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BIOPHYSICS
ELECTRORFCERTORS
There
aro
throe basic tyres
of lateral line organs.
The first
are
the
: ordinary LL0,
second
and first order neurons canned with micro-
villi.,
in general only mechanically sensitive,
however ,
still piezo-
electric
.
Ampullae,
viz ampullae
of loren*ini
,
are
canals, usually 1cm long
by 1mm
in diameter,
filled with
an acidic mucopolysaccharide
,
and
are
directly connected
to
the epithelial
layers.
The
wall resistance
of
the
canal
is
6M A “cm^
with
a capacitance of 0.4 -0.8 PR* cm-2
The
jelley however,
has
a resistivity of
25-31
n *cm?
an<3-
a composition that
varies' somewhat
spatially,
but
in general
is,
in mM«kg“-L
HjpO,
2fa(2)
445
Ca(2)
and Mg(2).50,
^l(-l)
580,
K(l)
12.5,
and urea 75
(MURRAY)
The
ampullae occur
in many varieitos of fish
and
amphibia,
In
general the
end organ
is
10-115
pm
in diamter,
with
groups or clust
ers
of 10-50 ampullae per organ .
The density of the organs range from
12-80* item"''
.
and may number
si total of 40,000
in
any given species.
Of specific
interest to us
is! that the
ampullae respond
to field strong
of 1-15
yV«m- l
if
and only
iff the
input frequencies
aro
low,
that
is
in general,
less than 300 Hz,
and most ofton,
less than
50 Hz.
The
third type
of lateral line organ
is called tuberous,
and
is
found
onljr
in weakly electric
fish.
The tuber ii
are located
in
invag-
inations below the
epithelium; and
surrounded by
_ supportive
colls.
The
tuber ii
responded
to fields
of 4-20
uV*m~l
.of high frequency, usually
up
to
1000 Hz,
specifically synchronous with t ho electric
organ: dis-
charge of that fish or of electric fish of
the
same
specie.
Figure 3.6
shows the comparative
' physiology of the three receptor
organs.
Ordinary lateral
lino
olectro-rccootors
are
innervated by
an
efferent neuron,
tho
ampuallab
and tuborii
are not.
a COCHLEA
(
/
\
AMRULL AE
TUBER I
\
w
afferent
efferent
iir'
Specific electro-sensitivity was demonstrated by Di jkgraaf
: and
Kalmijn .(1963,
1966,
1971)
in
a series of experiments omnhasizingboth
electrolocation
and conditioning stimuli
phenomena.
At first they
noticed that dogfish resnondod
to
a
5 Hz 10” 5 V»m“^ field by a twit chin
of the eyelids,
interestingly; enough,
when
the dogfish wore
drowsy.
(Altered
States'
researchers
,
take note!)
Later they noticed that when
tho ECU .was monitored,
the heart rate
slowed from
1 Hz to 0.7 ^ z when
stimulated
bjr 10”° V*m”^ fields.
Late
in
I 96 S dogfish wore trained
to
seek food with
a 4X10”5
V.m-1, field.
In
all
cases,
tho responses
to
stimuli!
or conditioning ceased when the
ampul 1 ary nerves were sections
even when the field strength was raised
30 db
(1000 X).
In addition
it was noticed that
the dogfish naturally avoided areas where
the field
strength
was
1- 0.1 mV
1 ®"-*-,
Three types
of responses
to
E fields were noticed.
Tho first
reaction was
a twitching of the body or appendage
, ^usually
in
' response
to
10 -
;l
7"m" 1 f iald
but
some time o
t.o low as
10“
V*m' -r
The
>cond
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response type
is galvanot axis,
a turning
into
or perpendicular
to
the
field.
The third response
type
is galyano-narcosis
,
an
immobilisation
of the tost
subject.
Pinal proof
of specific
electrosons.it ivity was provided by Kalmij.
who sequestered
a flatfish
' prey
'
in
sand,
and with
other controls
excluded visual,
mechanical
or chemical
clues,
yet
the
sharks
and rays
were
able
to locate
the flatfish.
Extra, weight
is
added
to
the
evident
when
an artifical current similar
to that
of the flatfish produced
the
samo location results.
.
Figure
3. 7 below
is
an adaptation of Table 2
(
in Kalmijn
(1974),
which plots
the electrosensitivities
of
the various species
tested, by
numerous
authors.
Two types
of electrical energy parameters
wore
presenr at^thc time of testing,
one being field strength,
in volts per
meter
(V*m
)
and
the
other being current density
in amperes per
centi-
meter
s quared
(A*cm"2).
Curve fitting reveals that the relation be-
tween the
two threshold parameter
is
(3. 8
)
log V»m”l
log
A*
for* 2
5,0
Figure
3 .8 below shows
that
in sharks
and
rays,
generally consider;
weakly electro
sensitive,
the relation between current density and
in
-
put frequency reveals that
the
threshold,
increases
as the frequency
increases,
a relation which
is
also observed
in man
(Cf.
3.6
-
3 . 9 ).
.
.
log:
Volts per
p
.
'
...
.
meter
-4
-6
-8
|-12
-11 -10 -9 -8 -7 -6 -5 -4 -3
-2"'
log1Q
amperes
per centimeter
squared’
Although specific eloctroreception has been demonstrated by Liss-
mann,
Bullock,
Kalmijn
and others,
of specific
interest
to us
is that
tho
ampullae have boon demonstrated
to
bo widely receptive
to other
electrical
and para-electrical Phenomena.
Tho reader
is requested
to
study the following closely,
for what has boon demonstrated
in the
ampullae
of fish has
also been observed
in man,
that
is both wide
and
n irrow electro-response.
Hero
the work
is
primarily due
to Murray,
specifically on the
ampullae
of Lorenzini.
Our
interest
in this organ
is due
to t ho fact
that
it
is widely distributed
in vertabratos,
and
its response
to
electrostimuli
is
a,
conic,
that
is frequency coded message
,
whereas
the tuber ii
,
round only
in fish possessing EOD,
responded with phasic,
burst-dispersive
andtonic-ph asic
codings
.
Figures
3. 9a to
3.9b
below show the following responses of
a
single unit of ampullae.
In 3.9a tho
input parameter
is voltage
and
polarity.
Notice
that
the organ has
a discharge
rate
of about
30 Hz,
and
tho output frequency therefore
codes polarity
and
intensity.
No tic
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B 10PHY3 IC 3
ELEC IR0RECEPT0R8
! 22
also
tho
3 kHz).
Pi
( infra-rod
)
and touch
(
reveals; the
sea, water
the
average
that of
sea,
range
of el
cutout
100'
frequency
impulses
per
;
75]
second
output frequencies
in
all
cases
are
in the SIP
r
guro #3.10
shows
the
response
of the
same unit
to
>
while figures
3 .11
abc record the
response
to
via the piezoelectric effect),
and finally,
figure
response
of
an ampul 1 ary organ
to
the
application
olution of various
salinities.
(It
should be noti
concentration of
the neural
solutions
are
two
to
water.)
Again,
what
is moot
important hero
is the
ctro-oonsitivitics
of the
ampul 1 ary organs
,
Pig.
3.9 Response
to electrical
stimuli
ange
(3
-
tompcrate
pressure
3. 12
of
a
ced that
three tim
wide
25)
o L
• A'
:*r-
"
"~
"0
an o dal
To
"“”'15
cathodal
Pig.
3.10
Temperature
impulses
per
second
J
'
I6.5°c
L
—16 . 0°C
/ i
stimulus
in
microvolts
Pig 3.11
Pressure
differential
40 1
I
\
imi
Si l
/
o
“
" i
_
Time
4 min
Pig.
312
!
100
cm Hg
Salinity
Change
of
frequency
in
:
impulses
per
second
(ie Hz)
-10
-20
••30
100
" 110
fo Sea water
120
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Returning to
specific oloctroreeopt ion ,
it
was observed that
stimuli
of duration 0.1 to 1.0 seconds wore
race b effective
in eliciting
electrorosponso .
. Waveforms of the
above durations
are equivalent
to
10 Hz to
1,0 Hz frequencies .
Also observed
was the phenomena that
stimuli that were either too brief,
or sustained for too long were lee
effective
in producing oloctrorosponso
.
Along the
same lino
,
it was
shown
that' stimuli
that
were either amplitude modulated or frequency
modulated gave response when the field strengths
were
as low as
The most effective frequency of frequency modulation was
3 Hz.
Let the render remember
this,
for
in the next section,
(on
audition),
it will be
shown the
the
auditory sense,
again,
histologies...
derived from t he LLO
also demonstrates
a propensity to sclectivly
respond
to
certain stimuli,
if
rand only
if that stimuli
is either
amplitude or
fre auency
raodu.la.ted
.
In addition to
the
above electro receptive characteristics ,
a
phenomena of value
‘o
the biofocdback researches occurs when two
olectrosensitivo
species with electric organ discharges
(BOD)
were
connected with contact electrodes.
It
was observed that when
the
two
fish of the
same
specie,
each with
the
same basic EOD frequency,
were
connected,
both fish we~c
observed to change their EOD
in such
a ma,nne:
to produce .dissimilar,
that
is divergent EOD.
Scbeich and Bullock
indicate
that
this
i; jamming avoidance response" occurs
so
that each
fish can discriminate
its own neural patterns,
such that
confusion,
and therefore less
self control may be
avoided.
This JAR accurs only
if the EOD’s
arc
within 15“ 20 Hz of each fish'
own
EOD.,
and
is most
pronounced when the difference
is
3-5 Hz.
This
is related
to
the
EM response
to
3 Hz,
since
two fish within 3-5 Hz of each other will
produce high amplitude
3-5 Hz modulation of each others EOD.
The phenomena of eloctrorceept ion has boon studied by the various
psychologists,
physiologists
and biologiosts,
who deem the phenomena
as necessary for
the survival of the
aquaric species observed.
They
call
the phenomena oloctrolocation and electrocommunication,
yet
a
perusal of their bibligrauhies
indicate
that none has cited or studied
the biophysical
or parapsycbological literatures,
and their conclusion
wore thus reached
independently^
In oloctrolocation,
for example
,
in Eigenmannia,
the males’
basic
frequency wan
250-600 Hz,
whereas
the females’
was 340-560 Hz.
Similar."
an adversary or prey may bo located
at
a distance
thru olcctroroccptio:
Again,
certain current voloocitios
in the oceans
or
streams con-
tain specific
life
forms,
and
it has been demonstrated that
sensitivity
to
current velocities
as low as
2.5 x l05m« sec-lexist .
Another report
indicates
that 10“° m*soc“
may be used
as
a conditioning stimulus.
In
a similar study,
a
2 vm vibration
at
15 Hz resulted
in
a 50 Hz sign
(Schwartz).
The
ampul 1 ary organs,
specifically electrorccptive
,
have boon
shown to
have, evolved
independently
in several species of vertabrates.
And,
nltho
the LLO
are found only
in water dwoliero,
evolution has
produced two organs specifically derived from the
LLO.
At
one time,
olf.action,
vision,
and certain cranial nerves were part of the Lateral
systems.
In men,
the vestibular canals
(not unco incident .ally named
Ampullae by anatomists),
and the cochlea are directly descended by
amphibia!
LLO.
In
the next section wo
shall show the very close
similarity between the oloctroroccptivc
ampul lary organs
in water
dwellers,
and
the
acountioo—oloo bro ruoovtivo vestibular
;m<3
cochlear
organs
in man.
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3.4
AUDITION
and MECHAN0-R3CBPTI0N
Hop ailing that
J.
Rhino has
indicated that GESP
is ultimately
processed thru the central nervous
system and that the
acoustico-
i ator alio system proceoooc
some clectro-rocopt ion ,
it behooves us
to
study
p.udition
and
the mochano-rcopot ive functions,
for
it will jbe show
that there
are gonerali sod responses applicable
to
all sensory modalitj
The sensitivity to sound
is
in fact the
sensitivity to mechanical
vibrations sot up
in the
atmoshhoro
•
Tho
inner ear
is
the
site
of this
ac ouo t ico-mochan i c al reception,
and those familiar with audition know
that
the mechanical vibrations
set up are relayed
to
tho
inner ear by
a. skin-like membrane,
the
tympanum,
and then thru throe
small bones,
the malleus,
the
incus,
end
the
stapes
to
the
inner oar which .contains
both the
semicircular can ala
and the
cochlea,
both of which innervate
the eighth cranial nerve.
It has boon shewn by von Bokesy,
who won
the
Nobel Prize
for his work,
that the
process
of audition includes that
of hone: conduction
as well
as the response
of
the whole body
to mechanc
vibrations.
And
altho
audition
is
a special zed sense
sequestered from
spurious vibrations,
wo
is
a kind of mechano-rcceptor
,
the
In
addition,
tho
skin
is
the
site
of contact electro-reception,
shall show later how the whole organ of
results
applicable
to
"dormo-
xne
s.o
optics
,
and
it
shall be shown the similarities between these various
sense modalitic
Specifically,
tho
organ of audition
is
in the
Center of
tho cochlear spirals.
The
the organ of Corti located
Organ of Corti
is
a layer
of hair cells connected tothe basilar membrane which connects directly
to
the
eighth nervo.
There
arc
about 15,000 hair colls on tho .basilar
membrane
and
a total
of
.about 15,000 neuron fibres
in tho coclear nerve
These hair cells range
in size from
about 130 pm,
which
arc high froque
cy responsive,
to
275
pm,
which
arc low frequency responsive.
Of un-
usual
interest
is
that
these nerve
fibres have afferent norve ending,
tho tho ire
is no motor function to bo
found,
and
in addition tho
ootenti
across
tho hair cells
in
the organ of Corti
is
about 140 mV.
Since
these
cells
aro
approximately 140 ym
in length,
the field strength
represented
in
the
orran of Corti
is
;
3 .
E
=
dV
• ds" 1
=
1.4 n„lCr^ v __ _
l.o*
10 5
V* nT 1
1.4
*lcH'm
The hair cells
in tho
organ of Corti relay the mechanical ivibratic
sot up upon them from the
stress-strain reactions caused by their bond:
a heat
set off by tho friction then stimulates,
thru this piezoelectric
process!,
the generator potentials.
Of interest
is
that
,
up
to
60 Hz,
tho output frequency response
is
an
exact' match of the
input frequency
response.
'Above
this frequency specialised
areas
in the cochlea at
specific distances from
the
stapes produce complex volleys responsible
for audition
at these
frequencies.
In addition,
the
three
semi-
circular canals,
which resemble
throe mutally pcndicular half-loop
antennae,
arc
also
innervated by cupulao
and microvilli which
also
respond! in piezoelectric process
to mechanical bending set up by
chango
in velocity and acceleration.
Those
semicircular canals
arc
about
cm
inj hoighth,
and thus may provide oloctrorocption form
EMR
in
this wavelength.
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Our
into root
in audition
ah in
to
tho
late -••alii
jtorac from that
oonso modalities* rolatio
system,
but
also from tho
fact
that audition
is
a sonoo modality concerned with tho frequency
and
tho
intonoity of
tho
inpinging stimuli.
Altho
the energy intensity by which audition
is measured
is usually Watts per centimeter squared
(W
*cm“ 2 ),
tho
various energy dimensions
o.ro interchangeable ,
like motors
into
foot,
and
it
can
be directly shown that
tho olectro-sonsitivty threshold
may be computed directly from
tho
audition thresholds.
(Cf 3.5
)
hnc;
as
vo
hrvo
indicated
that
that audition
is
a skin membrane
,
there exist several formulae
re-
mass,
and
tho
proportion of bone mass
bone conduction phenomena,
and
lating
akin area to height
and
to
total
orvan ism mass,
it
is not
surprising that
a mathematical
relat-
ion ship exists between auditory thresholds
and physical body moasurom
monts.
Mastorson
and Diamond have
ompiriCcally produced
tho following
relationships,
Eq.
3*
shows that the lowest frequency responded
to
is related
to
threshold
it
threshold.
body mass,
and that
in addition,
the high frequency
a mathematical function
of the low frequency cutoff
3.10
3.11
low
*^^Snio.,G8
(kg)
1high
"
20 kHz x log
f]_ ow
2.5
Other physiological
aspects
of
.audition illuminate processes
relevant
to
extra,
or
as
ono professional
calls
it,
expanded sensory
•
perception.
Von Bekosy studied t ho
auditory thresholds with
respect
to throe variables;
heredity,
ago,
and real
time.
He found for
instan
that
the
audiograms of members within
a family wore significantly
similar,
whereas
an audiogram of someone not
in that family differed
well
significantly.
Vis,
a proof of the heredity of sensory
and
perceptual abilities
and responses.
In addition
it was found that
with increasing
age,
the
ability
to perceive
sound frequencies
and
intensities was
attenuated
at
tho high frequency response band,
but
was basically unchanged
at
the low frequency end of tho
acoustical
spectrum.
Of very specific
interest
to perception was von Bokesey
* 3
observation of the following.
Audiograms wore
ma.de
of subjects'
threshold sensitivities
at
two
frequencies,
200 Hz
and 1000 Hz,
and
observed for
a duration of twenty minutes.
It was found that
the
thro
holds varied
to
db with periodicities
of 8 minutes for the
former
and
6-J- minutes for
the
latter.
It would prove
to bo highly illumin-
ating
to discern
if
ea.ch frequency had
its own periodicity of thres-
hold sensitivy,
for
in addition,
periods
of 8 minutes
and
6\ minutes
are
in effect frequencies of 1/480 seconds
and 1/330 seconds,
and
it
is well demonstrated that higher frequencies
are harmonica of lower
frequencies,
and
we may summize ultra-low frequency modulation
or
driving of higher frequency ncoustico-mechano-electro-sensitivitios
,
In this
section we
shall observe
tho various response character-
istics
of tho
auditory system
in order
to establish the limits of the
energy perceived.
It has been adopted
as convention that
tho root-
mean-square threshold for audition
is 0.0002 dynes*cm" 2
at
2000 Hz,
the fcquoncy of highest sensitivity.
However,
von Bekosy has observed
sensitivity thresholds
of 0.00005 dynes. cm-2
at
2200 Hz.
Therefore,
while
wo shall
_ discuss
standard auditory thresholds,
but
again tho
reaaor
is cautioned
th.at
tho demonstrated human perception thresholds
may vary
as much
as
to
15 db below the
50$ population standard
threshold.
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Like the olectrorocoptivo senses
(ELF,
infra-red,
and vision),
audition
is orgotonic,
that
is,
its sensitivity of energy
(ergo)
is
a function of the frequency
(tono)
of the
impinging stimuli.
Also,
like
the
elec trorecentivo
senses,
audition shows the property of
oxcitory and
inhibitory response.
That
is,
most neurons have
a base,
that
is
a static discharge
rate,
so
that
,
while
some
stimuli
canoe
an.
increase
in the neural discharge rates ,
some stimuli
cause
an
inhioitic
that
is decrease
of the
static discharge
rate.
This
is
. a
^ very inter-
esting aspect
of sensation and perception,
for the percipient must be
aware that the
information
is
"the
absence
of information, ",
or
to^ put
it crudely,
the percipient
is required to be
aware not of the dpnut,
but the hole.
!
Figure
3 .I 3 below
ic the
oft
reproduced
and now standard auditory
intensity-fro quoncy response
chart.
The
sound pressure level
is given
in
dynes'* cm"
"
and Joules per
esqunre motor per second on the left hand
ordinate
and (tentatively)
Volts
per meter per
square root
of the froqu-
-1
nocy
(y*m-lHz“T)
on the
right.
It
is convenient
to use J*m“2
rather than Watts •cm” 2 because
wo
should like
to bo
able
to calculate
the energy
in
a given fro qu.0 n.c3r waveform.
(See the next section for
conversion factors.)
10 2
-2
10-4
Figure
3.13
;
-
-
-
- 8
-10
-12
Auditory frequency-energy threshold response
\
\
X
J '
I
\
\
\ \
rises
p'ain
thundei
6.6
symphor
4.6
traffic
2.6
convert
0.6
1.4
tegs
125
log Joules
m 2, sec
250
500
1000
2000
frequency
(Hz)
response
-
inh i b it ory _ 3 ,
4000 Hz
log V
iff
exper:
Additional
evidenco
for the
coding of sensory information Into
the
ELF band
can be
seen
in Figure
3.14
below,
which charts
the frequency
of the noural
impulses
in response
to
.
a. given stimulus
intensity
input
for neurons with fro quoncy-snoc if ic responses.
Notice that
thol hyper-
bolic tangent
(tanh x) growth rate
saturates
at
about 450 Hz.
nerve
impulses
per
second
500
|
j
400
;
i
300
|
f
200
i
100
V'
/
5600
4050
/ /
/
/
/z f 700 Hz.
-12
/
-10
Approve^ For Release 2000/p
/
8/07_: CIA-RDP96-Q0787R000500230002-2o
_6
-4
-2
loS J*m-2 see"1
-8