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(extended version)
Linda Palmer
Department of Philosophy, Carnegie Mellon University
Contact information:
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lpalmer@cmu.edu
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Shortened version in press in Review of General Psychology: last revision February 16, 2008
Extended version (restore figures 1, 3 and associated text, etc): last revision Oct 2008
Kant and the brain: A new empirical hypothesis
Linda Palmer
Carnegie Mellon University
Immanuel Kant’s three great Critiques stand among the bulkier monuments of
Enlightenment thought. The first is best known; the last had until recently been rather
less studied. But his final Critique contains, I contend, a remarkable development of
Kant’s theory of how human beings use and create systems of knowledge. While Kant
was not himself concerned with the neuronal substrates of cognition, I argue this
development yields a novel empirical hypothesis susceptible of experimental
investigation. Here I present the Kantian motivation and describe experimental work
aimed at testing predictions arising from the new hypothesis.
1 Of course considerable detail and nuance will have to be set aside in this brief overview, for both Kant’s position and
the neuroscience involved.
KANT AND THE BRAIN 2
2. Kant’s basic theory of cognition concept of cause (which Hume pointed out could
2.1 Metaphysical and epistemological background be neither deductively nor inductively grounded);
In his account of cognition, Kant opposes the rather, we presume the world is causally connected,
empiricist view that all mental contents are drawn and the project of science is to look for what those
from experience. In the Critique of Pure Reason causes may be. “Cause” is thus an a priori concept.
Kant famously introduced the notion of “a priori Hereafter, however, we will be concerned with
conditions” of cognition which, he argued, cannot ordinary, a posteriori concepts, and their use in
be derived from experience but must instead be cognition.
given prior to it.2 Certain aspects of our knowledge
of the world, he argued, are not based on but are 2.2. The theory
instead put into our experience of the world by the
Kant was by no means an empirical
mind, in constructing our mental representations of psychologist; his cognitive theory was motivated
an objective world.3 His theory identifies two basic by epistemological and metaphysical concerns like
kinds of conditions: two “a priori pure forms” of those sketched above.6 At the same time I think it
space and of time, and twelve original “a priori worthwhile to inquire into the potential
pure concepts” (such as “substance”, “quantity”, neurobiological substrates of the functions he
and “cause”); these were to be regarded as a priori identifies. A basic sketch of the theory Kant gives
constraints of thought rather than results of in the Critique of Pure Reason follows. Take first
investigation or experience like a posteriori, the case of object recognition. Kant holds the
empirical concepts such as “water”.4 mental representation of some given external
Kant intended his position to solve certain object is by no means a direct “copy” of the
metaphysical problems like the applicability of sensible impressions, as held by the earlier
mathematics to nature and a defense of “cause” empiricist philosophers. To the contrary, Kant
from Hume’s attack. Our knowledge of argued that data received through the senses - the
three-dimensional Euclidean space is not based on “manifold” of sensory material, as he termed it -
experience of objects; rather, in order to represent must be transformed into our cognition of an
an object in the first place, the mind arranges given object, through the constructive work of two
sensory material into the three spatial dimensions. further mental capacities beyond the sensory.
The world we experience conforms to the formal These he called the “imagination” and
structures of Euclidean geometry because we “understanding” respectively (see e.g. pp.
construct it as such.5 Similarly, Kant holds we do A78/B103; pp. A120-121; p. A126).
not look to the natural world to discover the
2 Kant here uses the traditional philosophical distinction between “a posteriori” knowledge, as based on or derived from
experience, versus “a priori” knowledge, regarded as not based on experience but known independently thereof. The
classic examples are mathematical and logical: “how many coins are in the tip jar?” is to be answered a posteriori, by
examining the jar in question; “what is the sum of seven plus five?” is not properly answered by empirical investigations
but is supposedly determined a priori. Note traditionally “a priori” was considered to be quite distinct from merely “in-
nate” in a psychological sense, as instincts or drives are innate. In what sense and how mathematics thus “transcends the
contingent” is an open question today.
3“It may well be that even our empirical knowledge is made up of what we receive through impressions and of what our
own faculty of knowledge (sensible impressions serving merely as the occasion) supplies from itself” (Kant, 1781/1965,
p. B1). Page references to this work use standard A/B form referring to the first and second editions, given in the margins
of this translation.
4For space and time, see the Transcendental Aesthetic (Kant, 1781/1965, pp. A19/B33-A49/B73); for the pure concepts,
see the first part of the Transcendental Logic, particularly the table of categories (Kant, 1781/1965, p. A80/B106).
5 This means in Kant’s view mathematics applies to the natural world only insofar as we represent it (the world); indeed
Kant held we can know nothing whatsoever about the world considered independently of the conditions of our represen-
tation of it - this is his controversial doctrine of “transcendental idealism”. The basis of the remarkable applicability of
mathematics to nature, as well as the nature of mathematical objects, is still subject to debate.
6 I can hardly overemphasize this point: this is not an empirical reading of Kant’s philosophy itself. Kant meant his work
to be transcendental philosophy, not empirical science. But there is no reason we may not derive some empirical hypothe-
ses from Kant’s philosophy. In what follows I also use present-day results to illustrate or elucidate Kant’s position.
KANT AND THE BRAIN 3
other information normally used by the brain to time postulated in his theory “the pure forms of
stabilize the resulting perception. intuition”.
Kant thus argued that imagination is involved
We have seen Kant argues perception requires
in all perception, not just (as in the colloquial use active reconstruction in space and in time; it will
of the term) the ability to recall previously- also require concepts, discussed below. Studies of
experienced scenes or objects or construct the neurobiological substrates of vision have begun
“imaginary” ones. He also took imagination to to identify such functions in brain. Ungerleider and
involve all sensory modalities, not just visual. Mishkin (1982; see Fig. 4) showed that spatial
Different sensory modalities are united in the location (i.e. “where”) is processed in the so-called
perception of an object: as you taste your morning “dorsal stream,” proceeding from early visual
coffee, you see the cup and the steam; you feel the sensory areas in occipital cortex to dorsal areas of
weight and the smoothness of the porcelain, and posterior parietal cortex, while object shape is
the heat, the scent, and the taste of the coffee itself. probably in the “ventral stream” proceeding from
According to Kant, this unification ultimately early visual areas to infero-temporal cortex, which
requires concepts, to which we turn in the next subserves object recognition (i.e. “what”). The
section. neuronal bases of time perception - both elapsed
Kant’s technical term for the spatio- time (duration) and location in universal historical
temporalized sensory materials, as produced by the time - are not yet well understood. Cannabinoids
imagination, is Anschauung, translated “intuition.” are known to distort time perception; basal ganglia
This is quite distinct from the lay use of the term in and cerebellum have been implicated in timing;
English, meaning something like “hunch” or “gut perception of elapsed time is reported to involve
feeling.” For Kant the term “intuition” has nothing areas in parietal cortex and attentional systems
to do with hunches or feelings, but refers to the (Iversen, 2001; Harrington et al, 1998; Eagleman et
spatiotemporalized sensory material produced by al, 2005).
imagination; thus we may refer to the sensible
The task of the imagination, then, is to take the
intuition of the glass of milk, that of the cup of raw sensory material provided by the senses, and
coffee, that of the table covered by cloth, or that of arrange it in space and in time. Kant’s recognition
the scene as a whole. The imagination produces a that the three-dimensional shapes and locations of
“sensible intuition” from given sensory materials sensed objects, and their time order as enduring or
by arranging them in space and time. Similarly, successive, are not simply already given by the
Kant calls the “pure a priori forms” of space and sensory input but must be reconstructed via active
representational processes, represents a
considerable advance over the views of his
contemporaries.
Posterior Do
Parietal rs
Cortex al 2.2.2 The understanding, or, the home of concepts
St
re
am
So far we have discussed imagination, but for
Kant cognition also requires an appropriate
concept, provided by “the understanding” (p.
Primary A78/B103). Kant emphasizes that neither the
Visual intuition alone, nor the concept alone, suffice; both
Cortex
are needed. The cartoons in Figure 5A may help to
highlight the conceptual contribution.
m Wittgenstein’s “duck-rabbit” illustration and other
rea
l St examples can be interpreted in different ways,
Infero- ra depending on the concept chosen. The point of
nt
Ve
temporal
Cortex such examples is not in the illusion per se, but
simply to bring our attention not only to the
Figure 4. Dorsal and ventral pathways for vision. imagination’s spatiotemporal reconstruction of
Upstream processing is represented by the arrows; given materials, but also to the point that this
recurrent feedback connections are not indicated. activity is guided by the concept used in
Adapted from Milner & Goodale 1995.
recognizing just what it is. Thus the construction of
KANT AND THE BRAIN 5
Kant emphasizes this is not only a matter of hypothesis is summarized in Figure 6 (the region
combining the several different senses (what is anterior to primary motor cortex, broadly
sometimes termed “the binding problem”) but corresponding to ‘frontal cortex’ - an area vastly
combining and separating, as appropriate, the expanded in human brain - is here referred to as
manifold of data within each sensory stream. This ‘anterior cortex’). Work in the past century has
“manifold” of sensory data, which involves not shown that each distinct sensory modality (vision,
only multiple senses (visual, auditory, taste, touch, touch, etc) inputs to its own distinct primary
etc) but also a multitude of sensory material within sensory area within the posterior cortex, cascades
each sensory stream, needs to be unified into a through further stages of processing in secondary
single stable perception of a scene (objects and/or areas, and eventually converges in the so-called
events). In Kant’s view, this act of unification “association areas” of cortex, which thus receive
requires the use of concepts. Look at the coffee cup multiple sensory modalities. At the same time there
on your desk: the visual data from the cup must be are extensive recurrent connections among the
separated from those of the table surface, and cortical areas: each stage of processing is not
combined together as belonging to the one object, merely sequential, as in an assembly line, but
perhaps with the scent of coffee and anticipation of involves recurrent connections from secondary
its taste as well; the table’s sensory data likewise sensory areas and association areas (Fig. 6A). As
bound together and perceived as a unitary object mentioned above, for example, it appears that
(see Fig. 5B for a similar example). In both cases, visual recognition of complex objects is at least in
this separation and grouping requires the part subserved in inferotemporal cortex (the
application of the appropriate concept. And both “what” pathway) while primary processing of
table and cup are placed within a unique location in specific cue features (edge-detection, motion,
all of space, and in all of time. color) are accomplished at earlier stages. The
Kant actually considers this unity of feedback influence of higher-level conceptual
perception or thought to stem from the unity of representations on lower levels of sensory
consciousness itself, and that from “the possibility processing proposed in the Kantian model, and
of the ‘I think’ ”, i.e. the representation of a self evident in the playing card experiments, would be
(pp. B131-136). Searle (1997), with his usual well subserved by the massive recurrent
dryness, speaks of Kant’s flair for catchy phrases in connections observed between higher association
his calling this the “transcendental unity of cortex regions and earlier sensory cortical regions.
apperception”. The neuroscientists Gerald Edelman This implies that a conscious perception would
and Antonio Damasio discuss the role of the involve regions of activation distributed across
representation of the self in perception (Edelman, different regions of cortex, as has been suggested
2004, Damasio, 2000); these are important by a number of neuroscientists. (Singer and
questions, which we can not pursue here. colleagues have identified recurrent synchronous
firing in the gamma band, around 40 Hz, as a
2.3 Neurobiological substrates correlate of conscious perception (Engel and
Kant’s views deeply affected the 19th century Singer 2001), while Edelman’s theory of
founders of neuroscience (Helmholtz being an “functional clustering” of activations across
especially clear example), and have ready distinct neuronal groups addresses the issue of the
correspondences in contemporary ideas about how integration and differentiation, or unity in diversity,
the cortex creates representations of objects and of mental contents (Tononi & Edelman, 1998).)
sequences in the environment. What follows is
Anterior association regions, sampling the
necessarily quite simplified,7 but it appears that output of the posterior (sensory) association zones,
“universalizing” functions would be subserved in feed back into them (Fig. 6B) and are assumed to
more anterior cortical regions, and “particulars” in provide the rules whereby the encoding of ‘what’
more posterior regions, where sensory inputs first occurs in those latter areas. In the human brain, the
arrive in cortex. This general and widely accepted anterior-posterior relationship is further evident in
7 The neurobiological substrates of judgment are of course far from fully understood, but even so, this account will leave
out a number of important subcortical systems also likely to be involved, notably the cortico-striatal loops, the cortico-
thalamic loops, the hippocampal system, attentional systems, and so on.
KANT AND THE BRAIN 7
A
primary
sensory
primary
sensory
primary
sensory
assoc
secondary
assoc sensory
input
anterior posterior
B
primary
sensory
primary
sensory
primary
sensory
assoc
secondary
assoc
Figure 6. Posterior-anterior recurrence in cortex. A. Sensory inputs arrive in the several distinct primary sensory
regions in posterior cortex (one for each sensory modality) and cascade upwards through secondary and higher sensory
areas, converging in multi-modal association areas, which themselves send recurrent connections back to sensory areas.
B. Anterior regions show extensive recurrent connections with more posterior association and sensory areas.
the processing of words and their assembly into according to extant or, sometimes, new concepts (a
sentences via the rules of grammar. The red spade!). But the question arises as to how the
development of language means that concepts are recurrent cycling of activity among anterior and
associated with words, which can be used in posterior cortical association areas results in a
statements. It is well known that two normally neuronal ‘decision’ that an appropriate
left-cortex areas are language related: Wernicke’s representation of ‘what’ has been attained. This is a
area, located more posteriorly, subserves content profoundly difficult question for brain theories and,
and comprehension, while Broca’s area, located as discussed below, is one Kant belatedly realized
more anteriorily, relates to syntax and sequencing was faced by his own system.
of thoughts. Damage to Wernicke’s area, near the
junction of temporal and posterior parietal cortex, 3. New developments in the Critique of
yields difficulties in comprehension and in speech Judgment
production that is syntactically fluent but
incoherent (i.e. the patient uses words in fluent
Like much of Kant’s work the Critique of
sentences but the content does not make sense). Judgment is notoriously difficult, and has inspired
Damage to Broca’s area, in the frontal cortex and considerable scholarly controversy over its
near motor systems, produces speech that has interpretation. Where the first Critique proposed a
relevant content words, but little or no syntactical set of a priori conditions of our representation of
structure. Propositional judgments can not be objects, the third Critique discusses the a
expressed if this anterior language center is posteriori side of cognition. In the search for
sufficiently damaged. empirical laws and concepts, Kant notes in the
Finally, it is clear the brain must somehow Introduction to the latter work, we systematize our
generalize on the basis of experience. For simple empirical knowledge, and he points out we must
stimuli and responses this can be understood in a assume there are suitable generalizations to be
fairly mechanical fashion; e.g. the entire neuronal found - that nature is not too chaotic or random to
circuit for eyeblink conditioning has been traced be comprehended at all. (We can only assume this:
(i.e. the circuit responsible for the conditioned this is not something we can conclude on the basis
response of eyeblink to an auditory tone previously of having already found such regularities, for it is
repeatedly paired with an air puff to the eye). But induction itself that is at issue). In this search, we
the brain is capable of far more sophisticated are looking for appropriate generalizations to
generalizations. In Kant’s terms, judgment capture or describe given particulars (the data or
organizes the contents of new experience, phenomena to be explained). In the Critique of
KANT AND THE BRAIN 8
8 This brief sketch must neglect Kant’s account of how fine art differs from natural beauty.
KANT AND THE BRAIN 9
cognitive powers, which normally results in
In his analysis of judgments of beauty, then,
subsuming an intuition produced by the Kant argues there must be a “common inner
imagination under a determinate concept of the sense”: a feeling of the coordination of the
understanding, i.e. in a cognitive judgment, in free imagination and understanding in judgment. He
play instead resonates and continues, in “a play argues this feeling is also found in cognition
which is such that it sustains itself on its own and (indeed, that we must assume the sense is shared
even strengthens the powers for such play” (p. by everyone, in order to avoid skepticism). Rather
313). This is the “facilitated free play of the two than external sensory cues, Kant’s common inner
mental powers (imagination and understanding) sense, it appears, is to report on the processing
quickened by their reciprocal harmony” (p. 219). state of the mind itself, in acts of judging.
Finally, this activity of the mental powers is felt,
Now to our experimental question: Is there
Kant claims, as pleasure (pp. 218-219). such a signal in the brain processing leading to
Note that exactly what aesthetic unity consists cognition? If so can its locus in brain be identified?
in is hard to say: Kant denies it can be captured by Some experiments directed toward these questions
concepts. Verbal descriptions (like those above) of are described below.
certain aesthetically pleasing aspects of a natural
beauty, poem, or painting, are cognitive judgments, 3.2 Empirical work
not themselves experiences of beauty. Such efforts,
As we saw, one would expect potential
Kant notes, can at best serve to point others in the neurobiolobigical substrates of Kantian
right direction, to call their attention in hopes of “universals” to be subserved in more anterior
awakening their own aesthetic judgments. cortical areas, in frontal and temporal association
In a judgment of beauty the imagination does areas, and “particulars” in more posterior cortical
not produce merely random or chaotic connections regions, where sensory inputs arrive and begin to
of the manifold of sense, but rather ones that reflect be processed. Is there a potential neurobiological
a certain unstateable unity or unities (Kant even substrate for the Kantian inner sense, as well? Can
calls it a “free lawfulness”). But it does so freely, we identify a brain region that subserves feeling
without any constraint of determinate rules or and is in a position to signal the occurrence of
concepts. What this means is that we have a coordinated activity between these regions? Figure
capacity to feel a certain unity in the given sensory 8 shows the desiderata: a brain substrate for
diversity, an order, connection, or coherence that Kantian “inner sense”, not an outer sense like sight
isn’t (or isn’t yet) captured by any explicit rule or or hearing but a feeling of the mind’s (or here the
concept. brain’s) activity, in a position to report on
The experience of beauty, then, is the pleasure coordinated activity between anterior associational
one takes in one’s own mental activity in regions in frontal cortex and the posterior cortical
apprehending an object, and doing so freely, associational and sensory regions.
without the normal constraints of taking it
I would like to suggest a particular region of
cognitively. Thus I take it that an activity of the limbic system that appears to offer a good fit
imagination and understanding needed for all with the above two criteria: the basolateral division
cognition is found in its purest form in the
experience of beauty: the imagination searches for
connections in the material offered to it, and the primary
sensory
understanding is stimulated by this in its own primary
attempts to codify, formulate, and formalize. In sensory
Frontal cortex Temporal cortex
For both these reasons, the basolateral division
And: parietal, And: agranular of the amygdala seems well positioned to signal
cingulate, insular,
insular, hippocampal, the occurrence of synchronized, coordinated
prefrontal, prefrontal, activity between frontal/temporal and posterior
olfactory olfactory
cortex cortex cortical regions. Basolateral amygdala thus
Basolateral
BLA
Amygdala
LA appears to be a suitable candidate both in terms of
BLA LA connectivity and as a potential substrate for a
feeling signal reaching consciousness.
Striatal systems Perigeniculate nucleus,
thalamus
Other amygdalar nuclei 3.3 Experimental tests of this idea
Figure 9. Major connections of the “frontotemporal”
A set of recent learning studies in rats (Hess,
amygdala in rat, adapted from Swanson and Petrovich Gall, Granger, & Lynch, 1997) are promising in
1998. Note the extensive bidirectional connectivity of
the anterior basolateral nucleus (BLA) with frontal regard to the hypothesis of the basolateral
cortex in particular, and of lateral amygdala (LA) with amygdala as a potential substrate of a cognitive
temporal cortex. Additional cortical regions with signal on the Kantian model. Figure 10 shows
reciprocal connections to BLA and LA are indicated.
(Outputs of both to the striatal system and directly or
sections from rat brains through amygdala under
indirectly to other amygdalar nuclei, and input from two different behavioral conditions. The
perigeniculate thalamus, also indicated). basolateral division is highly active in one of these
of the amygdala. The amygdala as a whole is an conditions: when the animal has just learned
evolutionarily ancient subcortical structure, in something new.
humans located at the anterior ventral pole of the
In the studies, rats were trained on a two-odor
hippocampal formation. The most well-studied discrimination task: a rat must learn to poke its
division of the amygdala is its central nucleus. The nose into one of two ports, marked with distinct
central nucleus is known to subserve a particular odors, to receive a water reward (they have had
kind of feeling signal: it is well established as a temporarily limited access to water prior to the
substrate for fear and anxiety responses. While fear experiments). When a rat has first learned the
itself is not the signal that we are interested in - to correct odor cue (it begins to nosepoke only to the
the contrary, we are interested in a feeling of correct port), the basolateral division of the
pleasure - it is certainly a feeling-signal that amygdala shows marked activation, the researchers
reaches consciousness. Could a different division discovered. Remarkably, they found the basolateral
of the amygdala be a potential substrate of the division is not active when the animal is engaged
pleasurable and cognitive signal of the sort Kant in the identical overt behavior later, after the task
suggests? has been well-learned. Nor was it active earlier,
In fact, the basolateral division of the when the animal is in the same apparatus and
amygdala has extensive reciprocal connections nosepoking, but before it has learned to choose
across cortex. As such it appears well positioned only the correct odor. (It is also not active in “home
to report on the occurrence of coordinated activity cage” control animals.) It thus appears that the
between anterior and posterior cortical regions. basolateral division activity correlates with the
Figure 9 shows a diagram of some major inputs condition that something new has been learned - in
and outputs of the basolateral divsion of the
amygdgala - that is, the lateral nucleus and the
anterior division of the basolateral nucleus, what
Swanson and Petrovich (1998) identify as the
frontotemporal system of the amygdala. (There is
some ambiguity in the nomenclature; I use
“basolateral division” or “basolateral amygdala” to
Figure 10. Basolateral amygdala activation in new
include both the lateral nucleus and the anterior learning. In situ hybridization to c-fos mRNA is used as
basolateral nucleus of the amygdala). Note the a marker of neuronal activity. Left: Central nucleus
extensive, reciprocal connectivity with frontal and (arrow) is active in this brain; this rat had been lightly
stressed by being given an injection for the first time.
temporal cortical regions shown by the basolateral Right: Basolateral division of the amygdala (arrow)
amygdala. shows distinct activation. This rat had newly learned a
two-odor discrimination task. Courtesy of Christine Gall.
Introduction I. Behavior
Subfields of hippocampus form a densely connected serial network: Rat placed in novel environment and allowed to explo
KANT AND THE BRAIN 11
entorhinal cortex dentate gyrus field CA3 field CA1 output via subiculum No explicit reward/punishment in this learning task
other words,
Thus a natural prediction would bejust when
that CA1 the
activity animal
levels hasclosely
will correlate “figured it
with activity in CA3, its major input. Yet earlier work has shown that relative GROUP
activity levels inout”.
CA3 andThese
CA1 in factresults are encouraging
change substantially, for the
during acquisition
of rewarded behaviors (Hess et al. 1995a,b; Gall et al 1998).
hypothesis of the basolateral amygdala as a diagram 1. Animal sacrificed to o
Here we tested potential
if this result also holds for unsupervised learning: ie learning in
of ! levels at time of
substrate for a cognitive signal. test
the absence of explicit external reward or punishment. ! behavior (AOB group
Further studies are currently underway. The environment
Rats placed in an open field with an attached refuge were found to adopt, after
above studies,
initial intense exploration, like
a posture of alertmost animal
observation on thelearning
border of thestudies, Active exploration groups:
two domains (lasting tens of seconds to minutes) before resuming exploration. II. Animal sacrificed to obse
involve the presence of an externally-provided " 30 min prior to AOB (-3
This learning-related
reward changecue in behavior
to shape was accompanied
the animal’s by a dramatic
behavior.shift Such
in the balance of activity between fields CA3 and CA1, as assessed by in situ trace of
hybridization toreward-c-fos mRNA:(or in rats punishment-)
sacrificed to measure based peak mRNA learning is
levels one rat’s III. Animal sacrificed to obse
at time of the observational behavior, CA1 showed relatively low activity levels " 15 min prior to AOB (-1
with higher levels classified
in CA3. This isas “supervised”
the same hippocampal activity learning,
pattern seen meaning
in trajectory
the earlier studies of rewarded learning.
learning based on the presence of external rewards 30 min Control:
Results in rats(or sacrificed to measure neuronal activityorwhile exploringresponses.
the IV. sacrificed directly from t
punishments) for correct incorrect
apparatus corresponded to a distinct pattern earlier shown in rats exploring a " no exposure to novel en
novel environment, We would like to examine unsupervised learning as
namely labeling in both CA3 and CA1 with CA1 slightly
predominant. Figure 11. Unsupervised learning: the novel
well, that is, learning in the absence of any such environment exploration paradigm. Top: diagram of the
These results:
explicitis associated
• confirm acquisition rewardwithora particularpunishment.change in the Much human 2. In situ Hybridization:
test environment. Bottom: trace of c-fos mRNA
one rat’s trajectorylevels reflect neu
pattern of neuronal activity in hippocampus: CA1 < CA3
learning and discovery is of this sort. for thirty minutes.
• extend this observation to include unsupervised learning i. Representative animal from each of four conditions
• indicate the
pertinent
Considerchangesstepping
occur within offminutes a train in an unfamiliar latter; perhaps it is also required and can be found
CA1 relatively lower
We hypothesizecity (orobserved
that the to takechanges a less romantic
reflect the formation example,
within fieldconsider half-sections showing
in the former. control
CA1 of “winner-take-all” modules, a learning-related effect seen in certain
neural network walking
models. into a brand-new shopping mall).
In thelabelling
presentdensities
studies, a well-handled rat is
Exploration of a novel environment is a standard placed in controla novel environment and allowed to
AOB
mammalian behavior. Well-handled, unstressed rats explore; this contains local cues, an enclosed
placed in aMethods novel environment will spontaneously “refuge”, and clear walls (see Fig. 11). After an
explore it,
Animals: 6 week old male Long-Evans rats; and learn objects
handled for four and locations
days prior to testing, in the initial period of exploration in this apparatus, rats
including daily transport in opaque cages to the behavioral laboratory.
environment. Spontaneous exploration of a novel exhibit a new behavior: holding still in or near the
Test apparatus: 18- by 36- inches with clear glass sides, two local cue objects (2 inch high
environment is an example of an “unsupervised
glass bottles), and an enclosed refuge box: visually opaque (dark inside) but transparent entry to the refuge, in a posture of alert
to infrared illumination. Various distal cues are visible in the dimly lit testing room.
learning” paradigm, for there is no explicit external
Infrared LED illuminators allowed an overhead network video camera (Axis observation. We propose that the appearance of this
Communications AB, Lund, Sweden) to track animals while inside the refuge.
reward signal to shape the animal’s learning. behavior may serve as a behavioral marker that
Behavioral sessions and treatment groups: A rat was placed near the entry of the refuge and
At this point a question may be raised
allowed to explore at will, monitored for occurrence of the first ‘alert observation’ learning has -30m taken place. -15m We predict-30m that activation
behavior (AOB) > ~ 45s, and sacrificed to measure peak c-fos levels induced by
concerning Kant’s focus on human cognition.
neuronal activity as follows: at time of AOB occurrence; or 15m or 30m prior to AOB (ie of the basolateral amygdala will be a necessary
during active exploration). Control animals remained in the transport box until
Kant’s theory involves conceptual subsumption,
removed for sacrifice. Brains were fresh-frozen. For each group n=3 except the 15 min concomitant of theii. Serial sections through
unsupervised learningseptal-temporal
in this hippocampus
group (n=2): 11 rats total. AOB was defined as the animal’s sitting, alert but motionless
but it is unknown to what extent rats have anything
(excepting occasional head scanning movements) near the entry of the refuge for a paradigm. These experiments are currently under
Mapping procedure schematic
Compare control to experim
period of > 40 seconds, excluding grooming, local exploration and sleeping. Anterior hippocampal maps of labe Anterior hippocampal maps from 11 animals Compare time course in ope
similar. In response to this question, a two-part way, and we await the results of the analysis of the Experimental condition
Control AOB Controls, in familiar transport box
A
In situ hybridizationassumption
to c-fos mRNA: Levels of c-fos mRNA are
underlying thelow atpresent
backgroundexperiments
and rise • A constant width selection along the cell body layer anterior
quickly after neuronal activity, peaking at about 30 min. 20!m thick brain sections were amygdala activation
is “unrolled” through CA3, CA2,patterns.
CA1. (We have pre-
sectioned in a cryostat
should be outlined. The first assumptiona is
onto glass slides and processed for in situ hybridization to 35S-this: in Unrolling Serial Hippocampal Sections posterior
labeled cRNA probe complementary to rat c-fos mRNA (i.e., to positions 583-1520 of liminary results of these studies in another brain
CA1
CA3
b
CA2 CA1
anterior
CA3 CA2 CA1
1998; Guthrie et al. 1996; Lauterborn et al. 1996; Palmer et al. 1997). Kodak Biomax MR area (Palmer 2007): briefly, a characteristic activity
c
a
CA3 b .
.
film was exposedin to sections for 3 days yielding
the detection film autoradiograms
and encoding of hybridization in the
of regularities .
posterior
density; commercial radiographic standards (American Radiolabelled Chemicals, Inc., pattern seen in the hippocampus at the time of
Paxinos and Watson Serial Section Diagrams Unrolled (120 micron intervals)*
microcuries per gram (!Ci/g) mRNA protein. initial learning in the supervised learning studies, – 30 min is – 15 min 30 m prior to OPB 15
Object-Image (http://simon.bio.uva.nl/object-image.html).
regularities in the environment Hippocampal
by the subfields
brain, citing
CA3, CA2, and CA1 were sampled at every section obtained through rostral the time of the alert observation behavior. This CA3 CA2 CA1
*Paxinos and Watson, The Rat Brain in Stereotaxic Coordinates, Fifth Edition, 2005
suggest “people continuously . . . recognize commentators would agree regarding what claims
patterns in the stream of sensations” and “detect Kant is actually making, nor on the implications of
potential content based on only a few aspects of the his views.9 In any event Kant’s account raises a
input (i.e. the gist)”, and that this results in “a number of interesting philosophical and
vague perception of coherence which is not epistemological issues. If feeling plays a role in
explicitly describable but instead embodied in a inductive judgments, what are the epistemological
‘gut feeling’ or an initial guess, which subsequently implications of such a view? If not, why does Kant
biases thought and inquiry.” In the gestalt closure describe “the principle of judgment” in both of
task, the authors found median orbito-frontal these ways? Does the inner sense need
cortex to be activated, as well as: the lateral portion development or training, and if so how? Kant
of the amygdala, anterior insula, and ventral actually raises this latter question: whether the
occipito-temporal regions. While the authors do common sense exists, or instead “must be
not focus on lateral amygdala, we find these results produced”; in this case it would be, in his view, a
quite encouraging with respect to the Kantian rational demand or requirement that we strive to
signal hypothesis. “produce such agreement in the way we sense.”
Kant’s account raises fascinating issues which we
4. Philosophical questions unfortunately have not space to address here.
9 Paul Guyer’s influential 1979 work Kant and the Claims of Taste argued Kant’s discussions of science and of aesthetic
judgment were unrelated, while Hannah Ginbsorg’s 1990 The Role of Taste in Kant’s Theory of Cognition argues they are
related but rejects a literal interpretation of “free play”. Many reject Kant’s identification of a common sense underlying
both taste and cognition. Also see Henry Allison’s Kant’s Theory of Taste (2001), the collection Aesthetics and Cognition
in Kant’s Critical Philosophy edited by Rebecca Kulka (2006), and Rachel Zuckert’s Kant on Beauty and Biology (2007).
KANT AND THE BRAIN 13
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