI
for the measurement obtained on the stimulated
side. Note that 15O PET decay is so fast that the
injection of the radioactive material and the psycho-
logical task of interest need to be conducted while
the subject is in the scanner. This is not necessary
with the 18F deoxyglucose technique, which is based
on the accumulation of the radiotracer over an
extended period of time, allowing for the injection
and the psychological task to be conducted before
placing the subject in the scanner. This enables the
use of a greater variety of tasks, although only a
summary image of total deoxyglucose accumulation
during the experiment can be obtained.
In the past decade or so, a number of new
radiotracer molecules have been developed, and
more are currently being tested. These molecules
allow investigators to map a number of important
chemicals in the brain. Specifically, radioisotopes
are incorporated into selective agonists (or antago-
nists) of various neurotransmitters and neuromodu-
lators (e.g., dopamine and acetylcholine), making it
possible to visualize in vivo the distribution and
quantity of the receptors for these substances.2 This
can be used to compare different subject groups,
such as those who differ along some important
physiological, psychological, or genetic dimension.
For example, a recently developed tracer (the Pitts-
burgh compound B, or PiB), allows for the measure-
ment of beta amyloid deposition, which can be used
for in vivo diagnosis of Alzheimer’s disease (e.g.,
Jagust, 2009; Nordberg, 2010). Also, by comparing
images obtained at different times, it is possible to
determine whether certain treatments (e.g., pharma-
cological or psychiatric therapies) lead to changes in
the quantity and distribution of receptors for spe-
cific neurotransmitter within the brain.
From a practical standpoint, several factors have
limited the diffusion of PET and SPECT. PET is very
expensive, with costs of more than several million
dollars. These costs are due to the price of the
scanner and to the necessity for both a cyclotron to
generate the radioisotopes of interest and the
machinery (and expertise) required to incorporate
the radioisotope into the molecule of interest in situ.
Although costs are lower for SPECT, its inferior
569
 Gratton and Fabiani

spatial resolution makes it a less than ideal alterna-

tive to PET. Finally, because both techniques involve
ionizing radiation, there are limitations to their
extensive use for experimental purposes. To guaran-
tee the safety of subjects, the amount of radiation
absorbed during a PET or SPECT experiment is
strictly regulated, as is the number of times a subject
can participate in studies involving these measures.
In summary, PET and SPECT remain important
imaging tools for psychologists in the 21st century,
although the types of questions that are now
addressed with these techniques are different from
those that were addressed 15 to 20 years ago. They are
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no longer the methods of choice to generate images of

brain activity (measured as a change in blood flow)
during a task (as they have been supplanted by fMRI
for this particular application), but their great chemi-
cal flexibility provides an extremely useful approach
for studying brain biochemistry in vivo.

Diffusive Optical Imaging

Diffusive optical imaging is a relatively new imag-
ing modality, with the first human data obtained in
the early 1990s (for reviews see E. Gratton, Fan-
tini, Franceschini, G. Gratton, & Fabiani, 1997; G.
Gratton & Fabiani, 2009; Villringer & Chance,
1997). It studies how light is transmitted and dif-
fused through tissue (in this case the head) to
determine properties of the tissue itself. Light in
the red to near-infrared (NIR) wavelength range
(650–900 nm) is used. At these wavelengths, there
is relatively little light absorption by most tissues
in the human head, permitting photons to travel
several centimeters through the tissue. However,
NIR light (unlike X-rays or gamma rays) is
strongly scattered by head tissues, with mean free
paths on the order of 5 mm or less. For this reason,
photons do not travel straight through the tissue
but tend to diffuse according to a random-walk
process. Therefore, if a source of NIR light is
located on the surface of the head, it will generate
a halo into the tissue itself, with a maximum pene-
tration of a few centimeters. If a detector is also
located on the surface of the tissue, at a distance of
a few centimeters from the source, it will pick up
some of the photons produced by the source (see
Figure 29.1 for an example of recording montage).
Figure 29.1. Typical setup for recording optical
imaging data from a large set of sources and detectors.
Interestingly, the photons generated from the
source and arriving to a detector located on the
same surface will most likely have followed not a
straight trajectory (as would occur in a nonscatter-
ing medium) but rather a curved trajectory reach-
ing some depth in between the source and the
detector. This occurs because the head tissue (a
highly scattering medium) is surrounded by air (a
nonscattering medium). Therefore, photons travel-
ing close to the surface of the medium are likely, in
their random motion, to reach the surface of the
medium before reaching the detector. If this
occurs, they will move out of the medium and
never reach the detector. Because of this principle,
diffusive optical imaging can provide data about
changes in optical properties that occur inside the
head, at depths up to 2 cm to 3 cm. This permits
measurements from cortical areas relatively close
to the surface of the head.

570

For this approach to be useful in functional brain
imaging, changes in optical properties of the brain
need to occur in association with neural activity.
There are at least two phenomena causing changes
in optical properties of the tissue. The first are
changes in the coloration of tissue caused by varia-
tions in the concentration of oxy- and deoxy-
hemoglobin, which have distinctive absorption
spectra in the NIR range (see Frostig, Lieke, Ts’o, &
Grinvald, 1990; Villringer & Chance, 1997). These
effects are related to blood flow changes studied with
fMRI and 15O PET. The second are changes in the
scattering properties of tissue associated with neu-
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ronal activity (G. Gratton, Corballis, Cho, Fabiani,
& Hood, 1995; Rector, Carter, Volegov & George,
2005). These changes are related to the movement of
ions across the neuronal membranes and are in some
way related to ERP and MEG measures.
Both signals have been investigated using optical
imaging, generating two types of imaging modali-
ties: NIRS (related to hemodynamic signals; Kato,
Kamei, Takashima, & Ozaki, 1993; see Villringer &
Chance, 1997) and EROS (related to neuronal sig-
nals; G. Gratton, Corballis, et al., 1995; see G. Grat-
ton & Fabiani, 2009). Two types of instruments
have been most commonly used: continuous-wave
(CW) instruments, in which light sources have a
fixed intensity over time (or are modulated at low
frequencies, < 10 kHz); and frequency-domain (FD)
instruments, in which light sources are modulated
at high frequencies (typically > 100 MHz). Although
CW instruments are simpler and less expensive, FD
instruments measure both the amount of light mov-
ing between sources and detectors and the time
taken by photon to cross this distance (on the order
of nanoseconds). The time measurement can be use-
ful in estimating the extent to which photons scatter
through the tissue. Furthermore, it can be particu-
larly sensitive to deep phenomena, increasing the
penetration of the procedure and making it more
specific for brain (vs. nonbrain) phenomena, thus
providing higher spatial resolution.
Near-infrared spectroscopy. NIRS uses spectros-
copy to estimate changes in the concentration of
The influence of scattering can be calculated using FD instruments, allowing in principle for the measurement of absolute concentrations of oxy- and
3
deoxy-hemoglobin.
Beyond ERP and fMRI
oxy- and deoxy-hemoglobin in tissue. This is done
by comparing changes in light absorption occur-
ring in different conditions at different wavelengths,
exploiting the fact that oxy- and deoxy-hemoglobin
have different absorption spectra within the NIR
range. This can be accomplished through a system
of linear equations on the basis of the modified
Beer-Lambert Law (see Villringer & Chance, 1997),
which states that the proportion of light absorbed
by tissue is related to the concentration of the
absorbers in the tissue (in this case oxy- and deoxy-
hemoglobin), their coefficients of extinction (i.e.,
ability to absorb light of a particular wavelength),
and the length of the photon path through the tis-
sue (which in most cases is estimated on the basis
of the source-detector distance and a correction
factor derived from the literature to account for
scattering).3 For these computations, a minimum
of two light wavelengths with distinct extinction
coefficients needs to be used; some instruments use
three or more. Because a number of source-detector
pairs can be used within a study, maps of oxy- and
deoxy-hemoglobin concentration changes can be
obtained. Maps can be obtained very quickly, up to
100 or more times per second. Thus NIRS yields
maps of brain activity with a spatial resolution of
1 cm to 2 cm and a temporal resolution limited only
by the time required for hemodynamic phenomena
to occur (a few seconds).
In the past few years, the use of NIRS has
increased. Compared with fMRI (the most compara-
ble imaging method), it has some disadvantages but
also some advantages. Unlike MR technology, cur-
rent optical methods do not provide anatomical
images. Optical data can, however, be coregistered
with MR data with relatively good accuracy
(Whalen, Maclin, Fabiani, & Gratton, 2008). Opti-
cal methods have limited penetration through the
head and therefore, unlike fMRI, cannot be used to
image deep structures (i.e., located more than
3–4 cm from the head surface). The spatial resolu-
tion of NIRS data (1–2 cm) is more limited than that
of fMRI (which in some cases can be as low as a few
millimeters but most typically is in the 5–10 mm
range). Optical data also have a number of advantages.
571
 Gratton and Fabiani
Their recording is flexible and does not require
bulky or expensive equipment. This makes them
usable in a variety of different environments and
populations, such as with children and other
patients who may be difficult to image with fMRI.
Optical technology is completely compatible with
MR (e.g., Toronov, Zhang, Fabiani, Gratton, &
Webb, 2005; Zhang, Toronov, Fabiani, Gratton, &
Webb, 2005), electrophysiological methods (includ-
ing electroencephalogram [EEG], ERPs, and MEG),
and TMS, allowing for the simultaneous recording
of different modalities. It is also possible to obtain
NIRS and EROS data (see the next section) simulta-
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neously with the same instrument, providing con-
current neuronal and hemodynamic measures. NIRS
can provide separate measurements of oxy- and
deoxy-hemoglobin concentrations, which is not nor-
mally possible with fMRI. In fact, if FD technology
is used, absolute concentrations of these substances
can be obtained, again something not possible with
fMRI. Finally, optical technology is less expensive
than MR technology, with costs between $200,000
and $400,000.
In summary, NIRS is a useful developing tech-
nique that has some specific practical advantages
(but also limitations) with respect to fMRI. Its use,
especially in the applied field, is likely to grow sig-
nificantly in the next few years.
Event-related optical signal. EROS is another
application of diffusive optical imaging. The main
difference with respect to NIRS is the time scale:
instead of looking at phenomena occurring several
seconds after a particular area of the brain is acti-
vated, it focuses on phenomena that occur at the
same time as electrophysiological (postsynaptic)
events, on a millisecond time scale. These phe-
nomena are likely due to the swelling or shrinking
of dendritic trees, which have been demonstrated
to occur with depolarization and hyperpolariza-
tion of neuronal membranes (Rector et al., 2005).
EROS signals are smaller than NIRS signals, requir-
ing more advanced methods for their recording.
Although EROS has been obtained with intensity
measures (e.g., Franceschini & Boas, 2004; G.
Gratton, Fabiani, et al., 1995; Maclin, Low, Sable,
Fabiani, & Gratton, 2004; Medvedev, Kainerstorfer,
572
Borisov, Barbour, & VanMeter, 2008; Wolf et al.,
2002), most studies have used measures of photon
time of flight (i.e., the time taken by photons to
move between source and detector) obtained using
FD instruments (e.g., G. Gratton, 1997; G. Gratton
et al., 2006; G. Gratton, Corballis, et al., 1995; G.
Gratton & Fabiani, 2003; G. Gratton, Fabiani, et
al., 1995; G. Gratton, Fabiani, Goodman-Wood,
& DeSoto, 1998; G. Gratton, Goodman-Wood, &
Fabiani, 2001; G. Gratton, Sarno, Maclin, Corballis,
& Fabiani, 2000; Tse, Tien, & Penney, 2006; Wolf
et al., 2002; also see G. Gratton & Fabiani, 2009,
for a review), which have greater localization and
depth sensitivity. In addition, EROS measurement is
facilitated by high-density recording systems (e.g.,
G. Gratton, Rykhlevskaia, Wee, Leaver, & Fabiani,
2009; Low, Leaver, Kramer, Fabiani, & Gratton,
2009), appropriate frequency filtering (Maclin,
Gratton, & Fabiani, 2003), and extensive signal
averaging (e.g., G. Gratton & Fabiani, 2003; G.
Gratton et al., 2006). When all of these techniques
are employed, data with spatial resolution of up to 5
mm and temporal resolution around 10 ms to 20 ms
can be obtained (G. Gratton & Fabiani, 2003).
EROS has been used in a number of paradigms
and conditions, including visual (e.g., G. Gratton,
Corballis, et al., 1995), auditory (e.g., Rinne et al.,
1999), somatosensory (e.g., Maclin et al., 2004), and
motor modalities (e.g., DeSoto, Fabiani, Geary, &
Gratton, 2001). In each modality, simultaneous
electrophysiological recordings indicate that the
latency of the EROS activity is similar to the latency
of the main ERP components (e.g., DeSoto et al.,
2001; E. Gratton et al., 1997; Maclin et al., 2004;
Rinne et al., 1999). Furthermore, the locations of
the responses correspond to those obtained with
fMRI (e.g., E. Gratton et al., 1997; G. Gratton et al.,
2000; Toronov et al., 2005).
In summary, the main advantages of EROS are
good spatial (5–10 mm) and temporal (10–20 ms)
resolution; the possibility of simultaneous recording
with ERP, fMRI, and TMS (in addition to NIRS); its
relatively low cost (between $200,000 and $400,000);
and its adaptability to a number of experimental pop-
ulations and paradigms. Its major disadvantages are
low penetration and relatively low signal-to-noise
ratio (compared with ERPs, fMRI, and NIRS).

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Figure 29.2. Typical setup for a transcranial magnetic stimulation experiment.
Transcranial Magnetic Stimulation
TMS is a method for influencing the neural activity
in selected areas of the brain (for reviews, see
Pascual-Leone, Walsh, & Rothwell, 2000; Wagner,
Valero-Cabre, & Pascual-Leone, 2007). TMS is
based on the induction of weak electric currents in
cortical regions through strong magnetic fields (of
the order of 1 Tesla or more) produced by coils
located just outside the head (see Figure 29.2). It is
commonly thought that the weak electric currents
induce brief depolarization of small cortical regions,
whose extent depends on the type of coil and may
vary between 5 mm and 10 mm or so, followed by
prolonged periods of excitability or more often by
inhibition, akin to long-term potentiation and long-
term depression phenomena, respectively. In
essence, TMS can be used to produce temporary
lesions in specified cortical regions, which, because
There are a few side effects, for the most part limited to people with a history of epilepsy, which is therefore considered an exclusionary criterion for
4
participation.
Beyond ERP and fMRI
of their limited duration, can be compared within
the same subject with conditions in which the tem-
porary lesion does not occur. This makes TMS a
unique technique, in that it allows scientists to
manipulate directly the brain from the outside, and
therefore to make causal statements about the role
that a particular brain structure may have in infor-
mation processing or in a psychological task.
Two different TMS approaches are possible. The
first, called single- or double-pulse TMS (sTMS) is
based on the presentation of very short pulses of
current (a few milliseconds in duration), which gen-
erate relatively rapid effects in the cortex that
quickly wane when stimulation ceases. The advan-
tages of this method are the possibility of testing the
role of activity occurring at a particular time within
a specific brain region in the performance of a task
and its limited invasivity.4 Its main limitations are
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 Gratton and Fabiani
that the effects are weak, often only visible after
averaging over a large number of trials. A possible
confounding factor is that the sTMS pulse may gen-
erate stimulation not only through the induction of
currents in the brain but also through the induction
of muscle contractions in the scalp, and it may do so
through auditory stimulation related to deformation
of the TMS coil during the pulse. The latter con-
found can be controlled by running a control condi-
tion in which the magnetic field is redirected outside
the head. The possible role of muscle artifacts is typ-
ically controlled for by using sham stimulation of
other regions of the scalp, corresponding to cortical
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regions that are not considered relevant for the task.
The second method, called repetitive TMS
(rTMS), is based on the presentation of very long
(up to several minutes) low-frequency (up to 10 Hz)
pulse sequences. These sequences induce sustained
depression of the cortical regions involved, which
may last 30 min or longer. The paradigm typically
consists of comparing results obtained in two ses-
sions: an experimental one in which rTMS is applied
and a control one in which a sham rTMS is applied.
Because of the long duration of the effects, rTMS can
provide spatial information about whether a particu-
lar segment of the cortex is important for the perfor-
mance of a task but does not provide specific
information about the time at which this role is per-
formed. rTMS is more invasive than sTMS: In some
rare cases, seizures were reported even in subjects
with no previous history of epilepsy (unlike sTMS),
leading to the establishment of standards for its
appropriate use (Rossi, Hallett, Rossini, Pascual-
Leone, & the Safety of TMS Consensus Group,
2009). In addition, short duration headaches are
often reported as aftereffects of rTMS.
Notwithstanding these potential issues, TMS has
attracted the interest of many scientists because of its
unique potential for allowing causal inferences about
the role of different brain regions. That said, there
are a few important limitations of the technique.
Only regions that are close to the surface of the head
can be subjected to the procedure. The extent of the
cortical region manipulated is not well established,
and it is also possible that other regions may be
affected through the transmission of the signal by
neural tracts. sTMS is difficult to combine with other
574
brain imaging methods (but see Parks et al., in press,
for concurrent EROS and sTIMS recordings; Thut &
Pascual-Leone, 2010, for a review of combined TMS-
EEG recordings). This is less of a problem for rTMS.
Finally, the possible side effects associated with the
procedure (especially for rTMS) may discourage
some investigators from its adoption.
In summary, TMS is an intriguing technique for
manipulating brain function nearly noninvasively.
Its practical use is partly limited by safety and exper-
imental issues. The possibility of making causal
statements and the contained cost of the device
(around $50,000), however, have made its popular-
ity grow exponentially in the past few years.
Comparisons Among Measures of
Noninvasive Human Brain Function
To understand the relative utility of current measures
of human brain function, it is important to consider
that they vary along a number of dimensions and
provide different types of information. We can distin-
guish the following dimensions: (a) type of brain
function measured; (b) spatial and temporal resolu-
tion; (c) selectivity for specific regions, cell popula-
tions, or brain systems; (d) extent to which the
procedure is a measurement method versus a manip-
ulation tool; and (e) practical factors such as degree
of invasivity, flexibility, cost, availability, and
population(s) to whom it can be most readily
applied. Each of these dimensions plays a critical role
in determining the suitability of a specific technique
in investigating a particular psychological problem.
In this section we briefly discuss each dimension and
present a table (see Table 29.1) summarizing how
different methods can be classified accordingly.
Type of Brain Function Measured
Although brain activity if often regarded as a well-
defined concept, in reality it is composed of a com-
plex and multifaceted set of phenomena, which can
be summarized as follows:
1. Membrane phenomena: Depolarization and
hyperpolarization of neurons, associated with
ion exchanges across the neuronal membranes
(important for integrating information coming
 Beyond ERP and fMRI

Table 29.1

Dimensions of Human Brain Function

Spatial/
temporal
resolution Measurement vs.
Method Type of function (scales) Selectivity manipulation Population Other factors
EEG Membrane cm Postsynaptic activity Measurement Infants to older Least expensive and
ERPs ms Pyramidal neurons; adults; patient most portable
open fields populations technique; very
limited spatial
resolution
Copyright American Psychological Association. Not for further distribution.

fMRI Metabolic/ mm Postsynaptic activity Measurement Older children to Expensive; requires

hemodynamic s Glutamergic cells? adults; some large footprint
restrictions equipment
and special
environments
PET Metabolic/ cm Very selective Measurement Adults Moderately invasive;
SPECT hemodynamic; min depending on tracer very expensive
synaptic and method used
EROS Membrane sub-cm Postsynaptic Measurement Infants to older Limited penetration
ms activity; superficial adults; patient and low signal-to-
structures populations noise ratio
NIRS Metabolic/ cm Superficial structures Measurement Infants to older Limited penetration
hemodynamic s Glutamergic cells? adults; patient
populations
MEG Membrane sub-cm Postsynaptic activity Measurement Older children to Expensive; requires
ms Pyramidal neurons; adults large footprint
open fields; equipment
tangential and insulated
orientation (i.e., environment
sulci)
TMS Membrane? cm Largely unknown; Manipulation Adults Moderately invasive
sub-s superficial
structures

Note. The methods with gray background (EEG/ERPs and fMRI) represent the gold standards with which other methods are
compared. EEG = electroencephalogram; ERPs = event-related brain potentials; fMRI = functional magnetic resonance imag-
ing; PET = positron emission tomography; SPECT = single-photon emission computerized tomography; EROS = event-related-
optical signal; NIRS = near-infrared spectroscopy; MEG = magnetoencephalography; TMS = transcranial magnetic stimulation.

from different neurons and transmitting this
information over long distances).
2. Synaptic phenomena: Various biochemical events
that occur at the synapses between neurons
(critical for interchanging information between
neurons and for modulating neuronal function).
3. Metabolic and hemodynamic phenomena: A variety
of corollary homeostatic and trophic phenomena,
such as oxygen and glucose consumption,
vasodilation, increased blood flow, and so on,
which often involve other cells (such as glial
cells) in addition to neurons but intrinsically
relate to neuronal function.
Each of these phenomena results in biophysical
and biochemical changes, which can be monitored
with different brain imaging techniques. EEG,
ERPs, MEG, TMS, and EROS are sensitive to

575
 Gratton and Fabiani
membrane phenomena; NIRS and fMRI are sensitive
to hemodynamic effects; and PET and SPECT can
be rendered sensitive to a number of different phe-
nomena, both synaptic and metabolic or hemody-
namic, depending on the type of tracer used. All of
these phenomena are intrinsically related to each
other, albeit in a complex and typically nonlinear
fashion. Therefore results obtained with different
techniques are likely associated with each other
according to complex and sometimes not well-
understood rules.
Temporal and Spatial Resolution
Copyright American Psychological Association. Not for further distribution.
The different methods vary significantly in terms of
the temporal and spatial resolution of the informa-
tion they provide. In some cases, the temporal or
spatial specificity of the information is limited by
properties of the physiological phenomenon they
probe. For example, changes in the receptors for par-
ticular neurotransmitters, which can be in some
cases measured with PET and SPECT, may occur rel-
atively slowly, over the course of hours, days, or
even longer. The vasodilation response that follows
neuronal activation (also called blood-oxygen-level-
dependent [BOLD] response), measured by tech-
niques such as fMRI, 15O-PET, and NIRS, requires
the relaxation of smooth muscle fibers surrounding
the arterioles, which takes place over several
seconds.
Other limitations come from properties of the
measurement system. A basic limit for all noninva-
sive imaging methods is that the size, number, and
extracranial location of the detectors used makes it
impossible to target individual neurons (or individ-
ual synapses). Rather, all noninvasive human brain-
imaging methods refer to the summation of signals
from a large number (on the order of at least thou-
sands) of cells and neurons. In this sense, these
measures are logically different from the single- or
multiple-unit electrophysiological recordings
obtained intracranially in animals. The statistical
properties of the imaging signals have further con-
sequences: They bias the measurements toward
those systems that are most likely to generate sig-
nals that summate over space or time. This is true
even for procedures with relatively good spatial and
or temporal resolution. For instance, ERPs are more
576
likely to be the result of (dendritic) postsynaptic
activity (which extends over tenths to hundredths
of milliseconds) than to presynaptic (axonal) action
potentials (which only last for a few milliseconds)
because the probability of temporal summation of
the activity of different neurons is higher in the for-
mer than in the latter case. In addition, the larger
size (diameter and number of arborizations) of den-
dritic trees compared with axons makes the import
of the physical and chemical events associated with
postsynaptic activity overwhelmingly greater (and
therefore more likely to be detected at some dis-
tance) than that of the axons. Finally, different
techniques employ methodologies that may further
limit the spatial or temporal resolution of the
results. For instance, the temporal resolution of
methods relying on radioactive tracers, such as PET
and SPECT, may partly depend on the time-decay
function of the particular isotope used. Similarly,
the spatial resolution of measures based on electri-
cal potentials may be limited by the conductivity of
the tissue, which spreads electrical signals over a
large area.
Selectivity. Various techniques are more sensitive
to certain forms of brain activities than to others.
In some cases, this is intentional: SPECT and PET
methods that are used to investigate the location and
numerosity of specific receptors are intentionally
selective for them. In other cases, however, the bias,
or selectivity, is not necessarily intentional and may
provide important limitations to the types of data
that a technology can provide. For example, several
techniques (such as ERPs, MEG, optical methods,
and TMS) are more sensitive to superficial than to
deep events. Finally, biases may depend on other
factors, such as dependence on particular biochemi-
cal (as may be the case for hemodynamic-based
measures) or geometric (as is the case for ERPs and
MEG) factors.
Measurement versus manipulation. Most brain-
imaging methods passively observe changes in brain
activity; only TMS allows for its manipulation. Thus
most techniques only provide correlational data,
whereas TMS can be used in support of causal infer-
ences, indicating the extent to which activity in a

particular region is critical for the performance of a
given task.
Practical considerations. Several practical reasons
often lead to the selection of which procedures to
use to investigate a particular psychological phe-
nomenon. Some techniques may be less acceptable
than others because they may be more intrusive or
even invasive. Similarly, some techniques may be
unfeasible for some subject populations (e.g., infants
and young children). Some, such as fMRI, involve
large equipment and require participants to refrain
from moving. This may be incompatible with certain
Copyright American Psychological Association. Not for further distribution.
environments (e.g., field- or bedside applications)
or certain behavioral tasks (e.g., those requiring
extensive motion). Finally, other practical factors
may include cost and availability of a particular
technique.5
Combining Different Imaging
Modalities
Because different modalities offer data varying along
a number of dimensions, many researchers have
combined them with the intent of providing a more
complete view of brain function (Barinaga, 1997).
The approaches used can be classified into two
major classes: (a) a combination based on the com-
parison of completely analyzed data from each
modality (e.g., Logothetis, Pauls, Augath, Trinath,
& Oeltermann, 2001) and (b) a combination of
relatively unanalyzed data from each modality that
is fused based on physical or statistical principles
(e.g., Dale, 2000). Both of these approaches can be
based on data that are recorded separately (e.g.,
Huettel et al., 2004; Logothetis et al., 2001) or
simultaneously (e.g., Mackert et al., 2008; Toronov
et al., 2005). In practice, the simultaneous record-
ing of different modalities often creates compatibil-
ity issues. When this occurs, such as for the
simultaneous recording of EEG or ERP and fMRI
data, special methods need to be used to minimize
artifacts that originate in each modality from their
concurrent recordings. Because of the complexities
This issue has lead to the adoption of a shared-instrumentation model (which involves access fees) for equipment that is too costly or difficult to
5
maintain or operate to reside in a single lab. This is almost always the case for PET, SPECT, and fMRI, but it is more rarely the case for the other
methods reviewed in this chapter.
Beyond ERP and fMRI
involved in this process, concurrent recording
(although preferable in principle) is often limited to
cases in which it is absolutely necessary. In many
cases, superimposition of data collected at different
times at the moment of data analysis is sufficient.
Some imaging modalities (such as optical) generate
a small amount of mutual interference with others
modalities (such as EEG or ERPs, fMRI, and TMS)
and therefore appear particularly suited for concur-
rent recording and for providing a bridge across
techniques.
In fusing data from different imaging methods, it
is critical that the differences among the modalities
be taken into account. As such, the relative roles of
the various measures need to be conceptualized in a
theoretical model describing their mutual relation-
ship. The best understood integration issues relate
to the spatial and temporal superimposition between
measures. Spatial superimposition requires two
steps: (a) three-dimensional reconstruction of the
data in each modality and (b) coregistration of the
two data sets. Similarly, temporal superimposition
requires (a) modeling the temporal relation between
the two measures and (b) their relative time-locking.
In other words, the two sets of measures need to be
placed in the same space-time reference frame. Data
fusion also requires understanding how differences
in the underlying biophysical phenomena (as well as
possible biases) affect fused data. For example, if
one technique is particularly sensitive to cells placed
in open-field configurations (such as EEG/ERPs or
MEG) whereas the other is not (such as fMRI or
optical measures), we need to consider whether the
structures involved have open or closed configura-
tions in evaluating the relationships between the two
measures. Similarly, differential biases toward super-
ficial versus deep structures need to be taken into
account when comparing NIRS data with MRI data.
In summary, data fusion requires careful consid-
eration of the various characteristics of the imaging
modalities involved. This may require a series
of studies comparing the measures. In the past
15 years, a number of studies of this type have been
conducted (e.g., G. Gratton et al., 2001; Huettel
577
 Gratton and Fabiani
et al., 2004; Logothetis et al., 2001), and presumably
several more will be conducted in the future. Hope-
fully these studies will pave the way for a new
approach describing brain activity as a complex,
multifaceted phenomenon involving membrane,
synaptic, and hemodynamic effects. The integration
of anatomical information within this description
may further enrich our knowledge of brain and psy-
chological function.
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