ACTA NEUROL. LATINOAMER.

1961, 7; 179

(
(

.l
(
(
I Electrore~inogram · of the chick embryo*
(
I. Onset and development

! E. GARciA-AUSTT *~ and
(Montevideo)
M. A. PATETTA-QUEIROLO ***

A NUMBER of aspects of ERG have

not· yet been definitely elucidated.
Such is the case with the origin anc~
significance of the various component
waves. The s~udy · of- retin-al electrical
( activity during embryonal development
may provide some helpful data design-
( ed to fill some of the. existing gaps.
Keeler, Sutcliffe and Chaffee 10 have
The data obtained from · this animal
by other authors are very incomplete.
Birds have been the subject of nu-
merous studies but so far as the
chicken .is concerned references may
be found only ·in Piper's 12 important
report.
The onset and de';'elopment of ERG
of the chick embryo will- be discussed

~
,,·
studied the ontogenic development of in this first report. ~l
lr
ERG in the mouse. Zetterstrom n has l

( reported on its development in the

child from birth to the age of one Material and Methods il
t'

\ year, age at which 'it is similar to the

t"•' .

Sixty-eight experiments wef~ made u.sing

I.

{ adult. He further reported 16 that in
( the premature it appears only on the
( 15th postnatal day.
Although the main objetive of the
t'
60 embryo_s at periods r~nging between
the 12th and 20th day of incubation,
5 chicks of ages from 1 to 28 days, and
3 adults.
In the cur:arized embryo both eyelids,

~
present research was the study of
chick embryo, a number of rec~rds
were take~ from young "and . adult
l chickens for the purpose of comparison.
J .
the nictitating membrane, the upper edge
of·· the orbit, the ocular capsule, and
corresponding ocular muscles were resect-
ed. Thus the whole of the upper portion of
the eyeball was exposed as -far as the
. :!: ., i :_
origin of the optic nerve •. These resections

t • Paper performed in the Departamento de

Biofisica. Facultad de Medicina, Montevideo, Uru.
aimed at a twofold purpose: (i) to expose
the eyeball extensively in order to faci-
l guay.
• • Present address: Laboratorio de Neurofisio-
litate the placement of electrodes and (ii)
to prevent the action ~f the ocular muscles,
( logia, Institute de Ncurologia, Hospital de Clinlcas, thus avoiding the record of the elec-
Montevideo, Uruguay. · .
t • • • Present address: Catedra de Biofisica, Uni-
versidad de Carabobo, Valencia. Venezuela.
tromyogram from these muscles and of
movement artifacts.

r
I ·I
 180 GARCIA-AUSTT. AND PATETTA-QUEIROLo

Non-polarized electrodes were used, the
absence of the photoelectrical effeCt being
checked. An electrode was placed on the
cornea· immediately laterally or medially
to the pupil and another one on the _caudal
portion of the eyeball, next to the origin
of the optic nerve. In these positions, at
both ocular poles, the ERG; attained its
a wave is far deeper and there is neither
c wave nor "off" effect. ERG have been
studied in man with flashes of the same
type 3, G and in the dog.2
The energy released by each flash of
the photostimulator wa.s constant at what-
ever frequency, inasmuch as neither the

maximal amplitude. This arrangement
was important, for at the beginning the
maximal amplitude of ERG was only a
''
intensity nor the duration was. Hence it
more reliable a method than the one
employed classically (mechanical interrup-
tion of a continuous light,4, s, 12 by which

I
few microvolts. the duration of the stimulus decreased
Experiments in embryos :were conducted with the increase of frequency.
at a constant. temperature of 38°C. In·
young and adult chickens exposure of the
eyeball and placement of the electrodes
were made . in the same manner. They
Results t
were anesthetized . with . penthobarbital, Onset of the ERG
the young chickens being placed at ·38°C.
AC amplifiers with 0.155- 0.2 sec. time As seen in Table I, ERG developed
constants ~ere used. As no resistance on the 18th day. Prior to this stage
coupled amplifiers· were employed, 'slow
all the experiments had been negative.
ERG (C·-Wave) phenomena were no~Jaken
into account. The recording system con- Particular attention was devoted to the
sisted of ink-writing oscillographs and 17th day, with 7 experiments, includ-
CRO. An upward displacement in the ing a· 17 ;.2 day embryo. On the 18th
record indicated corneal. eleCtrode .·posi- day, 2 out of 12 (17 %) ·experiments
tiveness.
were negative. On the 19th and 20th
A gaslight flash photostimulator was
days all the results were positive.
used. The energy released by each flash . .
of light was con5tant, 0.3 joule.· Duration · The ERG of the chick embryo pre-
-100 sec.- was also constant. The light sented a number of features definitely
beam -of parallel rays- had· an· area of distinct from the ERCf ·~ecorded after

l
200 · cm2 ahd ·was placed 10-12 em from
hatching.
the ·eye, at all times illuminating the ma-
ximal retinal area. Pupillary diameter
was permanently kept ·at its maximum ERG of the "embrym~al"
throught .local atropinization~ One single and "postnatal" types !'t

I
flash caused dazzling in man during se~
vera} .· seconds · and was . -sufficient · · tt:) When ERG first appeared on the
· · disadapt;·: during several minutes, 'the 18th day of incubation· and; partly, on
darkne.SS adapted eye (Cobb and Morton a).
the :19th, it showed peculiar characte-
~'
Single· flashes and repetitive stimulation
were ··employed at 0.5-75/sec. frequencies. ristics that enabled their classification
· Experiments were performed in a dimly as belonging to the "embryonal" type.
lit room, except when the eye was adapted Late~ on, partly on the 19th day of \
to' da.tkness or light.
· Stimulation of 'the retina with intense,
short flashes provokes an ERG with cha-
·incubation and on the 20th, the retina
responded to light ih the same fashion I
I

racteristics · · slightly· different from that as at the postnatal stage, although

formerly obtained with · tungsten lamps; with different temporal characteristics.
'~ ,:'
 ELECTRORETINOGRAM OF THE CHICK EMBRYO lSI

TABLE. I teristics: (i) it developed with repetitive

Onset of ERG in chick embryo stimulation and failed to do so with
isolated stimuli. (ii) Repetitive stimu-
lation had a progressive effect, ERG
Experiment Incubation Result amplitude increased gradually. (iii)
days
Following intense high-frequency sti-
1 12 mulation or continuous light stimula-
2 13
3 13 tion (illumination) the ERG conside-
4 14 rably increased in amplitude and chan-
5 14
6 15
ged in form and duration to reassume,
7 -15 after a certain lapse, its previous
8 16
9 16 features. (iv) It was not elicited
10 17 with· repetitive stimulation with wave·
11 17
12 17
lengths below 0.51 ~t.
13 17
The ''postnatal" ERG type showed

I
21 17
31 17 none of these characteristics and the
14 17
17 18
changes provoked by repetitive stimu-
37 18 lation were similar to those observed
15 18 +
( 16 18 +
in adult animals and man. The use of
the term "postnatal" does not imply
20 - 18 +
30 18 + that the ERG can be superposed over
35 18 +
36 18 + that of the newborn chick or adult
39 18 + chicken. Quite the opposite, the "post- I,_
40 18 +
45 18 + natal" ERG type presented a longer
46 18 + duration, a lower amplitude and a
18 19 +
19 19 + considerably broadened a wave, · cha-
22 19 + racteristics that set them· widely .~part
( 24
25
19
19 +
+ '
~.

from ERG recorded aft~i - hatching.

26 19 +
! 27
28
32
19
19
19
+
+
+
In figure -1 the differential qualities
of both types of ERG are seen. The
range between what is considered iso-
i' .
I:
'
...
33
34
19
19
19
+
+
+
lated stimulation and repetitive- stimu- 1

l
38
lation, is an arbitrary one. For prac-
41 19 +
__ +
47 19 tical purposes, stimuli induced at in-
48 19 + tervals longer than one second were
23 20 +
29 20 -f regarded as isolated stimuli; those
.( caused to develop re~ularly with a
1 second period or less as repetitive.
\ The ERG was similar to that of the
postnatal· st'age and was classified as
The enhancement of the response
elicited after the onset of repetitive

'
of the "postnatal" type. stimulation (Fig: 1, A 2) may be in-
( ERG of the "embryonal" type was terpreted in two different ways: ( i)
;~~;~
made distinct by the following charac- it would be the case of a recruiting
182 GARCIA-AUSTT AND PATETTA-QUEIROI,Q

phe~omenon, that:. is,. ever more. nu- true phenomenon of postinhibitory

merous receptors responding to each rebound, ( ii) it might be due to
stimulus; (ii) it would otherwise be photic potentiation, to an increased
a staircase phenomenon, i~e., that the adaptation to light. As substantiated !
same number of receptors woul~ elicit below, both phenomena intervene. For t
an enhance response to~- ·subsequent reasons of expediency, this change will
sti!J1u1i. There is also·.· the~ ~~sibility be called "rebound". t
of. r an .. a_SSO!!iation.' of r both. phenomena.' In Table II, i~ seen the. distribution I
Its. exact. mechanism· was.. not establish-.
ed but, for the ~ake of expediency, it
of "embryonal" and "postpatal" ERG
with reference to incubation days.
.
r

will be termed "recruitment". . In 2 experiments ( 20 on the 18th day f

The ~arked increase in the response
following bight-frequency stimulation
or intense illumination has likewise
two explanations: ( i) it mig}lt bt?.. a
and 38 on the 19th day) the ERG was
classified as "mixed". Thus these
embryos gave frank, sustained res-
ponses with isolated flashes as in ERG
l
~

IIII II ~ JIII I JIII1111111111 Jl JJJ~ J.J.lllllll ~ 1 'I I I ' I , II o I I IIIII 111111111 II I II II I

..
,. ' lll.:'l "
t
~·· .
(
)..
{
~

l
of·
sa
re
-; 1 ... t t t ·t t ..., r 1 .J
.f' cla
\' res

~
FIG. 1 . - Chic:1; ERG of the "embryonal" an·d "postnatal~' types. A) "Embryonal"-type ERG. ch
B) "Postnatal"-type ER<;;. 1) Response _to isolated stimuli. No response at A. 2) Recruit- an
ment. No recruitment at B. 3) Rebound. No rebound at B. 4) Stimulation with 0.39-0.51,
micra wavelenghts. No response at A. Calibrations, horizontal', 1 sec. for 1 and 2, 2.5 sec·_
. for 3 and 0.5 sec. for 4; v~rtical, 100 microvolts. In this figure and Jn the following ones,
( 17
the upper tracing is the ERG and the lower the stimulus.
 ELECTRORETINOGRAM ·OF. THE · CHICK . EMBRYO 183

TABLE II "embryom!l" or "mixed" type. These

Distribution of "embryonal" type
comprised 100% of the 18 day embryos
and "postnatal" type ERG and 9 of the 19 day ones, out of a total
of 15 (60 % ). The two 20 day embryos
Experiment Isolated Recruit- Rebound Type showed a "postnatal'' type ERG. Con~
stimuli ment
sequently, ·"embryonal" type records
18 days • represent the early ERG appearing on
15 + + embryonal
the 18th day ·of "incubation, while post-
16 + + embryonal
30 + + embryonal natal records reflect a transition bet-
35 + + embryonal
ween the earlier form that· seen after
I 39
40
+
·+
+
+
embryonal
embryonal hatching.

~
46 + + embryonal

20 . +. + + mixed
Characteristics of the ERG
19 da~s _before and after hatching
18 + + embryonal

I
19 + embryonal
22 + + embryonal a) . Form;
duration and amplitude.
26 + + embryonal
These features varied from embryo
28 + + embryonal
33 + + to embryo _and even within one single
embryonal
32 + + embryonal
( 48 + +
experiment, they changed co-nsiderably
embryonal
on varying the qualities of the stimu-
38 + + + mixed
lus. Nevertheless, when these qualities,
24 + postnatal as well as experimental conditions,
25 + postnatal were kept constant, the ERG was the
27: postnatal
34 postnatal same throughout each experiment.
41 + postnatal With intense, brief stimuli two basic
47 + postnatal
t
t 20 days
waves were elicited, similar to those
described in animals· and in man:
23 postnatal
\ 29 postnatal surface negative a wa\fe~ and positive
b wave.- These waves tended to be-
) • Experiments 36 and 45 were eliminated because
of low voltage.
come more complex usually presenting

! notches of varying depths. ';I'he ERG

had at times a poliphasic appearance

i
of the "postnatal" type and at th~. _and it was difficult to differentiate the a
same time.presented the recruiting ·and and b multiple waves. By a wave is
rebound phenomena as in the "em- meant the first negative deflection with
bryonal" type. Experiments 32 and all its notches, until it reaches the iso-
f 48 (on the 19th day) could not be electrical line. Beyond this line, only b
classified with accuracy inasmuch as waves (b 1 , .b 2 , etc.)· ar~ present.
l
, response _to isolated stimuli was not In figure 2 a schematic diagram is
checked. Both showed the recruiting pr~sented of the _most frequent forms,
and rebound phenomena. the mean duration and ampl:tude of
Out of 25 positive experiments, embryo ERG on the 18th_ and 19th day
(
,- 17 ( 63 %) presented ERG of the as well as of the ERG elicited after
184 GARCIA-AUSTT AND PATETTA-QUEIROLo

hatching. Figure 3 shows the records

from 2 -embryos (A and B) and from A B c
a newborn chick (C). The duration of
embryonal ERG was much Ionge_r than
that of the newborn chick. The most
striking feature was the long duration
of a wave and the multiplicity of b.
The amplitude of embryonal ERG was
considerably lower and ranged within
__...._ ___
very broad limits ( 10-70 microvolts).
b) Latency. 'rable III shows the
readings correspond:ng to 8 experi-
Fie. 3.- Chick ERG before and ·after hatching.
I
ments with embryos and to those of
A: ERG on 18th day of incubation. Broad a wave
5 experiments from birth to over 1 and multiple b wave. B: ERG on 19th day of in-
year. Both series are compared statis- cubation. Shorter a wave and shorter, simpler b
wave. C: ERG of newborn 10-day chick. High
tically in table IV according to the amplitude, sharp a wave and simple, short b wave.
method of Dean and Dixon G for small Calibrations. 1 sec. and 50 microvolts.

samp[ngs. It is seen that in the

embryo the latency was twice as great TABLE III
or even greater than after birth and
that the range was far .wider. Latency of the ERG before and after hatching

Before hatching

n Experiment Incubation Latency

)..JV days

+ 1 38 19 10
2 46 18 16
3 41· 19 ~·· , 16·
4 48 19 ~ 0. 20
...... ,..1--- 5 35 18 25
6 - 40 18 25
7 32 19 35
8 39 18 uo•

After hatching

FIG. 2.- Chick ERG before and after

T
hatching.
n

1
2
3
4
5
Experiment

50
44
51
49
42
Days of age

1
12
365 or more
l
20'
Latency

8
9
9
10
11
l
,(
Diagnostic representation. Ordinates: microvolts.
Abscissas: msec. Full tracing: ERG of embryo on
the 18th day of incubation. a wave is notched. • For the statistic calculation of Ta- t
b wave is multiple and ERG is of low amplitude ble IV this value is eliminated
from the series, for the rejection l
and long duration, 300 msec. Bold dotting: ERG
at a more advanced embryonal stage. a wave coefficient is 0.75 and higher than .
)

->
.

decreases in duration. Fine dotting: ERG of new- that demanded (Q = 0.47 for 8 ex-
born 10-day chick. High amplitude, short duration,
sharp a wave and simple b wave.
perfments). The latency is expres-
sed in milliseconds.
1
~~,
ELECTRORETINOGRAM OF THE CHICK EMBRYO 185

TABLE IV. type only in its first part. In some

Latency of the ERG before
and after hatching
w M
Before hatching 25,0 . 20,0
After hatching 3,0 . 9,0
w. range. M, median. Sw, standard deviation
Between the 18th and 19th days no
significant differences in latency were
found.
c) a wave. As seen in figures 2
and 3 its form was var.:.able. On the
18th day it was often broad and cua-
drangular with a notch in its mid-
portion. On the 19th day it was usually
less wide and resembled the previous
instances only the first portion of a
was present while in others, only the
last. Its duration was not very long
Sw and was extremely variable compared
9,2 with that found after hatching. The
1,3
maximal duration was observed when
isolated. flashes. were employed. Dura-
tion with repeated flashes was lower
and var.:.ed with frequency, during re-
.I
bound, during recruitment, during and r.
after illumination and in general, with
the state of adaptation to light. Am-
plitude likewise showed great :ndivi-
dual var:ations. It also changed with
I
the state of adaptation, it increased I.
during rebound and diminished during
recruitment and with the increase of ~ !
frequency.

~·· '
E L L
I ,---
'L_'·.
--. -=-·

8 c

I I
I
F1c. 4.- Ocular movements elicited by photic stimulation. ERG, ocular movements and elec-
tro-retinogram. E, flashes of light. A-C, records at different speeds. At the onset of A, spon-
taneous movement is seen. Movements provoked by light produce similar artifacts with the
ERG super-imposed. Calibrations, 1 sec. and 50 microvolts.
; I

186 GARC'IA-AUSTT AND PATETTA-QUEIROLo

d) b wave. Its shape was. likewise sectioning this nerve, the movements
irregular and usually presented mul- elicited by flashing light disappear
tiple notches both in . its ascending although ERG and spontaneous move~
branch and in the descending one. ments still persist.
Particularly during rebound the notches Likewise, clonic corporeal move-
became more marked and as many as ments were observed as a result of
five b waves were identifiable (Figs. 2 photic stimulation at the same stage
and 3). Its duration was longer than of development.
that of a with isolated flashes of light,
probably exceeding 400 msec., although
this finding could not be accurately Discussion
ascertained owing to the characteristics
The outstanding features of the ERG
of the amplifiers .used. With repetitive
before hatching are: longer duration
stimuli its duration diminished progres-
anc;I greater complexity of waves as
sively until it disappeared altogether
well as highly reduced amplitude.
shortly before fusion frequency. Its
The a wave constitutes the salient
amplitude was likewise variable. These
phenomenon and in this respect, the re-
features of b were different with those
- ported data coincide with those obtain-
found after birth, where the shape was
ed by- Keeler, Sutcliffe a'i1d Chaffee 10
more regular, the amplitude conside-
from the embryonal mi.ce. a wave is
rably wider and notches were seen
always larger in man and in all adult
only occasionally in t h e ascending
animals when high-intensity flashes of
branch (Figs. 2 and 3).
light are ~utilized but its duration, on
the other hand, is very brief. In the
I
I
I

Movements elicited embryo, a wave has a much longer I

duration. b wave is of a multiple pat- !
by photic stimulation i
tern, particularly after·' adaptation to I
I
In 15 embryos a study was made of light. This finding coincides with that
ocular movem·ents elicited by photic reported by Granit,8 who described a l
stimulation. In non curarized embryos surprising variability of b wave to a
the flashes of light caused- steady sustained stimulus. The .latter author
ocular movements similar to sponta- upholds the view that component PII
1
neous ones, starting from the 18th dB:_:}': ___-_- which determines b wave- repre- ·
of incubation. Prior_ to __ this ..time·- -re- sents the summation of a certain
. suits were always negative. In figure number of processes with different
l
+
4 is observed the constancy of these periods of latency and speed. This
movement artifacts. They are distinctly multiplicity of PII :D,lay be even more !
I.
distinguishable from the ERG on which safely contended for the embryo.
they are superposed. Adrian 1 has reported that in man
It was established that these move- tl)reshold flashes eliciting sensation t:
ments are of a reflex origin and that may be unaccompanied by ERG. Con- I
their afferent pathways course along versely the onset of ERG does not r
;
· the optic nerve inasmuch as, after necessarily involve ·optic nerve dis- t
r
 t:LECTRORETINOGRAM OF THE CHICK EMBRYO
charge and the corresponding sensa-
tion. Granit 7 has demonstrated that
under certain conditions, the ERG is
not followed by the optic nerve dis-
charge. _The primary process in the
onset of b wave (PII of Granit) is the
one triggering or at least consistently
accompanying optic nerve discharge."· 7
This process disappears, as a result of
the action of ether and of anoxia 8
prior to the development of the pro-
cess eliciting a wave (Pill of Granit)
and particula~ly, by the action of po-
tassium.H When PII disappears, optic
nerve discharge is absent. Hence the
onset of ERG does not necessarily
\ entail visual sensation. Furthermore,
I
in the chick embryo, there is a frank
prevalence of Pill with a wave con-
siderably widened, the whole process
being inhibit?ry. These two events
might lead to the assumption that the
onset of ERG be not accompanied by
the transmission of se-nsory messages
toward the central nervous system.
However the determination of reflex
I movements which disappear following
I section of the optic nerve conclusively
l demonstrates that when ERG appears
in the embryo on the 18th day of
J incubation, the optic pathways - are
I
i
permeable.· This contention agrees with
the observation of Kuo, 11 who, by
I
-t
direct study found motor_ responses _in
the chick embryo by photic stimula-
tion at the end of the 17th day of
I
incubation in 5 % of experiments, on
the 18th day in 12 % and on the 19th
in 51 %. In addition, Hunt and Gol-
drjng 0 demonstrated in the rabbit that
~ .the optic pathways are permeable
l before the retina is found to respond
( to photic stimulation. Actually they
recorded- evoked potentials from the
187
visual cortex by electrical stimulation
of the optic nerve before the onset of
responses to light. Response to electri-
cal stimulation was apparent at birth,
before the myelinization of the optic
pathways; latency was quite large and
fatigability very fast. Response to light
appeared on the 7th day, when the
pathways are already fully myelinized.
On the whole it may be stated that
in chick embryo the central nervous
system is already well developed when
sensory inflows to higher levels have
already started. This finding further
substantiates _Carmichael's 1 a w of
"anticipatory function", which states:
"functional capacity may be demons-
" trated in many action systems of the
" growing organism well b~fore the
"time when the function in question
" is normally called upon to play an
" active and significant part in the
" vital economy of the organism."
Sufficient experimental evidence has
been presented as to establish the
existence of two types of ERG in the
embryo, namely, "embryonal": and
"postnatal". The former.. represents the
earliest stage of retinal··aevelopment
and -the latter an inter~ediate stage of
maturation. The differences· between
both are explained by adaptation to
light, which requires very intense
flashing in order to become elicitable
in the "embryonal" type of eye. The
"postnatal" eye, such as that of the
newborn chick and the adult eye
become immediately ~a,dapted to light
as a result of intense stimulation.
These views will be discussed in a
forthcoming report.
Rebollo 1 ~ studied the histological
and histochemical development of chick
embryo retina. She concluded that
188
there :s a critical period of ~evelop­
ment coinciding with the onset· of ERG
and· its early evolution. "Nissl bodies
and alkar ne phosphatase first appear
in ganglio"nic cells on the 17th . day,
in the amacr:ne on the 18th day and
in the bipolar on the 19th day. By the
18th day the paraboloid of visual cells
is well developed while the metachro-
matic stain of its external segment is
no longer present."
Summary
I. A study was conducted of the electrical acti-
vity of the retina in chick embryos, with flashes
of light, a comparison being made with the ERG
of newborn chick and adult chicken.
2. · EE.G appeared in the embryo on the 18th
day of incubation.
3. Two types of ERG were dilferentiated in
the embryo, the "embryonal" type obtained from
all 18-day embryos and part of 19-day ones and
the "postnatal" type partly recorded from 19-day
embryos and all of 20·day ones.
4. The "embryonal" type ERG was elicited
only after intense adaptation to light. It was not
obtained with isolated stimuli and presented the
recruitment and rebound phenomena.
5. The duration of embryo ERG was greater
and the amplitude lower than in adult ERG. The
long duration of a wave was the most salient
feature. In the "embryonal" type, a wave was
wider than in the "postnatal" type and in the
latter was always greater than in the adult.
b wave was generally multiple and extremely
variable.
Resumen
1. Se estudi6 la actividad electrica_ de ·- 9.
la retina en embrlones de polio; co.ri·cf~te-
llos de Uimpara de gas y se compar6 con
el ERG del polio reciE'in nacido y de la 10.
gallina adulta.
2. El ERG apareci6 · en el. embri6n a
los 18 dias de la incubaci6n.
3. Se diferenciaron dos tipos de ERG: 11.
el tipo "einbrionario" que· se obtuvo en to-
dos los embriones de 18 dias y. en parte
en los de 19; y el tipo •·postnatal" que se 12.
registr6 ·en parte en los de 19 dias y en
. GARCJA-AUSTT AND PATETTA·QliEIROLQ
todos los embriones de 20 dias. El primer
tipo represent6 Ia etapa mas precoz del
desarrollo funcional de la retina.
4. El ERG de tipo "embrionario" se ob-
tuvo solamente despues de una intensa
adaptaci6n a la luz. No se obtuvo con· es-
tfmulos aislados y present6 los fen6menos
de reclutamiento y rebate.
5. La duraci6n del ERG del embrion
fue mayor y Ia amplitud menor que el del
adulto. La gran duraci6n de la onda a
fue el car<icter mas saliente. En el tipo
"embrionario" la onda a fue mas ancha
que en el tipo "postnatal" y en este fue
siempre mayor que en el adulto. La onda
b fue generalmente multiple y sumamente
variable.
References
I. ADRIAN, E. D.: The electric response of the
human eye. J. Physiol., 1945, !04: 84-104.
2. BoRRAS, A.: El electrorretinograma del perro
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