about the associations between a cell's peak spatial frequency ... of vision has arisen . In Eqs. (16) - (18), f is the frequency giving.
Young, R. A. (1982a) The Gaussian Derivative Theory of Spatial Vision. Unpublished manuscript. 161 pages.
theory
.....,is i o n
Department of
and Neuroscience Institute
Psycholo~y
University of Oregon, Eugene, Oregon 97403 ABSTRACT
This stAgas
paper
proposes a new theory of the
spatial
of'
visual system. in
fields
through
the
processing
receptive
primary visual system
least
at
cortex can be characterized as
of a. Caussian function. and
the
in
The theory maintains that
striate
derivatives
information
early
confirmed for
the
The theory
various
up
is
tested
on
cat
and
monkey
receptive fields found in the literature
and
a sample of monkey cortical cells.
found
I t is
that the shapes of receptiv.e fields along one spatial dimension can be completely characterized by just one number
-- a
positive
integer
Gaussian derivative term. derivatives previous
shown
a
a
significant
Fourier-like
just as Fourier to
ba
improvement
analysis of
~nalysis
had been
closer to the visual l
the
I t is shown that Caussian
characterizations of the visual
performing scene,
offer
representing
over
system the
as
visual
previously
mechanisms
for
spa.t ia.l
vision
derivatives
than
Gaussian
are a.lso shown to be superior to several
other functions tested, which
detectors.
bar
are
a
except for Gabor
close
approximation
functions,
to
Ca.ussian
derivatives. Extensions
of Gaussian derivative theory led to
a number of predictions, all of which were confirmed, about the associations between a cell's peak frequency, width,
bandwidth,
and
providing
underlying
Gaussian
such
a
derivative
close fit
to
spatial
receptive number.
field Besides
neurophysiological
data, Gaussian derivatives were also shown to predict human
psychophysical
results
better
than
other
proposed functions. In
short,
the
derivative
provides
a
receptive
fields and human psychophysical.
He
close
Gaussian
approximation
to
theory
single
~nit
channels.
believe the Gaussian derivative theory represents
another
small
step
in
the
mathematical
characterization of the visual system. Key
words:
Gaussian
derivative
theory,
spatial
vision, receptive fields, Fourier, bandwidth, cortex, psychophysics, spatial frequency.
2
~--·····-··--···~----------------------
• Gaussian Derivative Theory
Introduction
J:NTRODUCTJ:ON
.L.. .I!Ul fou.rj1r Tb•orx .2f. Yl1t9n Fouri•r an
area
analysis of the visual system has of
scientists,
owing
application visual
chtafly
to
to
the
visual
successful
fi•lds
and
human
of
psychophysical
Indeed, one fundamental sign of our period
b•an
the
oscillating Ke
interest
of this technique to the description
rec•ptiv•
channels. has
considerable
been
high
amplitude
wave
of
around the meaning of such applications.
see the phasing in of terms such as
"spatial
research
frequency
sensitivity"
tuning",
"ba.ndwtd.th",
and
to describe the spatial
"contrast
properties
of.
the visual system, as had. been done with the temporal properties
since Iv1s C1922>.
An impr•sstve body of
literature
jn support of the utility and
predictive
power of the Fourier analysis of tha spatial of vision has arisen .
The system Many
question of Fourier analysis
ll.x the
has bean a co-sign of this wave of visual scientists,
Jfestheimer
,
hav• 3
visual
research.
despite aarly_cautioning by adopted what we
call
th•
-
·----------
~-
--~--
I ntr od.l.lc ti on
Gaussian Derivative Theory
"strong form" of the Fourier theory of vision,
which
maintains that the visual system itself carries out a Fourier transform of visual space .
Campbell stated. that the
neurophysiological findings in cat and monkey suggest that
the
"spatial frequency content
is ..• extracted.
stated:
masking visual the is
11
11
of
the
image
In the psychophysical a.rena, .. These
summation,
experiments .•. provide
Gr•ham
adaptation,
evidence
that
system does do a crude Fourier analysis." weak form" of the Fourier hypothesis,
made that the visual system itself
frequencies,
but
only
uses
and the In
no claim. spatial
that the visual system to
a
first approximation can be treated. as a linear system and Fourier analysis provides a valuable tool for its study. The hypothesis that there are "multiple frequency and
Robson,
form 11 been
channels" 1969>
in tha visual system is associated with
of Fourier theory. precisely
spatial
specified
frequency
Although it has what the
channels" 4
the
means,
term as
spatial
otherwise
Fou.rier spectra of Y,,ussi&n deriy&\iv11 What Fourier the
most
do
Gaussian
space?
derivatives
look
like
This question is important
reliable data so far collected
on
in
because visual
cells has been in the Fourier domain, due to the ease of
producing
multiple
24
repetitions
with
drifting
Introdu.c ti on
Gaussian Derivative Theory
gratings.
If visual eel l receptive fields really a.re
Gaussian
derivatives,
then
we
shou.ld be
able
to
recognize their shapes in the cell data collected
so
far. For Fou.r i er
mathematical convenience in solving for the transform,
definition
of
G
it
is
useful
to
·slightly by adding a
the
alter in
tr
the
0
denominator, as in Eq. . G (x)
o
The
= -1 - e-x
/2cr
2
(13)
er l2iT
derivatives 0¥ Eq.
are given by Eq.
2
C13> with respect to
( 1'4 >.
'') - - = - - 1 H {x) G \.)(, n n cr o • n cr WC
The Eq.
Fourier· transform of Eq.
( 15). \
is given
by
Details are given in Campbell and Foster 2
p. 84, pairs 707.0 to 708.4 with
imaginary value "i" in Eq.
8=2•~
>. The
is dropped to obtain
the absolute magnitude of the Fourier spectrum, which is what we are most concerned with here.
25
Introduction
Gaussian Derivative Theory
(15)
A six
plot
of the Fourier transforms of the
Gaussian
derivatives as given by
Eq.
first
is
presented in Fig. 2. insert Fig. 2 about hare These height
transforms have all baen sat to the
and
centered
at
f
which
is
the
same peak
0
frequency.
It
derivatives
can
be seen
that
the
get progressively narrower
higher-order
and
in
the
bandwidths
Gaussian
derivative domain are also predicted to
completely
independent of the size of the th the n Cau.ssian derivative has
field
bandwidth regardless of'
fl".
be
receptive the
same
This independence from absolute size might
be
u.sefu.l for a higher level 33
analysis
which
Introduction
Gaussian Derivative Thaory
must somehow deal with changing receptive field sizes as a function of retinal eccentricity. level
logarithmic
conjunction
mechanism
with
a
Such a micro-
might
operate
macro-level
logarithmic
retinotopic mapping scheme as postulated by
in
to explain overall object
Schwartz constancy
with changing image sizes. In sum, wa have given a theoretical development of Gaussian derivatives which suggests that they have a
number of interesting properties which would
them
useful
machanisms
of
equations visual
if
they
derivative
Empirical and
spatial
vision.
Also,
normal various
wera developed which should hold true
cells
Gaussian
tools for helping to understand
make
themselvas
are
performing
analysis of the retinal
tests of these aquations were
tor
image.
then
·made,
will be presented in the Results section of this
paper.
ll.L.. O\htr Fynctlon1 Various derivatives
functions were
functions for vision.
other
examined
as
than candidate
Legendre,
basis
Functions rejected for various
reasons included the fallowing types: Laguerre,
Gaussian
Chebyshev, 34
Bessel,
Airy,
and sine functions.
Introdu.ct ion
Ga.u.ssia.n Derivative Theory
Yet, to
four functions other than sine waves ware found have
sufficiently close affinities to
fields to warrant further examination. we
could
safely
functions
reject
the
by inspection,
receptive
We found that
first
two
of
and the la.st two
these
required
empirical testing, &s did sine waves.
A... Pv&bol ic cxl ind1r .m:.
I in ear harmonic
osc i 1 lator
fu.nct ions The closely ha.ve
cylinder
a number of interesting features.
shape
which
linear
harmonic
oscillator
.to Harmuth .. Pulses of use the
'best',"
small pa.rt
percentage of their energy." shown
functions,
allied to the Gau.ssi&n derivative functions,
physics, as
parabolic
of
time-
and
of
the
the
they time-
cartai n
a.
Their orthogonality was 4 graphs the first
seven
para.bolic cylinder functions. insert Fig. 4 a.bout here Parabolic
cylinder 35
functions
also
have
the
Gau.ssian Derivative Theory
In tr odu.c ti on
int•restinq prop•rty that th•Y ar• th•ir own transforms
.
Fou.rier
Therefore,
one
0
wou.ld the if
expect inhibitory and excitatory sidebands Fou.rier domain as w•ll as in th• spatial
domain
receptive fields were in fact parabolic
fu.nctions.
Ind•ed,
have
reported
bean
also report two cortical
calls with secondary maxima in the frequency domain. As
mentioned,
idantic&l linear 1978>
239>. the thesa
to
h&rmon i c which
quantu.m
parabolic cylinder fu.nctions are
osc i 11 a tor
simpl~
e iqenfu.nc ti ans
so that no
frequencies
in
maintained video
the
neural
activity
raster
or
could
spike
regular train
due
responses to th• contaminate
to hertz
60
the
high
data.
by
masquerading a.s low frequencies, a.n artifact known a.s aliasing. To rad.u.ce the neura.l data. from the drH'ting gratings into a. meaningful form, th•
post-stimulus
programs
we
time
Fourier analysis of
histograms was
done
wrote to extra.ct th• amplitude
using of
the
fund.a.mental and the mean response. These measurements ware usually ma.de a.ta. number of spatial and
at a number al contrasts.
frequencies
To obtain sensitivity
functions, th• contra.st required to achieve a. certain criterion
response
amplitude 43
from
the
cell
was
Gaussian Derivative Theory
Methods
estimated
•Y••
Tha
;ave
the
by
reciprocal
;raphical curv•-f itting by of
this
contrast
level
sensitivity at that fr•quency.
lL. Hatbtat.ic&I
H1thpd.I
Unfortunat•ly, working
with
th•
math•m&tical
methods
for
Gaussian derivatives hav• not been
as
wall developed as with Fourier analysis. For example, no
computer
program
was
found
which
would
automatically estimate th• co•fficiants of a Gaussian der i vat i va sari es when step-by-step
r
procedur•
was unknown. was
Therefore
d•vised
which
a
worked
sat i sf actor i ly.
ill!. J.n .1bl. I pt. t. l I I Me
d OJI& i D
first wished to fit Gaussian derivatives
rec•ptiv• f ialds measured in the spatial order
to
test
the
validity
of
our
to
domain,
in
methods~
We
therefore us•d data in the literature which are known to
have been measured with a large number of
so
as to ensure reliability.
points
Me obtain•d the
by measuring the published curves
trials data
.
in
The
c e 11
number is given in the column marked "CELL". insert Tabla III about here As
can be saan from Fig.
9 and
Tabla
single Gaussian derivative term produced a close fit to in
onl~
ever~
III~
a
reasonabl~
receptive field measured.
Indeed,
9 cases out of 55 was there a fit in
which
less than 75% of the varianca was explained. The mean "variance cells
explained"
for the entire
sample
of
55 was
85.5% + 1.8% standard error.
At
the time the
anal~ses
57
of Fig.
9 were
done,
Gaussian Derivative Theory
the
G
term
Results
was not tasted for a fit to
the
data
0
because
of the common assumption in
the
literature
that cortical cells always have low spatial frequency fall-off , which G
does
0
not.
In hindsight, it app•ars plausible that sevara.l
of the cells in the G group in Fig. 9 and ·
in the Introduction enable a chart to be drawn 10>
a
which
illustrates
the
translation
between F and BW
negative
partialled
of
( 1962).
bacama
when the variable hald
,
partial correlation given
Technically
statistical it
= -.59>
Cr
out
technique
Results
speaking.,
an
was
via
the
in
McHemar as
a
which means that
stronger
even
tr
is acting
fT
"suppressor" variable,
counteracts
even
dee line
in
that is max Calls with relatively high f hidden in th• data. max should have relatively low tr by common assumption, logarithmic
but
bandwidth
with increasing f
low o 's result empirically in
logarithmic
as
well
relatively
as arithmetic
high
bandwidths
for
Thus the influence of a declining tr as
one
I
calls. to
goas
hidden peak
higher frequencies serves
t~
counteract
empirical very sharp decline in bandwidths as fraquancy
influence
of
constant>.,
a
then
increases. changing
If r
one
removes
is that it is
still unknown how the important cell
classifications
of X-like and Y-like , simple and complex, and
oriented
and non-oriented,
non-color opponent, derivatives.
nomenclatures and
ate.
are
These
associated
classical
have had tremendous
~enetits
characteristics
over
the
color opponent
cellular
predictive
years
in
with
power
explaining
the
that visual neurophysiologists
have
most easily noticed when they initially looked at the properties derivative
of
visual
theory
calls.
has not yet
Although been
Gaussian
considered
in
relation to these classifications, or for that matter to the detailed anatomy of Area 17 know
, there would be uncorrectable chromatic aberration affects that would cause two images -- one rad
and
one
blue
communication>.
,
Center and
Monasterio
mechanisms
F.
M.
of
macaque
1394-1417. surround
of opponent-color X and Y ganglion
cells
of retina of macaques. ;[,,:,. Meurophvs. 41, 1418-1434. De Monasterio properties
F. M. and Gouras
of
can
K.
Functional
ganglion cells of the
retina. ;[,,:,. Phvsiol., Land. De Valois
P.
K.
~,
rhesus
monkey
167-195.
Spatial frequency adaptation
enhance contrast sensitivity.
Vision
Res. ,
.!.Z,
1057-1065. De Valois (1979>
K.
K.,
De Valois
R. L.
and Yund
Responses of striate cortex cells to
and checkerboard patterns.
~
fh~siol
•.
E. W. grating
Land. 291,
483-505. De Valois (1978>
R. L., Albrecht
Cortical
cells:
113
D. C. Bar
and Thorell
detectors
or
L. C. spatial
Gua.ssia.n Derivative Theory
frequency
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.!:L._
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Physiol. 226,
... Gua.ssia.n Derivative Theory
Tyl er
w.
C.
.
independent
observed
agreement
frequency
and
trequ.ency
is
Gaussian
der iva.tive
generally
orderly progression ot symbols from
indicating
lower
number
observed
the
A scan column
peak
between
predicted
good:
The
the
observed down
any
indicates
frequencies
to
a.
those those
indicating higher observed trequ.encies .
147
Gaussian Derivative Thaory ,
Fig.
12.
Fourier
Figura Lag ends
parameters of a large sample
monkey cortical cells,
of
based on data from De Valois,
Albracht, and Thorell , plotted on a chart for
going from the Fourier domain into the
derivative domain, as in Fig. 10. the
solid
circles
Gaussian
The digits next to
represent the
number
of
cells
observed with the peak spatial frequencies and median bandwidths
indicated
respectively. over
cells
the
fall
at
each
deviation>.
bandwidth with increasing which De Valois
.!! A!·
as statistically significant.
open
Note
peak
the
spatial
cu
>
:;:: 0 4J
a::
1.00
o.eo
Ct
0
..J
0.0 -0.50..___________-.....-.---.---..-----.--.--....---..---. -0.14 -0.10 -0.06 -0.02 0.02 0.06 0.10 0.14
20
.,
"'c0
••
•
B
10
0. in 4J
a:: 0
•
0
.,"":s
z
-10
20 -1.4
-1.0
-0.6
-0.2
0.2
0.6
Spatial Position (Degrees)
7
1.0
1.4
Fourier Domain
A
Gaussian Domain
A*
BW 1.4
10
10
5
5
3 2
3 2
1
1
r· .3
.5
1
2
5
0 1
10
s* 10
5
5
3 2
3 2
1
1
BW 0.9
B
2
5
10
1
2
5
10
0 1
2
5
10
r-
.3
.5
c
1
2
o
5
c*
BW 0.8
10
10
5 3 2
5 3 2
1
.3
.5
1
2
5
Frequency (cpd)
Derivative no. ( Gn)
8
\
~
.,
54
•
30
•
42
•
45
• 38
>
36
• • ~
43
A
(/)
ZI
.:::: (l)
(./)
·~
17
57
20
0
44
49
6
50 If)
c
8
(l)
(/)
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