JOURNALOFN EUROPHYSIOLOGY
Vol. 46, No. 6, December 198 1. Printed in U.S. A.
Tactile Spatial Resolution. I.
Two-Point Discrimination, Gap Detection,
Grating Resolution, and Letter Recognition
KENNETH 0. JOHNSON AND JOHN
Sensory Processes Laboratory, Department
Parkville, 3052, Australia
SUMMARY AND CONCLUSIONS
1. The experiments reported here are part
of an investigation of the spatial neural
mechanisms underlying tactile sensation.
The first step in such an investigation must
be a broad, accurate characterization of tac-
tile spatial discrimination. More particu-
larly, the investigation requires specification
of the discrimination behavior that depends
strictly on spatial neural mechanisms. The
results of four experimental designs are re-
ported here. No single experiment, taken in
isolation, provides a basis for ascertaining
which aspects of spatial discrimination are
based on spatial information in the neural
discharge patterns rather than on intensive
or temporal information. Taken together,
the results of the four experiments provide
a consistent basis for examining the neural
mechanisms underlying tactile spatial dis-
crimination.
2. Experiment I, a modified two-point li-
men test, showed that subjects could reliably
discriminate between one and two OS-mm-
diameter points even when there was no sep-
aration between the two points. The result
demonstrated a high level of spatial resolu-
tion but discrimination may have been based
on any of a number of neural codes. The
contact area and the overall dimensions of
the two stimuli being compared were differ-
ent and, therefore, discrimination may have
been based on a difference in number of ac-
tive fibers, in total number of impulses, or
in the spatial patterning of the afferent dis-
charge evoked by the single- and double-
point stimuli.
0022-3077/8 1 /OOOO-0000$01.25 Copyright
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R. PHILLIPS
of Physiology, University of Melbourne,
3. In experiment II an attempt was made
to eliminate contact area and overall dimen-
sions as variables between the stimuli being
discriminated. Subjects were required to dis-
criminate between stimuli with and without
gaps. The overall dimensions of the stimuli
were constant and much larger than the
gaps. The d’ function was constant for small
gaps and then rose steeply (d’ = 0.86 for
gaps, g, ranging from 0.2 to 0.6 mm; d’
= 4.25(g - 0.55) for g > 0.7 mm). d’ is de-
fined here as the separation between two
unit-normal (a2 = 1 .O) distributions that
would, with an appropriate discrimination
boundary, produce discrimination judg-
ments with the same probabilities as those
observed experimentally. Gap size at thresh-
old (P = 0.75) equaled 0.87 mm. Some am-
biguity arises in the interpretation of these
results. Neurophysiological experiments have
shown that one major mechanoreceptive fi-
ber class is very sensitive to edges. Subjects
may have discriminated between stimuli on
the basis of the increased discharge evoked
by edges when gaps were present.
4. Experiment III was designed to hold
overall size, contact area, and edge content
constant between stimuli. Subjects were re-
quired to discriminate between square-wave
gratings (equal gaps and bars) that were
oriented along or across the axis of the fin-
ger. Subjects could not discriminate grating
orientation until the gaps exceeded 0.5 mm
(d’ = 0.0 for gaps < 0.5 mm). For gaps
greater than 0.5 mm (spatial periods > 1.O
mm) the d’ function rose steeply with a slope
of 4.0 SD units per millimeter gap size (d’
= 4.o(g - 0.5)). Gap size at threshold
0 198 1 The American Physiological Society 1177
 M. 0. JOHNSON AND J. R. PHILLIPS

(p = 0.75) equaled 0.84 mm. Neurophysio- stimuli are reported in the two following
logical studies have shown that some pri- papers (23, 24)
mary afferents are sensitive to grating ori- A key issue in the study reported here was
entation. Subjects may have discriminated to design experiments that would allow in-
the stimuli on the basis of differences in dis- ferences concerning the spatial neural mech-
charge rate evoked by the gratings with dif- anisms underlying tactual spatial discrimi-
ferent orientations. nation. The study of spatial discrimination
5. The results of each of the preceding requires the use of two or more stimuli that
experimental designs provided simple mea- are spatially different. The problem is that
sures of spatial resolution but they allowed it is difficult, if not impossible, to devise stim-
the possibility that they were based on non- uli that evoke different spatial patterns of
spatial neural cues. Experiment IV, a tactile neural activity but not different total num-
letter-recognition task, yielded results that bers of impulses or different temporal pat-
could not readily be explained by any non- terns in single fibers (when the stimulus is
spatial neural mechanisms. Subjects’ per- moved laterally). When that is so, how can
formance was as good or better than pre- it be inferred that one type of spatial dis-
dictions based on the results of experiments crimination (e.g., two-point discrimination
II and III, indicating that the results of those or grating resolution) is based on one of these
experiments were not based on some simple cues and not the other? A significant part
nonspatial mechanism. On the basis of in- of the study is concerned with this problem.
formation measures, X0-mm letters were The results of four experimental designs
discriminated at a level midway between are reported here. Each is subject to the dif-
chance and perfect discrimination behavior. ficulties referred to above but, together, they
provide a basis for separating the tactual
INTRODUCTION discrimination results based on spatial neural
mechanisms from those based on intensive
The fingertips and the fovea of the retina mechanisms.’ There are no possible tem-
are analogous in many ways. Each region is poral cues since the stimuli are stationary.
an anatomically specialized part of a two- The first three experiments use experimental
dimensional receptor sheet and each is the designs whose treatment is well developed
behaviorallv preferred site when maximum (16, 17) and they provide simple measures
spatial act&y is required. A fundamental of spatial discrimination. However, each de-
problem in tactile and visual neuroscience sign involves relatively few different stimuli
is to understand how spatial variables of a and training, which means that subjects
stimulus are represented in the afferent re- might have used intensive cues or spatial
sponse (the representation problem) and cues. The fourth design, a letter-recognition
how the afferent response is used in task, makes no attempt to control the inten-
spatial discrimination behavior (the coding sive cues; rather, it employs so many differ-
problem). ent geometrical patterns that it is not rea-
Investigation of the coding problem is sonable to hypothesize that they are classified
ideally based on a detailed map of discrim- ~-
ination behavior across a broad range of * Throughout the paper we distinguish between spa-
tial and intensive mechanisms (and cues, codes, etc.).
stimulus conditions coupled with detailed .A mechanism extracting information from the afferent
characterizations of the discharge patterns discharge patterns is purely intensive if it depends on
evoked by those stimuli. Any hypothesis con- the impulse rates in single fibers but makes no use of
cerning neural coding must then account for the exact locations of the afferent terminals relative to
each other or to the exact timing of impulses relative
a broad range of psychophysical results. This
to each other. A cue or code is purely intensive if it can
paper reports the results of four psycho- be extracted by such a mechanism. A mechanism is
physical experiments designed to measure purely spatial if it depends on the exact locations of
spatial resolution, which employ a variety of active neurons but is unaffected by uniform changes in
spatially configured stimuli applied to the impulse rate and makes no use of the detailed temporal
structure of the impulse patterns in single afferents. A
skin without lateral movement (hereafter code or cue is purely spatial if it can be extracted by
called stationary stimuli). Experiments deal- such a mechanism. A similar definition might be applied
ing with the neural representation of such to temporal mechanisms.

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 SPATIAL TOUCH: PSYCHOPHYSICS

accurately on the basis of some unidimen- bilities (with an appropriate, decision boundary).
sional intensive cue (22). The results of that The response bias is defined as the location of this
experiment, which are complex and yield no boundary (in normal deviates) relative to the
simple measure of spatial discrimination, midpoint between the two distributions used to
define d’ (16).
serve to distinguish which results in the first
In experiments I-III subjects were trained with
three experiments derive from intensive cues feedback until their thresholds (d’ = 1.35) stabi-
and which derive from spatial neural cues. lized: this required only 2 h for each of the subjects
in experiment 1, but about 20 h each for those in
METHODS experiments II and III.
EXPERIMENT I: TWO-POINT DISCRIMINA-
Experiments I-III TION. The stimuli (Fig. 1) consisted of OS-mm-
Each subject sat with his right index finger im- diameter steel pins with flat ends embedded singly
mobilized in a support that allowed stimuli to be or in pairs, near the edge of a large horizontal
applied to the distal finger pad from below. The disk that was rotated to bring the appropriate
stimuli were driven onto the skin by a linear motor stimuli into place. The disk was mounted on a
that allowed either force or displacement to be counterbalanced, pivoting cantilever that allowed
controlled (5). Between stimulus presentations the the whole disk to be raised and lowered. The un-
stimuli fell free of the skin so that the next stim- derside of the disk was supported by rollers. At
ulus could be positioned beneath the finger. The 8-s intervals the linear motor raised the counter-
period between stimuli and the stimulus duration balanced cantilever and disk (rise time, 50 ms;
in each experimental design were adjusted to duration, 3 s), pressing one or two pins onto the
achieve a comfortable stimulus-response rhythm finger pad with a controlled force ( IQ, 30, or 80
for the subject and to allow the experimenter time g). The sequence of two stimuli preceding each
to position the next stimulus, which was done judgment consisted of either S(l) (a pair of pins
manually. The stimulus apparatus and the sub- followed by a single pin) or SC*)(a single pin fol-
ject’s hand were shielded from his view, lowed by a pair). The pairs were aligned along
The experimental designs used in experiments the axis of the finger when brought into place and
I-III employed two possible stimulus sequences, were separated by gaps of 0.0,0.2,0.6, or 1.0 mm.
each containing two stimuli, and allowed two pos- The position of the finger was adjusted so that the
sible responses (16). After every second stimulus two pins of a pair contacted the skin as simulta-
in a train of stimuli, the subject was required to neously as possible. Contact time differences were
make a judgment. The sequence of two stimuli monitored electronically; they were generally less
preceding each judgment was one of two possible than 1 ms and they were never allowed to exceed
stimulus sequences (designated here as S(l) and 2 ms,
SC*)), which were presented in random order and
with equal probability. The order was taken from
a table that had been constructed prior to the
experiment using a random-number generator
The judgments were likewise restricted to one of
two possible responses (designated here as R”“and
R(*)), which were judgments that the two preced-
ing stimuli were presented in the S%equence or
the SC*)sequence. The raw data derived from the
experiments were the frequencies of R(l) and R(*)
following S(l) and SC*). These data are reported
here as sample estimates of the conditional prob-
abilities p( R(*) I St*)) and p( R(*) I SC”), which rep-
resent the frequencies with which the subjects re-
ported that SC*) occurred when, in fact, St*) and
S(l) were presented. All of the psychophysical
measures used here were derived from these two
sample probabilities, which fully define the sub-
jects’ behavior (16). The psychometric functions II I
are direct plots of p( R(*)IS(*)) and p( R(*)IS( ‘) ) ver- FIG. 1. Diagrams showing the relationship between
sus experimental variables. The behavioral sepa- the subjects’ right index finger and the stimuli employed
ration index, d’, is defined as the amount by which to measure tactile spatial resolution. I, two-point dis-
two unit-normal distributions would have to be crimination. 111, gap detection. III, grating resolution.
separated to produce the same response proba- IV, letter recognition (edge of letter shown).

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 K. 0. JOHNSON
EXPERIMENT II: GAP DETECTION. The stim-
ulus consisted of a 30-mm-diameter, 2-mm-thick
plastic disk that was pressed edgewise onto the
finger pad (Fig. 1) by the linear motor. Radial
slots (gaps), 2 mm deep, were cut at 15mm in-
tervals around the circumference, leaving a 30-
mm segment of the rim with no gap. Two such
disks were employed so that segments containing
gaps of 0.2,0.4, 0.6,0.8, 1.0, 1.5, or 2.0 mm could
be presented. The disk was rotated between trials
so that the rim segment below the finger contained
either a gap or no gap (i.e., the 30-mm segment
containing no slot). The rim of the disk was
pressed onto the skin for a period of 3.2 s once
every 10 s, reaching a skin displacement of 1,400
pm (indentation velocity, 755 pm. s-l; final re-
action force, 40 g average). The long axis of the
contact region between the rim and the skin was
colinear with the axis of the finger and was 2 mm
wide and 12 mm long overall. The sequence of
two stimuli preceding each judgment consisted of
either S(l) (a stimulus with a gap followed by a
stimulus without a gap) or SC2)(a stimulus with
no gap followed by one with a gap).
EXPERIMENTIII:GRATINGRESOLUTION. The
stimuli consisted of square-wave gratings cut into
the faces of 25-mm square plastic blocks that were
pressed onto the skin (Fig. 1) by the linear motor.
Gratings with spatial periods of 1.0, 1.5, 2.0, 2.5,
3.0, 4.0, and 5.0 mm were constructed by cutting
slots (depth, 1.5 mm; width, half the spatial pe-
riod) at regular intervals across the face of the
plastic block. The face of the grating lay at right
angles to the axis of the linear motor and was
mounted on a pivot so that it could be rotated
between stimulus presentations. The sequence of
two stimuli preceding each judgment consisted of
either S(‘)(the grating bars aligned along the axis
of the finger during both the first and second stim-
ulus presentations) or SC2)(the grating bars aligned
along the finger during the first presentation and
across the finger during the second presentation).
Indentation levels of 500,900, and 1,200 pm were
used (rise time, 50 ms; duration, 2.5 s; repetition
period, 7.0 s), and in some experiments vibration
at right angles to the face was superimposed on
the 900-pm step indentation for a period of 2.0 s
centered in the 2.5-s stimulus period.
Experiment IV
Experiment IV employed a pattern-identifica-
tion design using the 26 letters of the English al-
phabet. Subjects were told that the stimuli were
in the form of embossed, capital letters without
serifs. After each stimulus the subject was re-
quired to specify which letter had been presented.
No equivocal responses were allowed. The five
subjects who took part in this experiment received
no training or feedback. No improvement in per-
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AND J. R. PHILLIPS
formance was evident during the eight 2-h sessions
required of each subject.
Four sets of 26 sans serif capital letters (3.0,
4.5, 5.5, and 8.0 mm high) with 0.5-mm line
widths raised 1.5 mm above the background (so
that the skin would not contact the background)
were machined in plastic. The letter widths were
held as closely as possible to 60% of the height
although that was not always possible (for the
letter I, for example). The letters M and W were
slightly larger, having widths closer to 80% of the
height. These letters were fixed near the edge of
a large horizontal disk that could be rotated to
bring any letter beneath the subject’s finger (Fig.
1). The subject’s right index finger was fixed in
a support that allowed only vertical movements.
At 15-s intervals a light signaled the subject to
depress the letter beneath the finger and to hold
it down for a maximum of 4 s in order to identify
the letter. At contact the vertical dimension of
each letter was oriented along the proximodistal
axis of the finger pad. The disk was counterbal-
anced so that the embossed letter produced a con-
stant reaction force of 60 g. Disk displacement
was monitored on an oscilloscope and recorded
on a chart recorder. The mean time to steady
displacement (from 80 trials selected at random)
was 350 ms. Trials in which the subject failed to
hold a steady depression, which were rare, were
discounted and run again later.
RESULTS
Data were derived from experiments con-
ducted on the distal phalangeal pad of the
right index finger of 14 paid subjects (12
females and 2 males between 17 and 25 yr
of age). Subjects were not selected or re-
jected on the basis of performance.
Experiment I: two-point discrimination
This experiment tested the ability of sub-
jects to discriminate between single and dou-
ble-point stimuli (each point having a OS-
mm diameter). Note that this test differs
from the classical two-point limen test,
which determines the least two-point sepa-
ration at which the subject feels (has the
subjective impression of) two points. As de-
scribed in the METHODS section, one of two
stimulus sequences, S(l) = (two points fol-
lowed by one point) or SC2)= (one point fol-
lowed by two points), were presented and the
subject was required to respond with the
judgment that S(l) or SC2)was presented.
These responses were designated as R(l) and
RC2), respectively.
The mean performance of four subjects
 SPATIAL TOUCH: PSYCHOPHYSICS 1181

is illustrated in Fig. 2 where p(Rt2) I S(l)) and p( R12’1 S” ‘) p (R’2’( St2’)

p( RC2)I St2)) are plotted against two-point
separation for each contact force. It is not
obvious that any experimental variable, in-
cluding two-point separation, had a statis-
tically significant influence on performance;
the four subjects performed at a high level
under all conditions. Therefore, the raw data m subject BB
(4,460 stimulus-response pairs), consisting 0 II DF
of the sample probability p(Rt2)) for each
combination of subject (four subjects), se-
quence (SC’)and SC2)),point separation (0.0, Gap width (mm)
0.2,0.6, and 1 .O mm), and contact force ( 10, FIG. 3. Gap detection. Subjects’ ability to discrimi-
30, 80 g), were subjected to a four-way anal- nate between bars (12 x 2 mm) with and without central
ysis of variance. Only three sources of vari- gaps applied to the finger pad (1,400 pm indentation)
ation were significant at the 0.01 level; was assessed by presenting the stimuli in either of two
sequences: So) = a bar with a gap followed by a bar with
namely, sequence (which accounted for no gap, or S2) = a bar with no gap followed by a bar
96.2% of variation) and the interactive terms, with a gap* Subjects were required to judge which se-
sequence x point separation (1.4%) and se- quence had been presented ( Rt2) represents the judg-
quence x subject (0.7%). The effect of force ment that St2) was presented). The left and right sides
of the figure show the probability of Rt2) following So)
was not significant, either alone or as an in- and S(‘) versus the width of the gap. Data are for two .
teractive term. trained subjects.
These results demonstrate the exceptional
resolving capacity of the tactile mechanisms
subserving the skin of the distal pad but they most probably differed in total number of
provide no indication of the nature of those active fibers and total number of action po-
mechanisms. Because of the difference in tentials, as well as in spatial profile. Thus,
overall dimension1 and contact area between central neural mechanisms with no spatial
the single and double-point stimuli, the af- resolving capacity might have discriminated
ferent discharge evoked by the two stimuli between one and two points on the basis of
such intensive cues.
p( R12’( S” ‘) p ( Rt2’1 S12’) Experiment II: gap detection
1.0
p;-
In this experiment the overall dimensions
of the two stimuli being compared were held
constant. Subjects were tested for their abil-
ity to discriminate between stimuli (12 mm
--0.5
x 2 mm bars) with and without central gaps
A log (see Fig. 1 ), which were presented in each
n 309 possible sequential order: S(i) = (gap, no
l 80g
gap), S2) = (no pap, lyP)* The conditional
11.1 ,111
probabilities p( R 2, I SC1) and p( Rt2) I SC2))are
0.5 0.0 0.5 1.0
Gap separating two points (mm) plotted against gap size in Fig. 3 for two
subjects. The data are based on a total of
FIG. 2. Two-point discrimination. Subjects’ ability 3,240 stimulus-response pairs obtained after
to discriminate between one and two OS-mm-diameter training was completed. Unlike the first ex-
points applied to the finger pad was assessed by pre- periment, subjects’ responses were less ac-
senting the stimuli in either of two sequences: S(l) = two
points followed by a single point, or S(‘) = a single point curate at small gap sizes, although they per-
followed by two points. Subjects were required to judge formed more reliably (p = 0.66) than chance
which sequence had been presented (Rt2) represents the at the smallest gap size (g = 0.2 mm). The
judgment that St2) was presented). The left and right threshold, based on the traditional criterion
sides of the figure show the probability of R(2)following
S(l) and Sc2) versus the gap separating the two points. of 75% correct judgments (d’ = 1.35), was
Performance (mean of four subjects) at three contact 0.87 mm. However, the threshold is not a
forces ( 10, 30, and 80 g) is illustrated. sufficient characterization of the results

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1182 K. 0. JOHNSON AND J. R. PHILLIPS

since the response probabilities were not de- tensive rather than spatial information.
scribed by any simple relationship such as Comparison of the results of this experiment
a cumulative normal function. The manner with the results of experiments III and TV
in which subjects’ behavior varied with gap will indicate that the flat section of the d’
size in the vicinity of threshold is illustrated curve was based on intensive cues, whereas
in Fig. 4. The separation index, d’, is roughly the rising phase was based on spatial infor-
independent of gap size at small gaps and mation in the afferent discharge,
then rises rapidly; the data are approximated
by two linear segments Experiment III: grating resolution
d’ = 0.86 g < 0.7 mm In this experiment the overall dimensions,
the contact area, and the edge content were
= 4.25(g - 0.55) g > 0.7 mm (0 held constant for the two stimuli being dis-
where g represents the gap size in millime- criminated. Subjects were required to judge
ters. The subjects displayed a slight bias to- whether two gratings with the same period,
ward the judgment Rc2) (mean bias = 0. I6 presented sequentially, were presented with
SD units). the same alignment, S(l), or with orthogonal
Although the overall dimensions of the alignments, S2! Two grating-resolution
stimuli with and without gaps were held con- studies were conducted. In the first study the
stant, it cannot be inferred with certainty gratings were stepped onto the skin and he1
that the discrimination performance was stationary, which preferentially ges th
based on the ability of central mechanisms slowly adapting afferent fibers In the
to use spatial information in the afferent second study the gratings were vibrated at
discharge. The gaps introduced edges, which 40 and 200 Hz to maximize the contribu-
probably increased the total afferent dis- tions of the quickly adapting afferents (at
charge (23) and they modified the total con- 40 Hz) and the Pacinian afferents (at
tact area, which may also have altered the 200 Hz).
total afferent discharge (see DISCUSSION The performance of each of the three sub-=
Thus, stimuli with and without gaps might jects who took part in this experiment was
have been discriminated on the basis of in- assessed by computing the conditional prob-
abilities p( R(2)IS(1)) and p( Rc2)IS2)) for each
2.5 of the six grating periods at three skin in-
dentations (500, 90 Y and 1,200 pm). The
a
1 individual data for the three subjects relating
2.0
I B to the stimulus intensity of 900-pm skin in-
dentation are illustrated in Fig. 5, Unlike the
previous two experiments, subjects displayed
1.5 /
the full range of performance from chance
to perfect discrimination over the range of
8 /
the primary stimulus variable (
d I /
riod), As in experiments I and 1
1.0 q / 0 little intersubject variability or response
m
- 8
n- - - - l-
AL- bias (mean bias toward Rc2) for three sub-
0 / jects = 0.026 SD units). In Fig. 6, d’ (mean
0.5 i / for three subjects) is plotted versus grating
/ period for the three indentation levels (data
. BE4 base = 5,200 stimulus-response pairs ob-
/ * DF tained after training was completed). It is
0.0 Lr #in
0.0 0.2 0.4 0.6
I
0.8 1.0
, apparent that increasing the grating period
Gap width (mm)
above about LO mm results in a near-linear
increase in d’, particularly at 900- and 1,200-
FIG. 4. Gap detection-behavioral separation index pm indentation Increasing indentation ap-
(d’) versus gap width. Data points represent the indi-
vidual performances of two subjects. Dashed lines rep- pears to lead to an increase in performance
resent a piecewise linear approximation of their behavior at all grating periods. However, the d’ curves
(see text). intercept the abscissa at a grating period of

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 SPATIAL TOUCH: PSYCHOPHYSICS

p( R’*‘l S” ‘) p( R’*‘l S’*‘) of gratings with periods below about 1.0 mm.
Similarity between the gap detection results
and these results can be seen by expressing
these results in terms of the gap between the
bars rather than the grating periods; then,
the results for 900- and 1,200-pm indenta-
tions are closely approximated by
$’ = 0.0 g < 0.5
= 4.O(g - 0.5) g > 0.5 (2)

Grating period (mm) where g represents the gaps in the gratings

FIG. 5. Grating resolution. Subjects ability to resolve (half the grating period). The threshold eval-
grating stimuli applied to the finger pad was assessed uated at 75% correct judgments (d’ = 1.35)
by applying a grating to the skin twice in succession in for 900 and 1,200 pm corresponded to gaps
either of two sequences: S(l) = the grating bars aligned of 0.92 and 0.84 mm, respectively. The rising
along the finger axis in both the first and second pre- limb of the d’ functions obtained for exper-
sentations, or S(*) = the grating bars aligned along and
then across the finger axis. Subjects were required to iments II and III are very similar (compare
judge which sequence had been presented (R(*) repre- equations 1 and 2).
sents the judgment that S(*)was presented). The left and The grating-resolution experiments were
right sides of the figure show the proability of R(*) fol- repeated at 900-pm indentation, but with
lowing S’) and S(*) versus grating period. The experi-
ment was performed using three levels of skin inden- superimposed vibration normal to the skin
tation (500, 900, and 1,200 ,um). The individual surface at the best tuning frequencies for
performances of three subjects corresponding to 900- cutaneous quickly adapting and Pacinian
pm skin indentation are illustrated. afferents (27). Stimuli designed to engage
the two rapidly adapting mechanoreceptive
about 1.0 mm for all indentation levels, im- afferents had no important effect on the sub-
plying th at no in formation is available to the jects’ performance. Vibration at 200 Hz and
decision process concerning the orien tation IO-pm amplitude had no effect; i.e., the d’
curve was virtually identical to the d’ curve
obtained without vibration. Vibration at 40
Hz, 200 pm and at 200 Hz, 35 ,um raised
the d’ function slightly at 1.0 mm and de-
pressed it at higher frequencies. Under these
conditions (40 Hz, 200 pm and 200 Hz, 35
pm), the d’ functions were closely approxi-
mated by d’ = 0.5 -!- 3.l(g - 0.5) for gaps
ranging from 0.5 to 1.5 mm (i.e., grating
periods from 1.0 to 3.0 mm).
although the overall dimensions, the con-
tact area, and the edge content were held
constant for the two stimuli being discrim-
inated in the grating-orientation test it can-
not be inferred with certainty that the sub-
jects’ responses were based on spatial
structure in the afferent discharge when the
results of this experiment are considered in
isolation. Neurophysiological experiments
1.0 2.0
Grating period (mm) (23) carried out in parallel with the psycho-
physical experiments showed that the dis-
FIG. 6. Grating resolution-behavioral separation charge rates of some single afferents are
index (d’) versus grating period. Each curve represents
the performance (mean of three subjects) for one of the greater when the gratings are oriented along
three indentation levels (500, 900, and 1,200 pm) used the dermal ridges than when they are ori-
in the experiment. ented across the ridges.

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 K. 0. JOHNSON
Experiment IV: letter recognition
The problems associated with interpreta-
tion of the data derived from experiments
I-III point to an intrinsic difficulty in the
general experimental design employed in
those experiments. They involve a small
number of stimuli presented many times in
succession, usually with training and feed-
back. When the discriminations are difficult
the subject may use neural information (if
it is available) other than that which the
psychophysical experiment is designed to
test. Thus, rather than try to devise a still
more carefully controlled two-sequence x
two-response design to eliminate the possi-
bility of spurious cues, a different design
strategy was used. If a subject accurately
identifies a spatial pattern that is one of
many possible patterns without feedback or
prior training, then it is unlikely that the
identifications are based on some nonspatial
aspect of the afferent discharge. That ratio-
nale underlies the letter-recognition experi-
ment.
Subjects were required to identify raised
capital letters by touching them without lat-
eral movement, as described in the METHODS
section. Letters with stroke widths of 0.5 mm
and heights of 3.0,4.5, 5.5, and 8.0 mm were
used. Subjects were given no training or
feedback. The raw data consisted of a 26
x 26 confusion matrix for each of five sub-
jects and four letter sizes. Figure 7 illustrates
a pooled confusion matrix representing the
responses of all subjects to 4.5-mm letters.
The entries represent the number of times
that each stimulus evoked each possible re-
sponse (e.g., the stimulus letter Y was pre-
sented 52 times and evoked the response T
24 times and Y 16 times).
The probability of correct identification
(mean over all 26 letters) versus letter height
for each of the five subjects is shown in Fig.
8A. Three aspects of these data are notable.
First, as in the previous experiments, the
performance of all subjects was similar. Sec-
ond, the subjects performed at levels well
above chance (the probability of correct
identification by chance was p = 0.038; i.e.,
l/26) at all letter heights. Third, the rela-
tionship between recognition accuracy and
letter height was linear over the range tested,
suggesting that subiects would have ner-
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AND J. R. PHILLIPS
formed significantly better than chance for
letters smaller than 3.0 mm. Furthermore,
this analysis, based on frequency of correct
identifications, underrepresents the subjects’
performance, as can be seen in the confusion
matrix in Fig. 7. Erroneous judgments were
predominantly restricted to a small number
of alternatives. This consistency is illustrated
in Fig. 8B where the curve labeled C rep-
resents the mean frequency of correct judg-
ments at each letter size over all subjects and
letters; C + 1 represents correct judgments,
C, plus the single most-frequent distractor
for each stimulus (e.g., T for V); and C
+ 2 represents C + 1 plus the second most-
frequent distractor (e.g., Y for V). Almost
60% of responses to 3.0-mm letters were con-
fined to the correct response plus the two
most frequent distracters. The logical exten-
sion of this approach is an information the-
oretic analysis (2), which assesses consis-
tency in the stimulus-response matrix rather
than accuracy.
In this analysis the subject was regarded
as a communication channel between a
sender (the experimenter presenting letters
in random order) and a hypothetical receiver
listening to the subject’s responses. Before
the subject’s response the receiver’s uncer-
tainty concerning the stimulus is 4.7 bits
(log, 26). After the response the possibilities
are narrowed (assuming that the receiver
knows the subject’s response probabilities)
and his uncertainty is reduced. The formulas
used to calculate the mean information
transfer per response are given in the AP-
PENDIX. The performance measured as in-
formation transfer (Fig. SC) is much higher
than that measured simply by the frequency
of correct judgments (Fig. 8A). That can be
seen by calculating the information mea-
sures that would have resulted if the errors
had been randomly (uniformly) distributed
among the 25 alternatives rather than con-
sistently allocated to a few distracters. The
curve labeled C (mean of five subjects) in
Fig. 80 was constructed by redistributing
the erroneous responses uniformly and re-
calculating the information measures. The
curve labeled I represents the mean of the
curves illustrated in Fig. 8C. At the smaller
letter sizes the consistent but erroneous re-
sponses accounted for most of the informa-
tion transfer.
 SPATIAL TOUCH: PSYCHOPHYSICS 1185

RESPONSE

ABCDEFGHIJKLMNOPQRSTUVWXYZ
A 40 1 .12 4. 1 2 1 52
B . 13414li3:..... 32i:36.:i::::. 52
C 3 522 14 211 1 3 52
D .8 28 2 i: : i i 3 2 52
E 2 i 23 !i i . 1 3 1 i 1 . i 1 1 1 2 52
F i i 130 . il . . li, 5 52
G li,i 5 2. i 1 9 2 3 4 5 i 52
t-l i3.23.6 9 : * 1 3 ii . 1 3 2 2 3 2 52
I . 52 . . . 52
J . 5i . 52
K i : i : ii i i i .l 9 1 5 3 6 52
ZL . . . l . . . . 149 .2 . . 52
L M li : 3 .2; i : : . i : i : : : 52
r N ii:ik:i6.i2.7u ii ' .l 3 2 . . 52
* 0 1715.13.. 3Oi ii . i 52
P 1 .4 i ;i:h32:21i..:::: 52
Q i i il ii:. I 61514 321... 52
R 322 i-41 i . i 8 1 513 2 . . i : l 52
S 28lizi4.:i....i23614 iI... 52
T . . 1 . 2 1. 1 4; 52
U .i. 8. i ii .i : : ij;;l;zzi*;::: 52
V . . . . .2. i . 1 . 1 4 . . 210 4lic 9 . 52
W . . l . . i i 2i 12 224 i 1 52
X . . . .S: 2::3ill.::ii:z 1 12 . 1s 52
Y . . 3i 2 . . 24 . i .16 1 52
Z . .i 1 i, .i 1: ii ..i : 1 : : ii . . : 7 1 30 52
56 69 46 70 79 53 50 39 56 62 33 58 40 58 69 58 19 55 45 92 53 26 43 31 30 62 1352

FIG. 7. Letter recognition-confusion matrix. Subjects were required to identify raised (embossed) capital letters
applied to the finger pad (application force 60 g) at four letter sizes (3.0-, 4.5-, 5.5-, and 8.0-mm letter height).
The confusion matrix illustrated relates to 4.5-mm letters; it was obtained by pooling data for five subjects. Each
entry indicates the number of times that the stimulus Si evoked the response Rj. Stimuli were presented in
pseudorandom order a total of 52 times each.

A close examination of the distracters as- letters (I, J, S, and R) fell outside this simple
sociated with each stimulus letter confirms grouping. I and J were identified incorrectly
the initial assumption that the subjects’ be- on only 1 and 5 of 200 trials, respectively,
havior in the letter-recognition task depends and thus they could not be placed in any
on spatial information in the afferent dis- cluster. The most-frequent distracters for R
charge and not some simple measure of the and S were N, E, A and B, G, respectively.
response like the total impulse rate. The As can be seen in the groupings, the most-
most-frequent distracters associated with frequent distracters are those letters that are
each stimulus letter were obtained by pool- closest in spatial form to the stimulus, in-
ing the confusion matrices (each letter was dicating that the subjects’ responses were
presented 200 times). That analysis is almost based on the spatial form of the neural rep-
completely summarized by the sequential resentation.
groupings BDOQGCE-KZX, HNM WU,
PF-TVY, and AL; the dash is meant to sig- Comparision of performance in
nify that there is significant confusion be- experiments II, III, and IV
tween letters of the two groups, but that the If the results of experiments II, III, and
confusion is not as consistent as the confu- IV all represent the human ability to resolve
sion within groups. With one exception, G, spatial components of tactual stimuli, then
the most-frequent distracters for each letter the results of the three experiments should
are given by the two neighboring letters in be related in a systematic way. Systems with
its group. The two most-frequent distracters limited spatial resolution have long been ap-
for G were 0 and B, which are found in the proximated by mosaics of elements whose
same group but are not neighbors. Only four dimensions represent the smallest resolved

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1186 K. 0. JOHNSON AND J. IX. PHILLIPS

0001 2 3 4 5 6 7 8

Letter height (mm)

FIG. 8. Letter recognition. A: probability of correct response (mean over 26 letters) versus letter height for five
subjects. Guessing behavior would have resulted in 3.85% correct responses. B: curve C represents the mean
frequency of correct responses over all subjects and letters versus letter height. Curve C + 1 represents the frequency
of responses that were correct or confined to the single most-consistent alternative for each letter (e.g., T for Y).
Curve C + 2 represents C + 1 plus the second most-consistent alternative for each letter (e.g., V for Y). C:
average information conveyed by subjects’ responses for each of the five subjects versus letter height (see text).
D: curve I represents the mean information transferred per response over all subjects versus letter height. Curve
C represents the information measure that would have resulted if there had been no consistency in subjects erroneous
responses (see text).

components (13), such as the gaps in exper-
iment II. On this basis gratings should be
resolved when the grating period approxi-
mates two unit spacings, allowing interunit
modulation to represent the peaks and
troughs. Such a mosaic only begins to rep-
resent letters at somewhat larger dimen-
sions. A 3 x 3 array may allow I, .I, and L
to be represented effectively but most letters
require at least 5 x 5 resolution units to re-
move geometric ambiguities (e.g., consider
a dot-matrix display of the letters B, E, M,
R, S, and W). Based on this simple analysis
it would be expected that the gaps, grating
periods, and letter heights would fall ap-
proximately in the ratio 1:2:5 when com-
pared at a common level of performance.
The results of the three experiments are rep-
resented in Fig. 9 in terms of the frequencies
of correct responses. Abscissa values in Fig.
9 represent dimensions of the stimuli (i.e.,
gap width, grating period, and letter height).
The dashed curves represent the expected
performance in the gap-detection and letter-
recognition experiments relative to the grat-
ing-resolution results. The abscissa values of
the dashed curves are scaled in the ratios 1:2
and 5:2. As can be seen, the results of the
three experiments lie in approximately the
expected ratios. It might be argued that the
letter-recognition results should have been
better than the 5:2 ratio because some letters
can be resolved by 3 x 3 arrays. As dem-
onstrated in Fig. 8, the probability of correct
identification underestimates subjects’ per-
formance in experiment IV. When the re-
sults of all three experiments are compared
using an information theoretic measure (bits

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 SPATIAL TOUCH: PSYCHOPHYSICS 1187

1.0 3
&
0.8 g
G
0.6

CE
-.-.- o, 0.6
-
%

0 1 2 3 4 5 6 7 8 9 10
Stimulus dimension (mm)
( gap width, grating period, letter height )
FIG. 9. Comparison of the results of experiments II, III, and IV. Points and solid lines represent results of the
three experiments (measured as probability of correct response) plotted versus the stimulus dimensions: i.e., gap
width (II), grating period (III), and letter height (IV). The dashed curves repeat results of experiment III with
the stimulus dimensions scaled in the ratio 1:2 and 5:2. They represent results for gap detection and letter recognition
that would be expected if the gaps, grating periods, and letter heights had fallen in the ratio 1:2:5 at comparable
levels of performance (see text).

per response) the letter-recognition results
lie in the range 3:2-4:2 relative to the grat-
ing-resolution results. On this basis, the let-
ter-recognition results are somewhat better
than predicted.
The consistency of the results from ex-
periments II, III, and IV indicate that they
all measure that facet of tactile spatial res-
olution that is based on spatial modulation
of the afferent discharge. No other hypoth-
esis accounts for the consistency of the data.
DISCUSSION
The psychophysical experiments reported
in this paper were conducted in order to pro-
vide behavioral data necessary for testing
hypotheses concerning neural coding of the
spatial details of tactile stimuli. When mov-
ing stimuli are employed a large number of
hypotheses are available ( 18), partly because
of the possibility of coding spatial informa-
tion in the temporal discharge profiles of
mechanoreceptive afferents. The use of sta-
tionary stimuli, as in this study, limits the
number of possible coding procedures to two
main types: I) discrimination between spa-
tially different stimuli may be based on the
difference in form of the spatial neural pro-
files representing the stimuli (i.e., a spatial
code), or 2) it may be based .on some dif-
ference in the magnitudes of the responses
to the stimuli (i.e., an intensive code). In-
cluded in this latter category is any aspect
of the afferent discharge that might be used
by a central mechanism with little or no spa-
tial resolution (e.g., the total number of ac-
tive fibers, the total number of action poten-
tials, the peak discharge rate, etc.).
The experiments reported in the RESULTS
attempt to define which coding procedure is
actually employed under various conditions
and to measure the limit of resolution when
a spatial code is used. Taken together, the
four experiments provide a broad, consistent
description of the spatial resolving capacity
of the tactile system for stationary stimuli.
In experiment I subjects discriminated
two OS-mm-diameter points applied to the
finger pad from one point with near cer-
tainty, even when there was no separation
between the two points. This result is similar
to Friedline’s (8) observations made on the
forearm (see later); both results imply that
the classical two-point limen is not a mea-
sure of the limits of spatial resolution (dis-
cussed later). However, little can be inferred
from the results of experiment I concerning
neural mechanisms when the results are con-
sidered in isolation. The results of experi-

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 K. 0. JOHNSON
ments II, III, and IV suggest that the sub-
jects might have been able to discriminate
two OS-mm points from one point on the
basis of a spatial code but with extreme dif-
ficulty; the ease with which they made the
discriminations suggests that they used in-
tensive cues. The tactile system is very sen-
sitive to differences in area and length (19,
3 1), which may well be based on intensive
information in the manner proposed by Hol-
den (14) to explain the classical two-point
limen.
The second experiment is, we believe, an
example of two neural codes underlying dis-
crimination behavior in one task. At 0.2-,
0.4-, and 0.6-mm gaps the subjects’ discrim-
ination performance was constant (d’ = 0.86)
and significantly better than chance behav-
ior. The following paper (23) shows that the
slowly adapting afferents are very sensitive
to edges, which provides at least one expla-
nation for this behavior; that is, that the
edges produced increased discharge rates,
which were used to discriminate stimuli with
gaps from stimuli without gaps. Increases in
gap size above 0.6 mm produced improved
performance, which might be accounted for
by some nonspatial mechanism but the sim-
ilarity between the gap-detection and grat-
ing-resolution results suggests that they were
both based on the same mechanism.
The grating-resolution results are, we be-
lieve, the key results. They were derived
from a two-sequence by two-response ex-
perimental design, which is supported by a
well-developed theoretical framework ( 16)
and they yielded simple measures of spatial
resolution. Of the experiments using this
design (I, II, and III), the grating-resolution
task included the most extensive controls for
eliminating intensive cues. Furthermore,
comparison of the results of experiments II,
III, and IV confirmed the hypothesis that
the results of experiment III were based on
spatial neural cues.
One possible ambiguity in the interpre-
tation of the grating-resolution results is that
the change in grating orientation relative to
the skin ridges caused differences in the af-
ferent impulse rates, which may have been
used in the discrimination task. The primary
.
evidence that this did not occur was the con-
sistency between the grating-resolution and
letter-recognition results (see Fig. 9) where
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AND J. R. PHILLIPS
explanations based primarily on intensive
cues are not tenable. Also, while the effect
of orientation between the bars and the skin
ridges on the discharge rates is significant
(23), the circular patterning of the ridges on
the human fingertip provide a mitigating
effect. Thus, the evidence indicates that the
grating-resolution data provide a measure
of the component of spatial tactual discrim-
ination that is based on spatial structure in
the afferent discharge.
The data indicate that for grating periods
less than 1.0 mm no spatial neural infor-
mation reaches the final discrimination
mechanisms. At spatial periods greater than
1.0 mm the behavioral separation index, d’,
rose rapidly, indicating the rapid develop-
ment of spatial information in the afferent
discharge, which was preserved by the cen-
tral pathways.
Comparison of visual and tactual
spatial resolution
The letter-recognition results provide a
basis for comparing visual and tactual spa-
tial discrimination. Confusion matrices rep-
resenting the performance of subjects in let-
ter-recognition tasks provide detailed maps
of spatial-discrimination behavior over a
wide range of geometric forms and they have
been obtained for vision (11, 12, 30) and
touch (6, 21; this study). Unless the same
fonts are used and the confusions are in-
duced in the same way, it is difficult to com-
pare data. Confusion in letter recognition
has been induced bY reducing the let ter size
(12; this study ), the exposure time (6, 11,
30), and by presenting blurred images (1).
The most similar published visual and tac-
tual data are those of Gilmore et al. (11)
and Craig (6) who used dot arrays and short
exposure times. The results of their experi-
ments were very similar, at least as measured
by the product-moment correlation of fre-
quencies of correct judgments (r = 0.88).
The visual letter-recognition study that is
most like the one presented here and that
provides a basis for comparing spatial res-
olution in the two systems is the study by
Hartridge and Owen (12) who used visual
letters with heights ranging from 1.4 to 3.4’
of arc. A scaling factor of 2.0-2.5 mm/min
of arc is required to obtain overlap between
their psychometric function (percent correct
 SPATIAL TOUCH:
versus letter height) and the ones presented
here (Fig. 8A). The imprecision in the ap-
propriate scaling factor arises because the
visual psychometric function was steeper
than the tactual function. Work of our own
(unpublished observations) using visual and
tactual letter-recognition tests in single sub-
jects verified this comparison. All the data
suggest that insofar as spatial neural mech-
anisms are concerned, 2.0-2.5 mm on the
skin of the distal phalangeal pad is equiva-
lent to 1.O’ of arc in the region of maximum
visual acuity.
This dimensional equivalence corresponds
closely to the receptor spacings in the skin
and the retina, implying that the relation-
ships between receptor spacing and resolu-
tion limits are also similar in the two sys-
tems. The mean center-to-center spacing of
cones in the fovea is approximately 2.0 pm
(25) which corresponds approximately to
0.45’ of arc. The innervation density for the
distal finger pad is much more difficult to
assess but two studies in man and monkey
(7, 15) estimate the mean center-to-center
spacing for both slowly and quickly adapting
afferents at aproximately 1.0 mm, which
yields a ratio of 2.2 mm/min arc (i.e., 1.0
mm/0.45’ arc). The letter heights yielding
50% correct responses (approximately 2.0’
of arc in the visual task (12) and 5.0 mm
in the tactual task, see Fig. 8) span approx-
imately five receptors in each system. Thus,
in both systems the limits of spatial resolu-
tion and of pattern recognition seem to be
set by the peripheral receptor spacing in the
regions of highest acuity; thus, those path-
ways that lead to spatial pattern recognition
must be characterized by optimum or near-
optimum spatial processing.
Classical two-point limen
It is not clear how the spatial discrimi-
nation results reported here are related to
the classical two-point limen devised by
Weber in the 19th century (26). In the clas-
sical test the separation of a pair of blunt
compass points is adjusted until the subject
reports that he feels two points; that sepa-
ration is usually 2-3 mm on the fingertips,
20 mm on the forearm, and 40 mm on the
back (32). It appears that the two-point li-
men is correlated with the relative spatial
acuities of these skin regions but it is not
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PSYCHOPHYSICS 1189
clear what is being measured at any one lo-
cation. For example, it does not represent a
subject’s ability to discriminate two points
from one point. That was demonstrated by
Friedline (8) when she showed that subjects
could reliably discriminate two points sep-
arated by 2 mm from two points with no
separation on the forearm where the clas-
sical two-point limen is 20 mm. The results
of experiment I showed that on the fingertip,
as on the forearm, subjects’ abilities to make
discriminative judgments involving punctate
stimuli are much finer than is indicated by
the two-point limen. Thus, in the classical
experiment subjects do not say that the stim-
ulus feels like two points until the separation
far exceeds a level at which they are objec-
tively able to discriminate two points from
one point. If that is so, subjects should be
able to subclassify and discriminate two-
point separations between the absolute
threshold (which was not reached in the ex-
periment reported here-the 0.5-mm pins
were too coarse) and the classical two-point
limen. Indeed, the earliest investigations of
tactual discrimination using punctate stimuli
(10, 20, 28) showed that subjects could re-
liably categorize two-point separations be-
low the classical two-point limen.
What do these observations mean in terms
of the underlying mechanisms? We believe
that this question can be answered by re-
phrasing the hypotheses of early investiga-
tors such as Titchener (29) and Boring (3)
in terms of the likely central neural repre-
sentation of two points (9) and the results
of this study. To quote Boring (Ref. 4, p.
482), “It means that there is a series of dif-
ferentiated perceptual patterns, that the cri-
terion for two must be placed at some critical
point in the series, and that the meaning of
the judgment two is indeterminate unless the
criterion has been established.” Paralleling
this series of perceptual patterns is a series
of neural patterns, all different in spatial
form and intensity. At the lowest discrimi-
nable levels the spatial differences are al-
most certainly too small to be discriminated;
therefore, the subject uses some intensive
aspect of the response and places his crite-
rion somewhere along that dimension. Then,
at separations of the order of 0.5 mm on the
fingertip he begins to resolve the spatial dif-
ferences. Finally, at 2- to 3-mm separation,
 K. 0. JOHNSON AND J. R. PHILLIPS

which is well above his limit of resolution, where p(Si I RJ represents the probability that the
the spatial neural patterns become suffi- ith stimulus &as presented given that the subject
ciently bimodal to satisfy the subject’s cri- gave the jth response, which is estimated by nij/
terion for the statement “the stimulus feels Ilj = entry in ith row and jth column of the
(Ilij

like (evokes the subjective sensation of) two Confusion matrix, nj = jth Column total)* The
uncertainty reduction following Rj (i.e., 4.7
points.” Thus, the two-point limen should be
- HWR,)) re p resents the information transfer
regarded as a boundary on a categorical
(1)
scale and not as a threshold of spatial acuity.
I = 5 p(Rj)(4.7 - H(S 1R.i))
APPENDIX i=l

Information theory measures used in = 4.7 - 2 El H(SIRj)

letter-recogn ition analysis i=l N (A3
The subject is regarded as a less-than-re-
liable communication channel between the in which p(Rj) represents the proability of Rj and
experimenter who is selecting letters for pre- N represents the total number of stimulus-re-
sentation to the communication channel and sponse pairs.
a receiver who is listening to the subject’s
responses. Before the subject responds, the ACKNOWLEDGMENTS
receiver only knows that the letters are
equally likely and his uncertainty concerning We thank Lyn Wood and Gloria Kruba who prepared
the typescript.
which letter was selected is given by H(S)
The work was supported by National Health and
= log, 26 (see Ref. 2 j. Following a particular Medical Research Council of Australia grants.
response, Rj, the receiver’s uncertainty con-
cerning any particular stimulus, Present address of IS. 0. Johnson: Dept. of Physiol-
by H(Si I Ri) = - log, p(Si I Rj). His average ogy, The Johns Hopkins School of Medicine, Baltimore,
MD 21205.
uncertainty concerning all stimuli is given
Present address of J* R. Phillips: Physiological Lab-
by oratory, Downing St., Cambridge CB2 3EG, England.
H(S I Rj) = 5 p(Si I Ri) log, p(Si I Rj)
i=l

Received 23 December 1980; accepted in final form

(AI) 30 June 1981.

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