Psychological Research (2012) 76:131144
DOI 10.1007/s00426-011-0408-6
REVIEW
Some unsettled problems in behavioral neuroscience research
Frank Rosler
Received: 2 November 2010 / Accepted: 21 December 2011 / Published online: 10 January 2012
 Springer-Verlag 2012
Abstract The goal of behavioral neuroscience is to map
psychological concepts onto physiological and anatomical
concepts and vice versa. The present paper reflects on some
of the hidden obstacles that have to be overcome in order to
find unique psychophysiological relationships. These are,
among others: (1) the different status of concepts which are
defined in the two domains (ontological subjectivity in
psychology and ontological objectivity in physiology); (2)
the distinct hierarchical levels to which concepts from the
two domains may belong; (3) ambiguity of concepts,
becausedue to limited measurement resolution or defi-
nitional shortcomingsthey sometimes do not cover unique
states or processes; (4) ignored context dependencies.
Moreover, it is argued that due to the gigantic number of
states and state changes, which are possible in a nervous
system, it seems unlikely that neuroscience can provide
exact causal explanations and predictions of behavior.
Rather, as in statistical thermodynamics the transition from
the microlevel of explanations to the macrolevel is only
possible with probabilistic uncertainty.
Introduction
Behavioral neuroscience intends to understand the biolog-
ical bases of behavior and subjective experience. The credo
is to eventually explain how such complex phenomena as
understanding language, perceiving objects, driving an
automobile or deciding on investments as well as subjective
experiences such as fear, happiness or insight do arise from
F. Rosler (&)
Department Psychology, University of Potsdam, Karl-
Liebknecht-Str. 24/25, 14476 Potsdam OT Golm, Germany
e-mail: froesler@uni-potsdam.de
the mere interaction of simple building blocksneurons
that interact by means of excitation and inhibition and
whose connective strengths can change due to learning.
During the last 70 years1 a vast number of findings on the
biological bases of behavior have been accumulated. These
provide many insights into how the neural machinery
works and which electrical and chemical processes enable
perception, movement, memory or language. Nevertheless,
these empirical findings and theoretical accounts are lim-
ited in scope. Judged by the richness of real-life phenom-
ena or by what psychology is actually interested in, the
material summarized in textbooks on cognitive or behav-
ioral neuroscience scratches the surface only. For the time
being we, as neuroscientists, are far away from providing a
clear-cut, biologically grounded explanation for complex
behaviorhow a novel is written, an airplane constructed
or why an individual becomes a genius or a terrorist. And
most likely this state of affairs will not change soon.
Neuroscience does provide many basic and domain-spe-
cific explanations, but it is still far away from providing
exact predictions of individual behavior or, even more
ambitious, explanations of the interaction of mind and
1
The term Cognitive Neuroscience was introduced around 1980 by
Michael Gazzaniga and George Miller to define an area of research
that has its focus on psychologically oriented neuroscience [in
contrast to the more (general) physiologically oriented neuroscience
program]. However, the topics of cognitive or better Behavioral
Neuroscience have a much longer history. Actually it was Wilhelm
Wundt who laid the corner stone with his Grundzuge der physiolog-
ischen Psychologie (Wundt, 1874). Before the term cognitive
neuroscience had been coined the topics were covered under the
labels Biological Psychology, Physiological Psychology, Biopsychol-
ogy, and Psychophysiology (e.g., early textbooks by Rosenzweig &
Leiman, 1982; Watson, 1981; Pinel, 1992, or journals as The Journal
of Comparative and Physiological Psychology whose first issue
appeared 1921, Behavioral Neuroscience (1983), and Psychophysiology
(1964).
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body. In this paper, I will briefly discuss some problems
which might help to understand why in behavioral neuro-
science explanations and insights are so difficult to
achievewhy this research, which tries to relate mind and
body, is much more difficult to pursue than most other
scientific enterprises. In so doing I will outline some
epistemological obstacles and I will discuss the question
whether it is possible at all to forecast individual behavior
by means of biological measures.
Mind and brain: distinct or identical?
In its essence the question on the relation of mind and brain
reduces to a mapping problem. How can concepts and
relations of the world of psychology, e.g., percepts, deci-
sions, reasons, attitudes, emotions, language scripts and
utterances, be translated into concepts and relations of the
world of biology or physiology, i.e., into states of activa-
tion and inhibition in a neural network or into the expres-
sion of transmitters at several brain sites?
Some philosophers and psychologists argue that it is
useless and irrelevant to consider neurobiology at all in
order to understand psychological phenomena and to pre-
dict behavior. Categories and concepts of both worlds are
seen as fundamentally distinct and, therefore, it is assumed
that it does not make much sense to search for mapping
relations between the two. In final consequence, however,
such a position can only be defended if one accepts an
ontological difference between mind and body, i.e., if one
assumes that psychological and physiological phenomena
are distinct and can exist independently from each other. A
body can exist without a mind and vice versa. While it
seems to be acceptable to subscribe to the first statement
a body can exist without a mindit seems more than
difficult to subscribe to the second: a mind can exist
without a body. The evidence in support of psychological
phenomena depending on the material basis of a nervous
system is overwhelming and it seems implausible to deny
this. Even hard-core idealists will not be free of doubts on
this issue unless they obey deep-rooted religiously moti-
vated beliefs. Most scientists adhere to a monistic position
and this also seems to be the more or less explicit creed of
common sense. It is accepted that the brain is the basis for
the emergence and development of what is addressed as
psychological phenomena. Mind and body are seen as an
ontological unity. This unity has distinct manifestations
and, accordingly, it can be described and investigated from
different perspectives: on the one hand by means of sub-
jective and objective psychological concepts and methods,
on the other hand by means of physiological and ana-
tomical, more generally spoken, by means of physical
concepts and methods. At bottom, one accepts ontological
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Psychological Research (2012) 76:131144
monism but allows for methodological and theoretical
dualism.
Given this basic assumption there must exist, at least on
principle, one-to-one mappings between psychological and
physiological phenomena and concepts. A thought, a
remembered event, a percept, or, in general, any psychic
state and process must have a physiological counterpart.
For each psychic state it must be possible to identify a
pattern of activated and inhibited neurons, as it should also
be possible to find a one-to-one psychological correlate of
any clearly defined physiological state or state change.
Equivalence on principle does not mean, however, that
such a mapping is actually possible with the available
concepts and tools. Mutual mapping of psychological and
physiological entities might be possible only to some
degree, i.e., with a more or less large amount of unex-
plained variance. I will give some examples below.
Concepts, levels and hierarchies
In order to do research at all it is necessary to categorize
the universe of discourse. Categorizations are provided by
our percepts and actions and categorizations seem to be the
result of some very basic properties of nervous systems as
such. Already most simple organisms categorize stimuli
and events, and neural networks can be efficiently descri-
bed as filters that categorize external and internal events
(van Essen, Anderson, & Felleman, 1992).
In the physiologicalphysical domain concepts are
comparatively transparent and clearly distinct. By and
large they are defined by means of their spatial (i.e., ana-
tomical) and temporal features. These entities form a
hierarchy according to size and complexity (Churchland &
Sejnowski, 1989). The peak of the anatomical hierarchy
starts with the all-encompassing category of the whole
organism, which is then differentiated into subsequently
smaller and more specific entitiesthe brain, the hemi-
spheres, areas of the cerebral cortex, nuclei of the brain
within these structuresnetworks (e.g., columns within the
cortex), neurons, synapsesand within these subsynaptic
entitiesion channels or single transmitter molecules.
Measures can comprise the whole lifespan (developmental
changes of the nervous system), days or hours (synaptic
pruning), seconds (metabolic changes), or milliseconds
(action potentials). As first mentioned by Churchland &
Sejnowski, 1989) each neuroscientific measurement oper-
ation and conceptual entity can be located somewhere in a
two-dimensional grid with two axesone is the anatomical
size of the measured entity, extending from nanometer to
meter, and the other the temporal extension of the mea-
sures, extending from microseconds to years. All these
entities are easily defined by an objective measurement
operation.
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With respect to functional variables that can be derived 2. Psychological concepts are not as objective and
from these measures the hierarchy is closely related to the independent from the observer as it is the case for
anatomicaltemporal grid hierarchy: metabolism of the biologicalphysical concepts. Rather, psychological
whole body, blood flow changes of brain regions, electrical concepts do always have introspective and social
activity of neural networks, single cell action and resting connotations, they areas outlined by Searle, 1987)
potentials, transmitter release and reuptake, etc. By means ontologically subjective.
of the available tools and knowledge about neuroana-
tomical, physiological and molecular mechanisms it has
Boundaries of psychological concepts
become possible to delineate many transitions between
distinct levels of this hierarchy, e.g., how the blood oxy-
Experimental psychology puts humans and animals into
gen level-dependent signal (BOLD) emerges from the
specified situations in order to evoke a certain type of
regulation of the capillary system and metabolic changes
behavior that varies in quality and intensity. How are the
(Magistretti & Pellerin, 2000), or how the scalp recorded
boundaries of an experimental situation being defined?
voltage changes in the electroencephalogram originate
Where does the situation start and end? And how is one
from electrical changes in underlying neuron populations
type of behavior defined and segregated from other types of
(Speckmann & Caspers, 1979; Nunez, 1981).
behavior? First of all and most obviously a situation is
Within the domain of psychological conceptsbehavior
defined by a set of physically given conditions: a lab room,
and subjective experiencea hierarchy can also be dis-
a computer monitor, some stimuli, e.g., words presented
cerned, however, the definition of distinct levels and of
sequentially on the screen, and a response key. But this is a
clearly distinct entities is less easily grasped (Rosler, 1983,
superficial description only. Much more important than
1984). It is only in some cases that the hierarchy becomes
these physical ingredients is the instruction. Depending on
obvious, while in most others it is difficult to specify. With
the instruction an identical physical experimental setup can
respect to language, for example, it is possible to distinguish
result in completely different psychological situations.2
between entities of different extensions: graphemes, mor-
The instruction can be, e.g., (a) Categorize the words
phemes, words, phrases, sentences, and text, and it is also
according to the distinction animate/inanimate. (b) Cat-
possible to order these elements in a hierarchy. Moreover,
egorize the words according to the distinction animate/
there is evidence that the elements of such a linguistic
inanimate and remember all words for a later recognition
hierarchy are not only theoretically definable but that they
test. (c) Categorize the words according to the distinc-
also have an empirical underpinning which becomes man-
tion animate/inanimate and remember all words for a later
ifest in behavioral effects (e.g., priming and interference
recognition test. For each correctly categorized word you
effects between elements within and between the distinct
will receive 50 cents, for each incorrectly categorized word
hierarchical levels; see Levelt, 1989; Dell, 1986).
you will loose 2 Euros. or (d) Categorize the words
A similar hierarchy might be discerned in the domain of
according to the distinction animate/inanimate and
visual perception, as there are visual features (color, orien-
remember all words for a later recognition test. We want to
tation, movement,), combinations of these features
investigate whether it is possible with this procedure to
(=objects), and combinations of objects (=scenes) (Treis-
measure students intelligence.
man, 2004). Much less transparent are hierarchies within
In each example the very same physical situation is
other psychological domains, e.g., emotions (is there a
embedded in a completely distinct psychological context.
hierarchy of emotional elements?) or social interactions
In (a) the focus is on psycholinguistic variables, e.g.,
(what are the elements of a conversation, the elements of a
whether word frequency affects decision time. In (b) the
promise, an instruction?). As a matter of fact concepts of
focus is on memory and the lexical decision instruction
psychology are most often far less precisely defined than
might only be used to distract attention from the experi-
concepts of the physiological-anatomical domain. This
mental question proper, viz., whether animate words are
ambiguity of psychological concepts has at least two causes:
better remembered than inanimate words. In (c) a moti-
1. The boundaries of psychological concepts are not vational aspect is added and the key question might be
defined by means of immediately perceivable measure- whether decision times and memory recall depend on
ments. There is neitheras in neuroanatomya simple monetary incentives, losses and gains during learning. In
measurement procedure which comprises always the (d), finally, a social context is defined and the question
same scale of measurement and by means of which might be whether decision times are affected, if an
concepts are easily defined, nor is there a clear temporal
segregation of states, events and processes which forms
2
the basis of psychological categorizations. In animal research the intention of the experimenter is induced on
the animal by means of an instructing learning procedure.

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imaginary peer group is introduced that will induce a
competitive attitude.
Where are, in these examples, the conceptual and tem-
poral boundaries of the experimental situation? The
instruction will be given at the beginning of the experi-
ment, it defines the superordinate context. Each monetary
gain or loss that is presented before or after an item, defines
a temporally shorter context, which, however, is not clearly
defined. How long will the aftereffect of an experienced
loss last? Will it affect only the immediate item, also the
next or even the next five items? And to what extent will be
a previous failureexperienced the day beforebe rele-
vant in context (d)?
The experimental situation and the set of variables that
may determine behavior is not easily and exhaustively
defined. Likewise, behavior as suchthe operation of a
response keyis meaningless. Its psychological meaning
does become transparent only, if the experimental context
is considered. Thus, the same key press and latency change
can mean many things.
In the daily routine of experimentation this ambiguity is
usually ignored. It is assumed that the ambiguity is elimi-
nated by averaging across different participants and across
replications of the same situation within the same partici-
pant. All indeterminacies of a single situationbehavior
combination are added to the error variance. This is a
feasible procedure in order to grasp statistical covariations
between general aspects of a situation and general aspects
of related behavior. It has to be kept in mind, however, that
this approach does not capture how an individual data of
behavior depends on a uniquely defined situation. The
approach provides nomothetic but not ideographic
statements.
Ontological subjectivity
As outlined in more detail by Searle (2000) there exists
another difference between physical and psychological
social concepts. On the one hand there are biological
physical measures and derived variables that can be
measured by clear-cut operations. Such measures, variables
and derived concepts are ontologically objective. These are
brute facts, which are almost completely independent
from the observer and which can be observed and
measured always in the same mannertoday, yesterday,
and tomorrow, in Asia, Europe, or Africa. In contrast,
psychologicalsocial concepts, variables, and measures
depend on the observer and his or her assumptions. These
concepts are ontologically subjective, as they do not exist
in an absolute sense by means of their spatialtemporal
reality, but only in a relative sense. They rest on intro-
spective experience and social agreement. Due to this they
cannot be fully grasped by objective measurements. In
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Psychological Research (2012) 76:131144
many cases these concepts exist only due to a collective
intentionality, this means, they are not brute facts, but
result from mutual understanding. Convincing examples
are, among others, money, contracts, promises, guilt,
remorse, etc. The true nature of money is not revealed by a
coin or a bill as such, it discloses itself only because of the
knowledge that people have who use money: it means
property, can be exchanged for goods, etc. The same holds
for psychological concepts as fear or pleasure, and for
psychologically defined experimental conditions. We do
understand these concepts only, because we have acquired
introspective experiences that are labeled accordingly. And
ontological subjectivity applies also to each and every
semantic meaning of a single word. A noun has a clearly
defined denotative meaning which most often is related to
perceptual and physically definable features (Barsalou,
2008), but each user of a word experiences additional
connotative shades of meaning which go far beyond the
basic and physically grounded meaning. Reading the word
cow may trigger associations of milk, a dairy, and
summer vacations in one person, but India and holiness in
another. In summary, even if we try to define an experi-
mental situation as carefully as possible and even if we
make a great effort with respect to how an instruction is
phrased, we cannot avoid that our psychological concepts,
as understood by the participant and by the experimenter, are
subject to idiosyncratic connotations and social under-
standings. This means that physical categories of physiology
and anatomy and psychological categories derived from
behavior and subjective experience are in some core aspects
distinct which will make it difficult to formulate one-to-one
relationships.
Relations and mapping problems
But let us ignore the problem of definitional ambiguities for
a moment and let us assume that there is agreement about
how a specific psychological situation has to be defined and
how the elicited behavior has to be interpreted. Even then a
fundamental problem remains, viz., which concept of the
one side of the mindbrain equation matches with which
concept on the other.
It is conceivable that precisely defined entities in one of
the two domains do not have equivalently circumscribed
counterparts in the other. For example, it is not possible to
map a sensation onto a single neuron or a set of neurons
right away, even if this sensation is temporally very cir-
cumscribede.g., a shock experienced in response to a
frightening, unexpected event. The shock does not corre-
spond with the activity of a single neuron only, but rather
with a specific pattern of activations that comprises mil-
lions if not billions of elementary activity changes within
the brain and the whole body. And this holds for all the
Psychological Research (2012) 76:131144
other psychologically circumscribed entities as wella
memory trace, a desire, a conflict, a movement, a word that
we read or hear. The mapping operation is not as simple as
translating from one language into another in which com-
parable entities can be defined on both sideswords,
phrases and propositions. Mappings between mind and
brain comprise much more complex relationships, i.e., a
psychic state, experienced or observed as a unitary event
has to be related to state or state change within a high-
dimensional dynamic system. Schematically such an
asymmetric mapping is depicted in Fig. 1 by means of the
continuous double-headed arrow.
To find a relation between a unitary concept on one side
and a dynamic process on the other is a hurdle not
impossible to overcome. It has to be seen as a problem and
the still often made error has to be avoided that a psy-
chological concept or process can be related to a narrowly
circumscribed anatomical location. It is still an often
implicit assumption that such simple localizations are
possible, e.g., relations between visual perception and
visual areas in the occipital cortex, or between parsing adjust concepts and categories. It could be that some
of syntax and Brocas area. In the light of more recent
findings such localization attempts seem to be nave. Visual
perception is not defined by the mere fact that filters are
Fig. 1 Mapping relations between elements of distinct hierarchical
levels of psychological (left) and biological (physiological, anatom-
ical) concepts (right). Shaded areas indicate concepts (states and
processes) which are distinct by definition, i.e., on the side of
psychology sensations, memory representations, etc., and on the side
of biology activations of neurons and neural networks. Long dashed
lines indicate mappings between unique concepts of biology and
ambiguous concepts of psychology (example, states of remembering
have distinct biological underpinnings but cannot be separated by
135
found in the occipital cortex being selectively sensitive to
color, orientation or movement. Such filters are a pre-
requisite for visual perception, but perception as experi-
enced or objectively measured takes place only, because
these filters interact with neural networks in prefrontal and
temporal cortex, structures which are involved in directing
attention and in combining bottom-up information pro-
vided by the receptors with top-town information provided
by memory (see Haynes, 2009; Quiroga, Kreiman, Koch, &
Fried, 2008). A comparably complex interaction of dis-
tributed networks has to be assumed for any other psychic
processbe it cognition or emotion.
Although it seems not to be impossible to relate dis-
tinctly defined entities with each otherpsychological
states and dynamic neuronal processesanother obstacle
has to be considered. On a priori grounds it is not known
which concept defined on one side does match with which
concept on the other. It is an empirical task to delineate
such optimal matches and the criterion for the goodness of
fit is mutually explained variance. It might be necessary to
concepts are not exact enough in order to relate them to
concepts of the other side. In such cases the mapping will
be ambiguous and mutually explained variance will be
means of behavioral indices). Short dashed lines indicate mappings
between unique concepts of psychology and ambiguous concepts of
biology (example, emotions may be distinguished by experience but,
due to a limited resolution of biological measures, they may not be
distinguishable on the physiological level). Continuous lines indicate
a unique mapping between a distinct psychological concepts (e.g., a
word meaning) and a uniquely defined distributed activation pattern
which extends over different hierarchical levels
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small. This problem will become more transparent when
some examples are specified.
Biological equivocation of apparently unique
psychological concepts
Psychology has a long tradition in distinguishing between
long-term and short-term memory (James, 1890) or, better,
between permanent and working memory (Baddeley,
Eysenck, & Anderson, 2009). It is the same word memory
that is used for clearly distinct phenomena. Common for
the concept of memory is the fact that something is not
perceptually present for some time but that it can be
recalled or recognized at a later point in time. The common
feature of the concept memory is the idea of storage and
later retrieval. Distinct is the time span that passes between
encoding and retrieval. However, does this apparent simi-
larity mean that the biological correlates of long- and short-
term memory are similar or almost identical and that it is
only the temporal aspect that makes memories distinct?
As a matter of fact this seems not to be the case. The
psychological concept memory has completely distinct
biological bases whether it concerns permanently or only
shortly stored information. What is addressed as long-term
memory has its biological basis in changed synaptic con-
nections in neocortex and other brain areas. These changes
are established in a long process of consolidation by means
of repeated activation of the involved brain systems.
Working memory, on the other hand, results from the
activation of a subset of these neural networks that house
permanent engrams. Moreover, this activation can be reg-
ulated by means of two distinct neural networks compris-
ing partially overlapping structures.
If a participant is instructed to remember a list of words,
such as camel, zebra, glove, scarf, rabbit and sheep,
these words are well familiar to the participant, i.e., the
words as such have not to be learned by the participant.
What has to be learned is an episodic tag that relates these
words to a particular situationthis experiment in this psychophysiological statements must be iridescent. The
room run by the attractive student. Experimental evidence
suggests that such episodic tagging results from an inter-
action between prefrontal structures and structures of
posterior cortex (see Fig. 2a). The posterior areas will be
involved in other activities in a delay period but the pre-
frontal areas maintain a kind of address code which enables
the system to reactivate the representations in posterior and
temporal cortex at a later point in time (Goldman-Rakic,
OScalaidhe, & Chafee, (2000); Druzgal & DEsposito,
2003).
However, after strolling through an unknown city and
reconsidering over a cup of tea what one has just seen, only
few already existing representations will have been episod-
ically tagged. In this situation primarily new associations
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Psychological Research (2012) 76:131144
will have been established and later reactivated. Such new
associations are initially stored in hippocampus and the
adjoining brain areas of the temporal lobe. Immediate rec-
ollection of these new impressions implies an interaction
between structures of the medial temporal lobe and filter
networks of the posterior parietal and occipital cortex
(Fig. 2b), as well as interactions between prefrontal, tem-
poral and posterior networks (Fig. 2c). All three scenarios
are correlates of storage and retrieval of memory contents,
but the biological bases are different.
This problem is not only notorious for constructs from
cognitive psychology. So far there is no evidence that
psychological or psychiatric disease categoriesdepres-
sion, schizophrenia or anxiety disorderhave immediate
biological counterparts, i.e., that there is one and only just
one biological cause in each case. Only a brief glance at
publications on this issue reveals that this must not be the
case. Rather, each syndrome which becomes manifest
maybe caused by a number of distinct biological deviations
from normality. Thinking does not take place at a narrowly
specified location in the brain. Thinking needs the whole
brain with all its diversified networks, and, accordingly,
disorders of thinking in schizophrenia can have many
causesan imbalance of transmitter substance in area x,
faulty connections between areas y and z, an overactivation
of area w, etc.
Even a meticulous categorization of symptoms accord-
ing to DSM (American Psychiatric Association, 2000) or
ICD-10 (DIMDI, 2010) will not be of much help, because
this diagnostic delineation of symptoms is subject to the
above-mentioned ambiguities and social agreements, and
each diagnostic category can have distinct biological bases.
Recent publications on the multicausality of autism provide
an illustrative example (Pinto et al., 2010).
The described situation shows that a psychological cat-
egory at issue is only apparently unambiguous or unique.
Considering the biological bases the concept is ambiguous
(see Fig. 1, the dashed connections). Given such a scenario
psychological ambiguity can be resolved, if the psycho-
logical categories are differentiated according to the bio-
logical distinctions, i.e., if subcategories are formed.
Psychological equivocation of apparently unique
biological concepts
Ambiguities into the other direction are also possible. It is
conceivable that a biological concept seen as a unity can be
related to different psychological concepts (in Fig. 1 this is
symbolized by means of the dotted lines on the second
level from bottom). For example, we can easily decide
whether we feel happy or sad and likewise whether another
person we are communicating with is more likely in the
Psychological Research (2012) 76:131144
Fig. 2 Hypothetical biological correlates of distinct forms of mem-
ory retrieval. a Episodic representations in a working memory task;
b Retrieval of new associations, which are transiently stored in the
medial temporal lobe by means of an address code; c retrieval of new
one or the other state of mood. Recording some peripheral
physiological measure reveals, however, that there seems
not much of a difference between the two states of sadness
or happiness, or of anger and anxiety. These peripheral
physiological patterns do not reliably differentiate between
different states of mood or emotion, just the general level
of arousal is reliably indicated by them. Only if additional
variables are considered, e.g., biochemical indicators or
brain activation states, emotional states maybe mapped more
precisely on biological states and state changes (Stemmler,
1989).
The last example reveals that it is not only a question of
finding the appropriate categories to be related with each
other, but also a question of the resolution and comprehen-
siveness of measurements that are considered to define a
particular concept. And this holds for both sides of the mind
brain equation. Progress in science increases the number and
the resolution of measurements and by means of this the
resolution of and distinctness of concepts will also change.
Inevitably this will result in a continuous change and dif-
ferentiation of psychophysiological statements. Progress on
this issue is an empirical affair. Different mappings have to
be tested until a maximum of covariance between mea-
surements and concepts will be achieved.
Translation does not mean substitution
and reductionism
The claim The total is different than the sum of the parts replacement of concepts of a higher level by means of con-
has a long tradition and can be traced back to Aristotle and
other ancient philosophers. But in particular modern brain
science gave substance to this claim. As already outlined it
is a characteristic feature of neuroscience that observations
and theoretical statements are made within distinct con-
ceptual domains and within each domain on distinct hier-
archical levels. And, as mentioned, translations between
these domains and hierarchical levels are difficult. But
assuming, just for the sake of argument, that one-to-one
137
associations which are also tagged as contents of working memory
(PFC prefrontal cortex, MTL medial temporal lobe, sens. and mot c.
sensory and motor cortices)
relationships between psychological and biological con-
cepts can be found, it remains a question whether psy-
chological concepts can be dismissed by simply referring
to the biological concepts, whether psychology can be
reduced to physiology. There are two counterarguments,
one is epistemological, the other one pragmatic.
Emergent properties
One can ask whether full reductionism is possible within
one domain alone (Wimsatt, 1976), e.g., between the dif-
ferent functional levels (and measurements) of brain
activitye.g., between single neurons and ensembles of
neurons. Obviously this seems not to be possible. The
properties of single neurons are well understood and can be
comprehensively described by mathematical formulas. In
short, a neuron is a nonlinear signal converter, which
integrates activation states on the input side in order to
generate a new signal on the output side, given a certain
activation level is surpassed. If many such nonlinear acting
elements are combined, as it is the case in neural networks,
the system states and state changes cannot be easily pre-
dicted by means of a simple set of mathematical equations,
i.e., the system built from such elements has its own
dynamics and is not fully predictable from the behavior of a
single element, the system has emergent or synergetic
properties (Basar, Flohr, Haken, & Mandell, 1983; Mitchell,
2009). Thus, complete reductionism in the sense of a full
cepts of a lower level is not possible, even within one domain
alone. Consequently, it seems to be more than questionable
to replace the concepts of one domain by those of another
distinct domain. Complete reduction of psychology on
biology would mean that there exist unequivocal one-to-one
relationships. Considering the limitations outlined above it
seems more than unlikely that such a complete translation
can ever be achieved, at least not with the now available
methods and tools.
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Since complete reductionism is an unrealistic goal,
neuroscience pursues a less ambitious program. It is
accepted that certain phenomena can be described, inves-
tigated, and theoretically grasped by means of different
methods (see above, methodological dualism). Such
observations are seen as complementary and the ultimate
goal is not that of conceptual replacement but of finding
correspondence between concepts, i.e., mapping and
translation rather that substitution (Fahrenberg, 1979).
Necessity of psychological concepts
As it will be explained in more detail below it might be
possible to describe certain high level processes by means
of a combination of low-level processes. But such
descriptions are of principle nature and not exhaustive.
Quite often it is already the mere number of microstates
that constitute one macrostate which prevents a complete
mapping. This is the same principle as discovered and
described by Ludwig Boltzmann more than one hundred
years ago for the domain of statistical thermodynamics.
The movement of a single molecule of water in the air
maybe traced with appropriate methods; however, the
tracing of all molecules in a cloud or the prediction of the
movement of a cloud, will only be possible by means of a
statistical statement. This means that an exact causal
statement is possible only on the level of the microstate
the trajectory of a single molecule colliding with other
molecules. This is a principle statement about how the
system works. But on the macrolevel such precise state-
ments are no longer possible, because of the limitation of
measurements and exhaustive description. And as it is a
necessity to use concepts of macro-physics to describe and
predict the world around use.g., to describe the function
of a combustion machine or to predict weatherit is
necessary to use (macro) psychological concepts in order to
describe and predict behavior of individuals and their
interactions.
Context dependencies
Above I have already alluded to the fact that psychological
concepts and variables are context dependent. They are
dependent on introspective and social reference systems,
or, in the words of Searle (1987), they are ontologically
subjective. One of several reasons why this context
dependency exists, and why such concepts are often not
fully explained by the immediately observable facts, is the
limitation of the time series of events that we normally
consider in an experiment (see Fig. 3). In an experiment we
must restrict the considered time series that determines the
behavior of a participant. We focus on one or two time
steps only, but the behavior shown at time t0 in its full
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Psychological Research (2012) 76:131144
uniqueness is a function of an extremely long time series of
conditional events that started somewhere in the past. The
encountered state changes that led to the behavior at time t0
have also determined how the wetware, the synaptic con-
nectivity was shaped, in other words it has determined the
memory of our participant. The time series of events starts
with birth (more precisely with conception) and each state
change will always be a consequence of the preceding state
changes. This results in a long chain of conditional proba-
bilities. In essence, the ontogenetic development of an
individual is subject to the mechanism of evolution. Devel-
opment comprises an accumulation of minimal, conditional
changes over a long stretch of time. On the short time scale of
a human life these changes bring about individuality of a
single person.
An example presented by Gromann (2007) illustrates
nicely how this mechanism of an accumulation of minimal
changes can bring about completely different final and
intermediate states (see Fig. 4). Let us assume that the
concentration of a transmitter at one synapse varies
between 0 and 1 and the state at time t ? 1 being defined
as xt?1 = 2 9 xt modulo 1. This means, the state variable
xt is multiplied by 2. If the result is smaller than one, the
result will be retained, if it is larger than one, only the
decimal rest will be retained. In Fig. 4 the function xt starts
with x0 = 0.2783, the function yt with 0.2784, i.e., the two
starting values differ in the fourth decimal. This is a min-
imal difference that should not matter much, but as it can
be seen, the two functions diverge after some time and two
complete distinct trajectories result. The difference is only
the consequence of a minimal difference at the start. And
this is not a question of the number of decimals considered
at all. The same effect can be seen if the two starting values
differ in the 9th or 10th decimal. Moreover, these distinct
trajectories develop free from any additional external
intervention. It is easy to conceive that such minimal dif-
ferences are amplified if external impulses are added at
some point in time.
A time series of conditional events and small accumu-
lated changes do not only exist for an individual. An
equivalent time series determines the environment and
which stimulus will be present at a particular time. Both
event sequences together determine the uniqueness of a
psychological state or state change of a person. Develop-
ment is always the result of a co-construction of environ-
mental and biological determinants (Baltes, Reuter-Lorenz,
& Rosler, 2006). And likewise the behavior and subjective
experience at a specific point in time is the consequence of
an interaction of these event series.
In order to capture behavior and subjective experience
in its full determination, it would be necessary to know
the complete time series relevant for a person. As this will
not be possible some of the behavioral variance must
Psychological Research (2012) 76:131144
Fig. 3 Outline of the contextual dependencies of situations and
persons and the temporal restriction of an experiment on only few
time steps or event transitions. The upper trajectory with white points
depicts the state changes of a person which are, seen from a
physiological point of view, engraved in the wetware of the nervous
system. The lower trajectory with gray points depicts state changes of
necessarily remain unexplained. This truncation of the
temporal perspective, the restriction on a few time steps
only does not only apply to psychological research. It is
likewise relevant for the biological side, the wetware and
all biological measures.
Dependency of behavior and experience
from previous context
Many examples show that current behavior can be better
predicted if not only the immediately preceding stimulus is
considered, but also stimuli and behavior that occurred
further in the past. Among others this is clearly shown by
experiments with task-set switches or backward inhibition
(Monsell, 2003). These reveal that behavior is a function of
a longer series of preceding trials. Such influences can span
two or three previous stimulusbehavior combinations and
more. In mental calculation, e.g., it has been shown that a
produced result can influence behavior until up to ten trials
later (Campbell, 1991), because results of the same mul-
tiplication table interact with each other. Assume the first
item reads 3 9 7 and the participant produces the correct
result 21. If now other items with multiplication factors 3
or 7 will follow, e.g., 3 9 4 or 2 9 7, it is more likely that
21 is produced as an error than other erroneous results. The
correctly produced number 21 increases the probability that
if an error occurs the same number will be produced again.
Expressed on a time scale such aftereffects can last up to
several minutes. It is a consequence of prevailing activa-
tions in memory.
Of course, it is not only a laboratory situation that dem-
onstrates long-lasting effects of previous stimulusbehavior
combinations. Some introspection shows immediately that a
139
the environment. Behavior is always a function of the states of both
trajectories (p and s). This interactive dependency is symbolized by
the arrows pointing from s and p onto behavior (the black points in
the center trajectory). Behavior changes the environment but also the
wetware of the behaving organism. Normally only a short historic
perspective is consideredlight or dark gray fields
Fig. 4 Illustration how small changes are accumulated and how this
can result in completely distinct trajectories. The two trajectories
differ in the fourth decimal of the initial value
certain event that occurred lately has a consequence on our
current behavior. In a meeting we heard something about
person x. We do not know x, but a week later we see the name
of this person listed as author of a paper in our favorite
journal. Now, this previously encountered information about
x will partially determine our behavior, whether we start
reading the paper or not. It is interesting that in laymens as
well as in experts explanations of extremely deviant
behaviorwhy someone became a terrorist or a shy, sub-
missive personvery often influences are cited that lie far
back in the past. Such references might be wrong, most often
they cannot be validated anyway, but the fact as such shows
that people are aware of their historic determination. Ironi-
cally in the lab of experimental psychologists and neuro-
scientists this historic dimension is often forgotten and it is
assumed that these factors, relevant as they may be, will
average out.
123
140
Dependency of biological signals from previous context
Whether such historic dependencies can indeed be ignored
as it is the case in daily laboratory routine has to be
questioned. Of course, it is possible to abstract from the
individual and to find general laws that relate input to
output or input to internal state changes. A question how-
ever is, how much variance will remain unexplained, if the
individual trajectory of events is ignored. Usually, we think
that this might only be a problem in the behavioral sci-
ences. But this is not the case, similar dependencies from
previous states and state changes as observed for behavior
can also be observed for physiological states.
A brightness change in the environment is a very basic
stimulus situation which can be very prominent and
attention catching if it is strong enough. Such a simple
stimulus situation does not imply, however, that the
induced changes of brain activity are easy to understand. A
brightness change evokes a cascade of brain activity
changes that can be recorded with distinct measurement
procedures, among others blood flow changes, changes of
the local field potential in visual cortex, spike activity of
individual neurons in visual cortex, etc. In order to extract
the changes that are systematically related to the stimulus,
the signal-to-noise ratio has to be increased and this is
normally done by averaging. Spontaneous fluctuations
of brain activity not directly related to a stimulus are seen
as noise that does not carry any systematic information.
Arieli, Sterkin, Grinvald and Aertsen (1996) studied on
a single trial basis the correlation between the brain activity
that prevailed when a light stimulus was presented and the
system states occurring later on. To this end they recorded
the responses of single neurons and of small neural net-
works in the visual cortex of the cat. The key question was
how much variance of a post-stimulus brain activity mea-
sure can be predicted from the brain activity measured at
the time of stimulus presentation. The results were more
than surprising: at many locations within a cortical area of
2 9 2 mm the correlation between initial measurement and
a measurement taken at a later point in time (e.g., 42 ms
after stimulus onset) lay between 0.8 and 0.9. Due to the
high correlations observed in this experiment it has to be
concluded that the activity of a neural net is often more
determined by the initial state in which the brain was when
a stimulus was presented than by the stimulus-induced
changes themselves. Correlations of the reported size
explain between 60 and 80% of common variance. Arieli
et al. (1996) also predicted the brain activity for a particular
time after stimulus presentation by means of the activity
measured at stimulus onset. These predicted activity pat-
terns were remarkably similar to what had been actually
observed in the visual cortex. All in all, these results
demonstrate very clearly that the initial statewhich is the
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Psychological Research (2012) 76:131144
endpoint of a long trajectory of previous state changes
maybe more important to predict ongoing brain activity
than the changes induced by an external stimulus. This
casts some severe doubt on whether it is theoretically
feasible to regard spontaneous fluctuations of brain activity
just as noise that can be eliminated by averaging over a
large number of trials.
Causal relationships
Humans have a strong tendency to organize their percep-
tual and conceptual world by means of simple causal
relationships (e.g., Newman, Choi, Wynn, & Scholl, 2008).
This trait is fundamental to nervous systems as it enables
an organism to survive in a world which is, by and large,
predictable on the basis of spatialtemporal contiguities
and contingencies.
However, monocausal relationships, as they are descri-
bed by simple ifthen statements, are not very suitable to
describe the functions of the brain and its interactions with
a complex environment. Boththe brain and the envi-
ronmentare continuously interacting and changing. And
these changes are the result of thousands of influences that
work simultaneously. Thus multicausality is a fact, but
everyday explanations as well as scientific explanations are
most often monocausal. The reason is that attention of
laymen and experimenters is usually focused on one input
output variable pair only while other variables are either
ignored or assumed to be sufficiently controlled.
Feedback
Moreover, within the nervous system many feedback
connections exist. For example, filter networks of the
sensory systems that follow after the first level of pro-
cessing feed back to these primary networks and influence
the following upcoming information (Lamme & Roelfsema,
2000). In such systems with many feedforward and feed-
back connections simple monocausal relationships do not
exist. System states develop due to mutual interactions of
many elements over several time steps and such dynamics
cannot be comprehended by means of simple ifthen
statements.
Multifunctionality
Many studies show that components of the nervous sys-
tema single neuron, smaller neural networks, and cir-
cumscribed cortical areasare not functionally unique,
i.e., the activity recorded from these structuresaction
potentials, local field potentials, or BOLD signals
respond to many qualitatively distinct experimental vari-
ables. For example, Platt and Glimcher (1999) recorded
Psychological Research (2012) 76:131144
action potentials in the parietal cortex of monkeys who
were involved in perceptual decision tasks. At the begin-
ning of a trial the firing rate of a neuron was mainly
determined by the expected gains and losses associated
with a learned stimulusresponse combination. At the end
of a trial, the activity of the very same neuron reflected
mainly the direction of the motor response executed to
achieve a reward, i.e., one and the same neuron coded
differently defined psychological variables, or, in other
words, one and the same neuron was found to be integrated
in functionally distinct networks. Reversing the perspective
this result shows that the firing rate of a neuron as such is
not diagnostic for a specific function. Without knowing the
experimental context, the firing rate is ambiguous. The
same holds for BOLD signal changes within circumscribed
brain areas. Areas in the prefrontal cortex, e.g., are acti-
vated by many conceptually distinct tasks. This seems to be
one of the reasons why functional claims about such areas
depend on the theoretical framework a lab is most closely
tied to (see discussions about whether the left inferior
prefrontal cortex, in particular the pars opercularis and pars
triangularis, are functionally more linked to working
memory, action perception, or language parsing, e.g.,
Grodzinsky & Santi, 2008).
Such findings suggest that a specific psychologically
defined intervention, state or process does not correspond
to the activation of a narrowly circumscribed brain area but
rather to an activation configuration within the distributed
networks. For the time being there are hardly any tools to
evaluate and compare such activity changes that simulta-
neously involve many cortical regions. The mainstream of
current research still tries to identify specific functions with
specified structures, while concepts of multifunctionality,
network configurations, and dynamic interactive processes
are hardly taken into account.
Ideographic versus nomothetic explanations
and predictions
The human brain is built from about 1,000 billion neurons
(1012 = 1,000,000,000,000; Nauta & Feirtag, 1986). Each
neuron contacts on average 10,00020,000 other neurons,
i.e., there are approximately 1016 (10,000,000,000,000,000)
synaptic contacts between neurons. Just for comparison:
the number of stars in the milky way adds up to about
2 9 1011 (200,000,000,000), this number is 5 magnitudes
smaller. These numbers are already gigantic but dimension
become even more dizzying if one considers the number of
possible activation states or state changes that are possible
in such an interconnected system. Assuming that there are
only two switching states at each contactthe synapse can
either be active or not, information will be transmitted or
141
notthen there could be (210)16 = 2160 system states. But
this is most likely not realistic enough, because synapses
cannot just toggle between two states but the transmission
characteristics will vary gradually. Therefore, about 10160
or 20160 different system states might result. This gigantic
set of distinct system states is the biological basis of states
and processes that become manifest in subjective experi-
ence and objective behavior. The complexity of such a
system is only partially described by the mere numbers; it
also results from the dynamic interactions of the elements
which continuously change the connectivities between the
elements.
The dimensions are difficult to grasp but they make
clear that states and processes of the nervous system will
never be described completely and exhaustively but only
exemplarily and on principle. A complete description
would mean that an observer is able to measure all synaptic
transitions at a particular point in time. Only if such an
exhaustive description of the system was available, an
exact prediction would be possible to anticipate how an
external signal would act on each element of the system
and which system output would then be generated. Such a
prediction without probabilistic uncertainty is not within
reach and as unlikely as a full description of the movement
trajectories of each and every molecule of some gas in a
container. In order to understand and predict the system
state in total, the microcausalities have to be aggregated in
statistical laws.
Acknowledging that relations between neuroscience and
behavior can only be statistical makes clear that it is not
possible to predict behavior of a single individual for a par-
ticular point in time in the sense of a strict causal relationship
and with full certaintyi.e., if biological measurement
x has a value of xl then under the influence of stimulus y the
organism will inevitably show behavior z. Predictions will
only be probabilistic, i.e., if biological measurement x has
value xl then under the influence of stimulus y the organism
will show behavior z with probability p.
Experiments to predict behavior on the basis
if biological signals
Benjamin Libet was one of the first who studied relation-
ships between physiological measures and the predictabil-
ity of behavior. He looked at the temporal relationships
between the readiness potential in the EEG that precedes a
button press and the report of participants about their
movement intentions and their feeling when they volun-
tarily started a movement. His main argument was that the
decision into which direction a movement will be executed
and the start of such a movement as indicated by physio-
logical changes lies far before the point in time when an
actor feels that he or she voluntarily starts the movement
123
142
(Libet, 1965). However, these experiments suffer from
some methodological and logical flaws that constrain the
conclusions that can be drawn (Rosler, 2008).
In the meantime, experiments have been designed which
provide clearer statements about the predictability of motor
acts on the basis of physiological measures. These show
very nicely what is and what is not possible when it comes
to such predictions. A prototypical example is a study of
Soon, Brass, Heinze, and Haynes (2008). In this study a
running memory task was combined with a self-initiated
free choice response task. The participant had to monitor a
sequence of letters. While these were presented one after
another on a screen the participant had to operate at a freely
chosen time one of two response keys. Later, at the end of a
trial, the participant was asked to report the letter that was
visible on the screen when he or she had decided to operate
the response key. By means of this the time interval could
be delineated in which the subjectively felt decision was
made. Averaged over the group of participants this time
preceded the actual key press by 630 ms. Brain activity
was monitored simultaneously by means of fMRI. The
BOLD changes of voxel clusters were used to predict
which of the two alternative response keys would be
selected by the participant. Surprisingly, the BOLD signal
of some brain areas revealed already 8 s before the actual
key press which key would be operated. In some sense the
finding is quite similar to that of Arieli et al. (1996) cited
earlier, i.e., the brain activity at the beginning of a time
series has predictive power with respect to what happens at
its endhere it is the spontaneous activity at the 3.
beginning of a trial that determines to some degree the
activity at the end and, by means of this, an overt response.
However, as impressive as this result might be, one should
not miss an important fact: the prediction is not causal but
statistical. The prediction of the response on the basis of
the recorded blood flow change was correct in not more
than 60% of the trials, this is significantly better than
chance but not overwhelmingly better. This statistical
value of the hit rate was aggregated over roughly 100 trials
of a single participant and 36 participants in total. Thus, the
method reveals a correlation between brain states and later
behavior but this predictive correlation is small (phi
amounts to approximately 0.20) and it does not allow to
predict what will happen in a single trial of a single par-
ticipant. Science is far away from permitting such ideo-
graphically relevant forecasts.
Moreover, with respect to the generalizabilty of such
findings towards other decision situations something else
has to be considered. In the study of Soon et al. (2008)
participants had to choose between two alternatives,
pressing either the left or the right response key. In real life,
without the restriction of the decision space to such a small
subset, the number of response alternatives and state
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Psychological Research (2012) 76:131144
changes will be almost infinite. Accordingly, an algorithm
that shall predict choice behavior from brain activity must
be able to discriminate between billions of system states.
This is not possible and the idea of mind reading in the true
sensethat biological measures disclose what a person is
thinking and what he or she is planning to do next
remains pure fiction.
Summary
Behavioral neuroscience has the goal to establish rela-
tionships between psychological concepts on one side and
biological concepts on the other. This mapping problem
comprises some basic difficulties:
1. Concepts of the two domains are not defined equiv-
alently. Psychological concepts are ontologically
subjective while biological concepts are ontologically
objective. This means that there exist physically exact
operationalizations for biological measures and obser-
vations while operationalizations of psychological
measures and observations depend on subjective and
social connotations.
2. Concepts on each side of the psychophysiological
equation do not necessary belong to the same level of
a conceptual hierarchy in which superordinate con-
cepts and their properties emerge from subordinate
concepts.
The mapping of concepts is most often not unique,
because a concept which is clearly defined and
separated from other concepts within one domain
maybe related to several distinguishable concepts in
the other domain.
4. Another restriction with respect to unique mappings
follows from the fact that biological systems comprise
complex feedback mechanisms and the features of
biological entities emerge from a continuous interac-
tion of millions of neurons. This means that one and
the same elementa neuroncan be part of func-
tionally distinct networks and, therefore, an element or
even a set of elementsa neural networkcan reveal
distinct features. Thus, the functional significance of
observed entities can be ambiguous.
Due to these restrictions it is not possible a priori to
decide which concepts of one domain match with which
concepts of the other. Psychophysiological mapping is an
empirical process and the criterion of the goodness of fit is
provided by the amount of mutually explained variance.
Moreover, psychophysiological mappings cannot be
understood in the sense of substitutions, but only in the
sense of a translation.
Psychological Research (2012) 76:131144
Even if clear-cut psychophysiological relationships are
given it has not to be missed that such relationships are of a
statistical nature only, irrespective of the fact that on a
microlevel causal relationships may exist. This restriction
follows from the gigantic number of elements from which
the nervous system is built and the even larger number of
possible state and state changes that can exist in such a
complex system.
As a consequence it has to be taken into account that
behavioral predispositionse.g., whether someone will
become a criminal offender or notcan only be deter-
mined in the sense of statistical forecasts not in the sense of
exact causal and ideographically valid predictions. Here,
the same indeterminacy holds as in thermodynamics.
Moreover, the question of whether a biological sign can be
used for predicting the predisposition for a pathological
behavior is not an issue of neuroscience alone, but also
depends on individual and political decisions that follow
from the social-historic context.
References
American Psychiatric Association. (2000). Diagnostic and statistical
manual of mental disorders DSM-IV-TR (4th ed.), Arlington,
VA: American Psychiatric Publishing, Inc.
Arieli, A., Sterkin, A., Grinvald, A., & Aertsen, A. (1996). Dynamics
of ongoing activity: Explanation of the large variability in
evoked cortical responses. Science, 273(5283), 18681871.
Baddeley, A., Eysenck, M. W., & Anderson, M. C. (2009). Memory.
Hove: Psychology Press.
Baltes, P. B., Reuter-Lorenz, P., & Rosler, F. (Eds.). (2006). Lifespan
development and the brain. Cambridge: Cambridge University
Press.
Barsalou, L. W. (2008). Grounded cognition. Annual Review of
Psychology, 59, 617645.
Basar, E., Flohr, H., Haken, H., & Mandell, A. J. (1983). Synergetics
of the brain. Berlin: Springer.
Campbell, J. I. D. (1991). Conditions of error priming in number-fact
retrieval. Memory and Cognition, 19, 197209.
Churchland, P. S., & Sejnowski, T. J. (1989). Neural representation
and neural computation. In L. Nadel, L. A. Cooper, P. Culicover,
& R. M. Harnish (Eds.), Neural connections, mental computation
(pp. 1548). Cambridge, MA: MIT Press.
Dell, G. S. (1986). A spreading-activation theory of retrieval in
sentence production. Psychological Review, 93, 283321.
DIMDI (Ed.) (2010). ICD-10Internationale Klassifikation der
Krankheiten, 10. Revision. Retrieved from http://www.dimdi.de/
static/de/klassi/diagnosen/icd10/index.htm.
Druzgal, T. J., & DEsposito, M. (2003). Dissecting contributions of
prefrontal cortex and fusiform face area to face working
memory. Journal of Cognitive Neuroscience, 15(6), 771784.
Fahrenberg, J. (1979). Das Komplementaritatsprinzip in der psycho-
physiologischen Forschung und psychosomatischen Medizin.
Zeitschrift Fur Klinische Psychologie Und Psychotherapie, 27,
151167.
Goldman-Rakic, P. S., OScalaidhe, S. P., & Chafee, M. V. (2000).
Domain specificity in cognitive systems. In M. Gazzaniga (Ed.),
The new cognitive neurosciences (pp. 733742). Cambridge,
MA: MIT Press.
143
Grodzinsky, Y., & Santi, A. (2008). The battle for Brocas region.
Trends in Cognitive Sciences, 12(12), 474480.
Gromann, S. (2007). Kausalitat aus der Sicht eines Physikers. In
Berlin-Brandenburgische Akademie der Wissenschaften (Ed.),
Kausalitat (pp. 1930). Berlin: BBAW.
Haynes, J. D. (2009). Decoding visual consciousness from human
brain signals. Trends in Cognitive Sciences, 13(5), 194202.
James, W. (1890). The principles of psychology. New York: Holt.
Lamme, V. A., & Roelfsema, P. R. (2000). The distinct modes of
vision offered by feedforward and recurrent processing. Trends
in Neurosciences, 23(11), 571579.
Levelt, W. J. M. (1989). Speaking. Cambridge, MA: MIT Press.
Libet, B. (1965). Cortical activation in conscious and unconscious
experience. Perspectives in Biological Medicine, 9, 7786.
Magistretti, P. J., & Pellerin, L. (2000). Regulation of cerebral energy
mechanism. In C. T. W. Moonen & P. A. Bandetti (Eds.),
Functional MRI (pp. 2534). Berlin: Springer.
Mitchell, M. (2009). Complexity: A guided tour. New York: Oxford
University Press.
Monsell, S. (2003). Task switching. Trends in Cognitive Sciences,
7(3), 134140.
Nauta, W. J. H., & Feirtag, M. (1986). Fundamental neuroanatomy.
New York: Freeman.
Newman, G. E., Choi, H., Wynn, K., & Scholl, B. J. (2008). The
origins of causal perception: Evidence from postdictive process-
ing in infancy. Cognitive Psychology, 57(3), 262291.
Nunez, P. L. (1981). Electrical fields of the brain. New York: Oxford
University Press.
Pinel, J. P. J. (1992). Biopsychology. Boston: Allyn and Bacon.
Pinto, D., Pagnamenta, A. T., Klei, L., Anney, R., Merico, D., Regan,
R., et al. (2010). Functional impact of global rare copy number
variation in autism spectrum disorders. Nature, 466, 368372.
Platt, M. L., & Glimcher, P. W. (1999). Neural correlates of decision
variables in parietal cortex. Nature, 400(6741), 233238.
Quiroga, R. Q., Kreiman, G., Koch, C., & Fried, I. (2008). Sparse but
not grandmother-cell coding in the medial temporal lobe.
Trends in Cognitive Sciences, 12(3), 8791.
Rosenzweig, M. R., & Leiman, A. L. (1982). Physiological psychol-
ogy. Lexington, MA: D.C. Heath.
Rosler, F. (1983). Physiologisch orientierte Forschungstrategien in der
Differentiellen und Diagnostischen Psychologie: I. Zur Konzeption
des Psychophysiologischen Untersuchungsansatzes. Zeitschrift fur
Differentielle und Diagnostische Psychologie, 4, 283299.
Rosler, F. (1984). Physiologisch orientierte Forschungstrategien in
der Differentiellen und Diagnostischen Psychologie: II. Zur
Systematisierung psychophysiologischer Untersuchungen. Zeits-
chrift fur Differentielle und Diagnostische Psychologie, 5, 736.
Rosler, F. (2008). Was verraten die Libet-Experimente uber den
freien Willen? Leider nicht sehr viel! In E. J. Lampe, M.
Pauen, & G. Roth (Eds.), Willensfreiheit und rechtliche Ordnung
(pp. 140166). Frankfurt am Main: Suhrkamp.
Searle, J. R. (1987). Intentionalitat. Frankfurt a. M.: Suhrkamp
Taschenbuch.
Searle, J. R. (2000). Consciousness. Annual Review of Neuroscience,
23, 557578.
Soon, C. S., Brass, M., Heinze, H. J., & Haynes, J. D. (2008).
Unconscious determinants of free decisions in the human brain.
Nature Neuroscience, 11(5), 543545.
Speckmann, E. J., & Caspers, H. (Eds.), (1979). Origin of cerebral
field potentials. Stuttgart: Thieme.
Stemmler, G. (1989). The autonomic differentiation of emotions
revisited: convergent and discriminant validation. Psychophys-
iology, 26(6), 617632.
Treisman, A. (2004). Psychological issues in selective attention. In M.
S. Gazzaniga (Ed.), The cognitive neurosciences III (pp. 529
544). Cambridge, MA: MIT Press.
123
144 Psychological Research (2012) 76:131144

van Essen, D. C., Anderson, C. H., & Felleman, D. J. (1992). Savodnik (Eds.), Consciousness and the brain (pp. 205268).
Information processing in the primate visual system: An New York: Plenum Press.
integrated systems perspective. Science, 255, 419423. Wundt, W. (1874). Grundzuge der Physiologischen Psychologie.
Watson, W. C. (1981). Physiological psychology. Boston: Houghton Leipzig: Engelmann.
Mifflin. Comp.
Wimsatt, W. C. (1976). Reductionism, levels of organization, and the
mindbody problem. In G. G. Globus, G. Maxwell, & J.

123
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