On the Functional Anatomy of Neuronal Units in the

Abdominal Cord of the Crayfish, Procambarus

clarkii (Girard)
C. A. G . WIERSMA AND G. M. HUGHES
Biology Division, California Institute of Technology,
Pasadena, CaliforniaZ

In a previous publication (Wiersma, '58)

it was shown that a number of the
neurons whose axons form the so-called
circumesophageal commissures of the
crayfish can be characterized by ( a ) the
nature of the sensory stimulation to which
they respond, (b) the type of response,
which distinguishes primary sensory fibers
and interneurons, ( c ) their location in the
cross section and (d) the size of their
action potentials, which is related to their
diameter. Using these criteria it was possible to establish 100 physiological entities, even though only two main types of
sensory stimulation were used, namely
touch of hairs on the body surface and
manipulation of joints. The same methods
have now been applied to the connectives
of the abdominal cord, in which the number of fibers is about half that in the
commissure.
A report has already been presented of
the types of physiological connections
made by axons of the cord, without entering into their individual properties
(Hughes and Wiersma, '60a). The purpose of the present paper is to classify the
data obtained for the cord in a manner
comparable to that for the commissure.
Only those entities are described ( A 1 to
A 75) which have been repeatedly found
in a number of preparations. They are
presented in table 1, to which the reader
is referred for all those cases in which
only the number of the entity is used in
the text. The results were collected from
about 120 preparations, the great majority
of these being isolated abdomens.

commissure, and have been described in

detail elsewhere (Hughes and Wiersma,
'60a). It was found that the isolated abdomen preparation had considerable advantages over that of the whole animal.
Its survival time was at least as long, and
in general it remained in better condition
than whole animal preparations. The best
results with the latter were obtained when,
in addition to exposure of the cord, the
circumesophageal commissures were also
prepared. It is likely that this difference
was due to improved circulation; replacement of blood by the perfusion solution
(van Harreveld, '36) is greatly facilitated
by opening the thoracic cavity. It may be
that clotting of blood is one of the factors
producing untimely deterioration of the
preparations.
The cord was exposed from the ventral
side by stripping off a median part of the
sternal area between the second and 6th
abdominal segments. Great care was
taken to keep all roots of the ganglia intact. First roots are especially easily damaged, and inspection and observation of
spontaneous movements of the swimmerets was routine in order to exclude this
possibility.
For several reasons splitting of the cord
was a source of possible experimental
error. The surrounding membrane is very
tough and it takes considerable force to
penetrate it. This probably leads to damage of a number of axons in each preparation along the line of split, which was
usually in the horizontal plane, dividing
the cord into about equal parts. The next

METHODS

'On leave of absence from Zoological Department, Cambridge University, England.

ZSupported by grant G-5461 of the National
Science Foundation.

The methods were similar to those used

for the analysis of units in the crayfish

209

210

C . A. G. WIERSMA

AND

G. M. HUGHES

TABLE 1
Short descriptions of all fibers running in the connective between the third and fourth
abdominal ganglion discussed in the text, are listed in numerical order
The location of the sensory fields to which the fiber responds is given after the number,
followed by the type of stimulation to which it responds. Next is indicated whether the
response is obtained from the same side of the animal as the prepared connective (Horn.),
from the other side (Het.), from both sides (Bil.), or that the response is asymmetrical
(As.). L. refers to location in cross section as presented i n figure 1. Id. C. indicates that the
fiber is identical in reaction with the commissure fiber with that number (Wiersrna, '58).
Abbreviations used in table
App., appendages
Dors., dorsal
Abd., abdominal
Vent., ventral
Seg., segment
Prim., bundle of primary sensory fibers
SRI, tonic stretch receptor
Med., medial location of the interneuron
in the cord
SRB,phasic stretch receptor
Sw., swirnrneret
Lat., lateral location of the interneuron
in the cord
Tels., telson
Urop., uropod
Pl.pl.1 or II., pleural plate hairs innervated
U. Seg., uropod segment (6th abdominal)
by the first or second root
A 1.
A 2.
A 3.
A 4.
A 5.
A 6.
A 7.
A 8.
A 9.
A 10.
A 11.
A 12.
A 13.
A 14.
A 15.
A 16.
A 17.
A 18.
A 19.
A 20.
A 21.
A 22.
A 23.
A 24.
A 25.
A 26.
A 27.
A 28.
A 29.
A 30.
A 31.
A 32.
A 33.
A 34.
A 35.
A 36.
A 37.
A 38.
A 39.
A 40.
A 41.
A 42.
A 43.
A 44.
A 45.
A 46.
A 47.

Abd. Seg. Joint. Tonic. Extension. Horn? L78

SRI-Ab 5/6. Horn. L78-79
Urop., Tels. Hair. Horn. L82
SRI-Ab 4/5. Horn. L78
Sw. 2-Urop. Basal Joint. As. L82
Urop., Tels. Hair Fringe. Prosser's. Horn. L85
Abd. Seg. 4-U. Seg. Hair. Horn. L82
SRI-Ab 6/T. Horn. L78
Abd. Seg. 5. Hair. Prim. L84
Sw. 1-Urop. Basal Joint. Phasic. Horn. L84
Sw., Pl.pl.1. Seg. 1-5 Hair. Bil. L84
Abd. Seg. 3-5. Hair. Horn. L83
Pl.pl.1. Seg. 2-5. Hair. Horn. L85
Abd. Seg. 5. Hair. Med. Horn. L78
U. Seg. Hair. Med. Horn. L78
Abd. Seg. 3. Hair. Prim. Horn. L83
Tels. Hair. Horn. L83
Abd. Seg. 4. Hair. Prim. Horn. L84
Sw., Pl.pl.1. Seg. 2-5. Hair. As. L80
Urop. Joint. Tonic. Horn. L84
SR2-Ab5/6. Horn. L79
Abd. Seg. 4. Hair. Het. L79
Abd. Seg. Joint. Phasic. Extension. Horn? L79
Sw. 1-Urop. Joint. Phasic. Bil. L81
SR1-Ab3/4. Horn. L79
Sw. 4,5. Joint. Prim. Horn. L83
Sw. 2-5. Basal Joint. Horn. L83
Sw., Pl.pl.1. Seg. 2-U. Seg. Hair. Horn. L84
Abd. Seg. 2. Hair. Het. L82
Abd. Seg. 3. Hair. Het. L82
Sw. 4. Basal Joint. Tonic. Horn. L81
Urop., U. Seg., Tels. Hair. Horn. L80
Sw., Pl.pl.1. Seg. 3 Hair. Joints? Horn. L81
Abd. Seg. 2,3. Hair. Het. L84
Sw. 5. Basal Joint. Tonic. Horn. L81
U. Seg. Hair. Quack. Horn. L81
Urop., Tels. Hair. Dors. Vent. Het. L81
Abd. Seg. 3,4. Hair. Horn.L79
Abd. Seg. 4. Hair. Med. Horn. L79
Abd. Seg. 2,3. Hair. Horn. L83
Abd. Seg. 2-5. Hair. Horn. L79
Abd. Seg. 3. Hair. Med. Horn. L79
Anal Valve. Joint. Horn. L82
Sw. 4. Joint. Prim. Horn. L78
Sw. 5. Joint. Prim. Horn. L78
Abd. Seg. 1-U. Seg. Hair. Dors. Vent. Horn. L83
Sw. 3. Joint. Prim. Horn. L83

Id. C 100
Id.. C
19 _
~
Id. C 89
Id. C 48
Id. C 91

Id. C 65
Id. C 38

Id. C 15

Id. C 93

Id. C 3

Id. C 71

21 1

CRAYFISH ABDOMINAL CORD NEURONS

A 48.
A 49.
A 50.
A 51.
A 52.
A 53.
A 54.
A 55.
A 56.
A 57.
A 58.
A 59.
A 60.
A 61.
A 62.
A 63.
A 64.
A 65.
A 66.
A 67.
A 68.
A 69.
A 70.
A 71.
A 72.
A 73.
A 74.
A 75.

TABLE I-(Continued)
Sw. 3. Basal Joint. Prim. Tonic. Horn. L76
Abd. Seg. 5. Hair. Lat. Horn. L80
Abd. Seg. 4. Hair. Lat. Horn. L80
Pl.pl.1. Seg. 3. Hair. Horn. L84
Abd. Seg. 2-5, U. Seg.? Hair. Dors. Vent. Bil. L78
SRI-Ab2/3. Horn. L78
Abd. Seg. 2-5, U. Seg.? Hair. Het. L78
Sw., Pl.pl.1. Seg. 2,3. Hair. Horn. L84
U. Seg. Hair. Lat. Horn. L80
Abd. Seg. 5, U. Seg. Hair. Horn. L80
Abd. Seg. 4,5. Hair. Horn. L85
Urop., Tels. Hair. Vent. Horn. L77
U. Seg. -5th. Joint. Tonic. Extension. Horn? L81
SRI. All Seg. Joint Tonic. Horn. L77
Abd. Seg. 3. Hair. Lat. Horn. L85
Abd. Seg. 2-4. Hair. Horn. L84
Abd. Seg. 2-U. Seg., Tels? Hair. Bil. L84
Abd. Seg. 5. Hair. Het. L78
Sw., Pl.pI.1. Seg. 3. Hair. Joint. Het. L76
Sw. 3. Hair. Horn. L76
Urop. Basal Joint. Horn. L77
Urop. Basal Joint. Het. L77
Urop. Joint. Bil. L78
Abd. Seg. 3-5. Hair. Bil. L80
Anal Valve. Joint. Het. L76
Abd. Seg. 4- U. Seg., Urop., Tels. Hair. Horn. L83
Sw., Pl.pl.1. Seg. 3. Hair. Prim. Horn. L83
Sw., Pl.pl.1, Seg. 5, Hair. Joint. Horn. L84

step was to peel the membrane surrounding one of the connectives, usually the
left, though sometimes the right or both
connectives were used. In general no active fibers clung to this membrane but
fibers adjoining the membrane may well
have been damaged during peeling.
Figure 1 shows the areas into which the
cross section of the connectives was divided in order to determine the locations
of the small bundles into which it was
split during an experiment. The accuracy
of this procedure was greater than for the
commissures since, in contrast to the commissures, the shape of the abdominal
connectives shows little variability between
preparations, which considerably reduces
a major source of error in localization.
However, another source of error, especially in the earlier experiments, was the
ease with which rotation of the major
bundles occurred once the sheath around
the cord had been removed. Localization
became much more precise after a number
of marker fibers had become established.
Most preparations were of the connectives between the third and 4th ganglia,
but in a large number those between the
second and third ganglia were used, and
some were made between the 4th and 5th.

Id. C 51

Id. C 45

Id. C 56

Primary sensory fibers present differ markedly in the different connectives, whereas
on the other hand few interneurons are
found in only one of these places and not
also in the others. All numbers used in
this paper refer to fibers present in the
cord between the third and 4th ganglia.
Determination of the properties of single entities was simpler than in the commissure, both because of a greater pre-

Fig. 1 Cross section of right connective between third and 4th abdominal ganglion with the
numbers of the areas into which it was divided.
Large structures on top are the medial (76) and
lateral giant fibers.

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C. A. G. W I E R S M A AND G. M. HUGHES

ponderance of fibers with simple reactions

and the large reduction in the extent of
the total sensory area which needed to be
explored. As a result the number of units
found in one preparation was generally
much higher than that in similar commissure experiments. Of the total 75 entities
established, as many as 30 have been recognized in a single preparation, and it was
routine to find more than 15. Hence the
number of times individual units were
found in the cord is in general higher
than in the commissure, although the total
number of preparations was considerably
smaller (about 1/5).

PLEURAL PLATE

,
SEGMENT 4
/
TERGUM

RESULTS

Primary sensory fibers in the abdominal

connectives form a much larger part of the
total number of axons than in the commissures. In both parts of the central
nervous system the fibers from a common
source and with similar function run together in bundles, With the exception of
the primary sensory fibers of the abdominal stretch receptors, which can be prepared as single units, a single number (A
9 etc.) has been given to such bundles of
primary sensory fibers. Both ascending
and descending bundles are found.
Primary sensory fibers responding to
touch of hairs on each abdominal segment
enter through both first and second roots
and will be divided into groups according
to this difference (see fig. 2).

Primary sensory hair fibers f r o m first

root innemation (figs. 3, 4)
A . Ascending. None found.
B. Descendin.g. A 74. Primary sensory
fibers from hairs on the third swimmeret
and posterior pleural plate ridge and anterior 4th pleural plate ridge, which are
known to enter through the first root of
the third ganglion, form a bundle in area
83. This bundle consists of a relatively
large number of small fibers. In the lead
between second and third ganglia a similar bundle of descending fibers from the
second ganglion replaces these fibers.
This bundle was called A 74 ( 2 / 3 ) . A 74
was found 4 times, A 74 (2/3) 4 times.
Summary. The primary hair fibers entering through the first root appear to have
a restricted distribution, compared to other

m d i c q m
THIRD GANGLION
Fig. 2 Schematic representation of the third
and 4th abdominal segments, showing the areas
innervated by the first and second root of the
right half of the third abdominal ganglion. Dark
shading, first root and areas innervated by it,
light shading second root and its areas. Arrow
shows the location of the abdominal stretch receptors, excited by bending the 5th on the 4th
segment, innervated by the second root of the
third ganglion. (From Hughes and Wiersma,
'60a)

sensory fibers. Like all others they are

restricted to the homolateral connective,
but run in this only in a caudal direction
for the length of a single interganglionic
space. There seems to be no histological
evidence for fibers with such a course.
From their function in local reflexes it
appears likely that they synapse with other
fibers both in the ganglion they enter and
in the one more posterior.

Primary sensory hair fibers from second

root innervation (figs. 3, 4 )
A. Ascending. A 9. Primary sensory
fibers of 5th abdominal segment. There
are a number of these small fibers, which
react separately to touch of small areas.
These fibers have also been found several
times between second and third ganglia.
Though they may all enter through the
second root of the 4th ganglion, they must
run well forward. Location 84, near 83.
Found 20 times, is used for marking the
area.

CRAYFISH ABDOMINAL CORD NEURONS

A18

A 47
A16
Fig. 3 Cross section showing the location of
the bundles of primary sensory fibers, responding
to touch of hairs and to joint movements of
swimmerets. Slight differences of location between diagrams and text (e.g. A 48) were
prompted by considering neighboring fibers in
the former.

B . Ascending a n d descending. A 18.

Primary sensory fibers of 4th abdominal
segment. Again there are many small
fibers in a bundle, which runs near the
A 9 bundle. In the lead between the second and third ganglia only ascending fib-

213

ers are found. Location 84. Found 24

times, often with A 9 and A 16.
C . Descending. A 16. Primary sensory
fibers of the third abdominal segment. In
the connective between second and third
ganglia they are present as both small
ascending and descending fibers. They
were not found in 4/5 leads. Location 83
close to 84. Found 15 times, of which
about half in second to third connective.
No primary fibers descending from the
hairs of the second segment were found
in the connective between third and 4th
ganglion, though they occur in 2/3 leads.
Summary. In contrast to the first root
hair fibers, those of the second root, representing hairs on the dorsal surface, run
much further through the cord. They descend only for a single segment, and these
branches may well be direct continuations
of the peripheral fibers, since Retzius
(1890) has noticed a number of fibers
entering through the second roots, which
bend immediately backward, without connections in the ganglion. Their forward
course can certainly span three interganglionic distances, as in the case of fibers
from the 5th segment found in 2 / 3 leads.
Remarkably, the dorsal hair fibers of the
uropod and telson segments appear to

74
Fig. 4 Schematic drawing of side view of abdomen indicating areas supplied by bundles of
primary hair fibers.

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C. A. G . WIERSMA AND G. M. HUGHES

movement of the swimmeret and are tonic.

They might well be fibers connected with
the muscle receptors of Alexandrowicz
('58), if such are present for the swimmerets, for they appear to react only during active contraction. However, it is not
completely excluded that they are interneurons. The responses can be inhibited
by moving the 5th swimmeret forward and
excited by the forward movement of the
second swimmeret. They undoubtedly play
a part in the generation of the metachronal rhythm of the swimmerts. They are
Primary proprioceptor fibers entering
located dorsal of A 26 in area 78 and one
through the first root (fig. 3 )
or more of them were found 12 times.
A 45 appears to be a similar small fiber
The decision whether a joint fiber is
primary sensory or an interneuron is much for the 5th swimmeret which fires tonimore difficult to make than for hair fibers. cally, especially during active backward
In the following enumeration there may contraction of the swimmeret. It can bebe some errors made in this respect. One come inactive although the swimmeret
criterion used, is that fibers forming very maintains its position passively, again inclear bundles, whose responses correspond dicating that a muscle receptor organ may
with that obtained when a peripheral lead cause the firing. It is a close neighbor of
from the root itself was taken, were con- A 44, running like the latter near the absidered as primary. In the cord other dominal stretch receptor axons in area 78.
proprioceptor fibers are more solitary and It was found 7 times.
B . Descending. A 47. Homolateral third
are sometimes of considerable size. When
they responded to the movement of what swimmeret, joint fibers both tonic and
appeared to be a single joint of a single phasic, different fiber sizes. These fibers
swimmeret, they were considered as pri- were found in the same area in 2/3 leads,
where they are ascending. Location 83
mary, otherwise as interneurons.
A. Ascending. A 26 and A 26A. A near 82. Found 10 times.
A 48 resembles A 44 and A 45, giving
bundle of small primary sensory fibers
from the 4th and 5th swimmerets. These a tonic discharge, especially during backare always close together and in most ward contraction of the swimmeret. As in
cases could not be split. Different types of the case of A 44 more than one fiber was
fiber are found, tonic and phasic and occasionally found being medium and
separate fibers for different joints. For small in size. Location 76. Found 6 times,
instance, a certain fiber in this bundle re- three of them in 2/3 leads (ascending).
A 47 and A 48 are certainly different
acts on spreading of the exo- and endopodite. In all these respects the bundle entities, as indicated by their locations,
gives the same kind of signals as can be and by the fact that they have been found
obtained with a peripheral lead from the in pairs in single preparations.
first root of these ganglia. These fibers
Summary. The absence from 3/4 leads
may well run for shorter distances in the of fibers from the second swimmeret may
cord than the following two entities. For indicate that this type of fiber descends
in contrast to the latter, there were indi- only one connective, whereas the presence
cation that only a few if any of the fibers of fibers from all swimmerets in 2/3 leads
from the 5th swimmeret also appear in shows that they ascend for at least three
213 leads. This bundle was found 19 times successive connectives. If the same held
in area 82, near 83.
for the uropod joint fibers, one would exA 44. Primary sensory fibers of joints pect to find them as readily in the 3/4
of the 4th swimmeret. These form a much lead, but this was not the case. Reactions
smaller bundle of medium-small fibers in in single axons to uropod joints were
area 78. They respond solely to backward found in several instances, and although

have a much more restricted course. They

have not been encountered even in 4/5
leads, except in a single instance. This
is in sharp contrast to the ease with which
all other bundles were found, so that, unless more evidence is obtained, their regular presence cannot be accepted. If so it
might indicate that the second root hair
fibers of the anterior abdominal segments
are separately integrated from those of
the posterior region, possibly in the first
abdominal or last thoracic ganglion.

CRAYFISH ABDOMIN'AL CORD NEURONS

it is uncertain whether these are definitely

interneurons, they will be regarded as
such in this paper. Two other fibers specific for a single swimmeret have been
found (A 31 and A 35) which might belong here, but which will be described
under interneurons.

Primary proprioceptor fibers entering

through the second roots (fig. 5)
Whereas previously (Wiersma, '58) the
neurons of the abdominal stretch receptors were named RM1 for the slowly adapting and RM2for the fasting adapting type,
the more accurate designation of SR1 and
SR, (Wiersma and Pilgrim, '61) will be
used in this paper.
A. Ascending. A 2. Tonic stretch receptor of the joint between 5th and uropod
(6th) segments (SRI-Ab 5/6). This fiber
enters through the second root of the 4th
ganglion. The fiber runs on to the brain
and is C. 100 of the commissure. It is large
and located as are all other abdominal
stretch receptor fibers in areas 78 and 79
under the medial giant fiber. Found very
many (21 recorded) times. A 21. Phasic
stretch receptor SR,Ab 5/6. This fiber is
even larger than A 2. Location 79, found
11 times.
A 8. Tonic stretch receptor of telson on
uropod segment (SR1-Ab 6/T). This is a
smaller fiber, which has so far not been
A 21

A 6

A4
Fig. 5 Cross section with location of the
primary sensory fibers of the stretch receptors
of the abdomen and those s o n s responding to
extension of abodminal segments.

215

found in the commissure. It enters, with

its partner the phasic receptor, through
the homolateral second root of the 5th
ganglion. Found 13 times, location 78.
Because the phasic receptor of this joint
was found ony twice, it has not been given
a number, but it was obtained in a 2/3
lead, indicating that all these primary sensory fibers go to the brain.
B . Descending. A 4. SRI-Ab 4/5, entering the cord through second root of third
ganglion. This large fiber is ascending in
2/3 leads, and is identical with C89.
Again it is accompanied by the fiber of
the phasic stretch receptor of this joint,
which was found three times. Location
78, found 11 times.
A 25. SRI-Ab 3/4. Found 13 times.
It is large and is identical with C93. Location 78.
A 53. SRl-Ab 2/3 is identical with C51
of the commissure. Found 6 times. Location 78. The descending branch appears
to be medium in size.
Because they are also most likely primary fibers, though this is not strictly
proven, some fibers which react to extension of the tail, but which do not enter
the cord through any of the roots of the
ganglia, and appear to be excited within
the cord will be next discussed. The reasons for ascribing extensor sensation as
the function of these axons has been given
elsewhere (Hughes and Wiersma, '60a).
A 1. Tonic extensor fiber for at least
the posterior joints and perhaps all joints
of the abdomen, but not of the one from
uropod segment to telson. Details of the
responses of this fiber are given in the
paper quoted (Hughes and Wiersma, '60a).
This fiber has been found many (16)
times. Location: area 78. There are in
reality two tonic fibers, a smaller and a
somewhat larger one, which run together
with the phasic fiber (A 23). The smaller
one is more tonic but stops when extension is too extreme. They resemble each
other so closely that it is not possible to
determine which one has been found when
a single signal is obtained.
A 23. Phasic extensor fiber of the abdomen. It is larger than the previous ones,
and located near them and the stretch receptor fibers. Found 15 times in area
78-79.

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C. A. G. WIERSMA AND G. M. HUGHES

A 60. Also reacts on extension, but for

this fiber it has not been proven that the
sense organs are not innervated by a root.
It is more restricted in its sensitivity, answering with low frequency exclusively to
extension of the joint between 5th and
uropod segments but quite tonically. Its
location is rather removed from that of
the previous fibers, namely area 81. It was
found 6 times. It is a medium-sized fiber,
found also in preparations in which A1
was present. This fiber may well be an
interneuron.
Summary. The finding of SR2 fibers in
the cord makes it likely that all of the
quick stretch receptor cells also send direct
branches to the brain. This was expected
from results in the commissures (Wiersma,
58). According to the histological findings (Allen, 1894; Alexandrowicz, 51)
these fibers should also have descending
branches, and there are two technical reasons for the failure to find their signals
in the present experiments. The anterior
SR,s are known to adapt quicker than the
more posterior ones, and quick bending of
the anterior joints is very difficult to obtain
in the isolated abdominal preparation as
used. The fact that the extensor fibers
run in the same bundle as the SR fibers
enhances the likelihood that integration
of all these signals takes place both in the
brain and in the 6th abdominal ganglion.
Such integration, of which the main function may be the regulation of the activity
of the central giant fibers, is indicated by
the existence of fiber A 61 (see below).
Interneurons
The distinction made above between ascending and descending fibers, was not
altogether appropriate for those of the
stretch receptors and perhaps other primary sensory fibers and can lead to confusion when applied to interneurons. Only
in some cases is there no objection to this
term, as for example the interneuron collecting from hairs on the telson is ascending throughout the central nervous system. But the term is quite inappropriate
when the interneuron collects impulses in
both the third and 4th ganglia so that impulses pass in both directions in the connective between these ganglia (Hughes
and Wiersma, 60a). Because relatively

few interneurons are excited by sensory

fibers which enter through both first and
second roots of abdominal ganglia 1-5,
these terms retain value for descriptive
purposes. However, the fact that the roots
of the uropod and telson ganglion (6th)
cannot readily be homologized with first
and second roots, diminishes their appropriateness.
As in the commissure, interneurons can
readily be divided into homolateral, heterolateral, bilateral and asymmetric depending on whether the sensory field to which
they respond occur on the same side
of the body as the connective in which
they run, on the other side, on both sides
symmetrically, or on both sides but with
an asymmetrical distribution of their sensory fields. They can also be distinguished
according to the type or types of sensory
input to which they react.
It should be realized that there are certainly a number of interneurons present
in the cord which receive their sole input
in the ganglia of the thorax and the brain.
These were encountered in whole animal
preparations but have not been found
enough times to be described here. It will
be noticed from the diagrams showing the
distribution of the axons found, that fewer
are present in areas 76-77 than would be
expected from a random distribution.
There is strong reason to believe that a
considerable number of descending interneurons, but by no means all, are located in these dorsal areas underneath the
giant fibers.
The order followed in describing interneurons will be to start with those responsive to hairs, next those responding to proprioceptive stimulation and finally those
activated by both these methods. First, the
interneurons stimulated by hairs on the
dorsal aspect of the abdomen, will be discussed, first all those with sensory fields
on one homolateral segment then on one
heterolateral segment, next the same for
two segments and then three and more
segments. The interneurons responding to
the ventral aspect will be similarly treated,
followed by those responding to both aspects.
Interneurons stimulated by hairs o n
dorsal areas. The remarkable distribution of the nerves on the dorsal surface of

217

CRAYFISH ABDOMINAL CORD NEURONS

each anatomical segment (which for all

abdominal segments, including the uropod
(6th) segment are innervated by roots of
two different ganglia, fig. 2 ) makes it uncertain in most of the following cases
whether the interneurons respond to the
external anatomical or the neurological
segmentation or to neither exactly. When
an interneuron is described as reacting to,
e.g., 4th and 5th dorsal abdominal segment, it means that it reacts to dorsal
stimulation of both segments but that reaction to the posterior part of the pleural
plate 11 innervation of the third segment,
innervated by the second root of the third
abdominal ganglion, is not excluded. It
will be clear from the following that interneurons can also react to only a part of
these sensory fields. Exact mapping is
very difficult, either because of the necessity of stimulating a large area in order
to make the interneuron fire or on account
of its extreme sensitivity to a few hairs,
and has therefore not yet been tried in
many cases.

Interneurons from dorsal areas o f single

abdominal segments (figs. 6, 7)
A. Homolateral interneurons. For each
segment two fibers have been found. They
differ, as f a r as is known only in their locations within the cord. There appears to be
distinct bundle formation, one bundle running near the abdominal stretch receptor

A I4
Fig. 6 Cross section with the location of interneurons responding to touch of hairs of dorsal
side of abdominal segments; 0,homolateral interneurons; (>, heterolateral.

HOMOLATERAL

HETEROLATERAL

29

30

22
65

I7
Fig. 7 Dorsal view of abdomen with areas
innervated by interneurons which cover only
small dorsal parts, limited to single segments.

fibers, and thus in the medial half, the

other bundle being more laterally located.
All these interneurons appear to react to
the sensory input of a single root, and thus
to the posterior pleural plate 11 area of
one segment and the dorsal area of the
next (fig. 2).
A 42. Medial interneuron for homolateral dorsal third segment hairs. Medium sized fiber located in area 78-79.
Similar to A 14, A 15 and A 39, of which
it is a neighbor. It was found 11 times.
A 62. Lateral interneuron for dorsal
third segment hairs. Homolateral, medium
in size, and located in area 85. Like the
previous fiber, it is found only in diphasic
and anterior recordings. Found 6 times.
A 39. Medial interneuron for dorsal 4th
segment homolateral hairs. It is a neighbor of A 14 and A 15 in area 78-79. Large,
it was found 10 times. This fiber can be
obtained in both anterior and posterior recordings, which makes it necessary that
the primary sensory fibers make synaptic
connection with it in both the third and
the 4th ganglion.
A 50. Lateral interneuron of the 4th segment active in anterior and posterior recordings. Medium in size, it is a neighbor
of, and similar in response to, A 49 and

218

C. A. G. WIERSMA AND G. M. HUGHES

A 22. Heterolateral interneuron of the

A 56. Found 6 times, and located in area
4th dorsal segment hairs. It is a large
80.
A 14. Medial interneuron for the dorsal fiber, located in area 78-79, near A 65.
5th abdominal segment hairs. It is very Found 13 times.
A 65. Heterolateral dorsal hair fiber of
like A 15 in its properties and location.
Large, and located in area 78, it was found abdominal segment 5. Medium sized, located in area 78 and found 4 times.
20 times.
Interneurons integrating dorsal hairs of
A 49. Lateral interneuron of dorsal
hairs of 5th abdominal segment. Very like two abdominal segments (figs. 8 and 9).
A 56 in properties and location in area 80.
Large, it was found 6 times, once in the
same preparation as A 14.
A 15. Medial interneuron of homolateral
hairs on dorsal aspect of uropod segment
only. A large fiber in area 78 which very
closely resembles the next one (A 56).
Found 21 times, it sometimes responded in
addition to hairs on the basal joint of the
uropod.
A 56. Lateral interneuron for homolateral hairs on uropod segment. Located
in area 80, this large fiber was found 7
times, twice in the same preparation as
A 15. Both A 15 and A 56 may receive
their main input by way of the second
root of the 5th ganglion.
A 34
\
A 36 (fig. 13). This very large fiber has
A 40
received the nickname of quack fiber. It
Fig. 8 Cross section with location of interusually has a high threshold and fatigues neurons responding to touch of hairs on two
rapidly. Its sensory field is somewhat un- adjacent abdominal segments.
certain, but after mild fatigue it responds
only to a quick brush stroke dorsally along
the base of the uropod near the 5th segment. Similar fibers, located in the same
part of the cross section, have occasionally
been found for more anterior segments.
Its location is near the border of areas 81
and 85, it was found 12 times.
B. Heterolateral interneurons. Inter34
neurons for single heterolateral dorsal segments have also been found. They form a
bundle very near the midline and like the
homolateral fibers, appear to respond to
the sensory input of individual second
roots.
A 29. Heterolateral fiber of second segment dorsal hairs. This fiber is a close
neighbor of A 30 and also large. Location
78-82; it was found 9 times.
A 30. Heterolateral interneuron for dorsal hairs of segment 3. This is also obtained in 2/3 leads as is the previous
3
37
fiber. Large fiber, located in area 78-82
Fig. 9 Dorsal view of abdomen indicating
near the other heterolateral single seg- areas innervated by interneurons covering areas
ment fibers. Found 10 times.
of two segments.

219

CRAYFISH ABDOMINAL CORD NEURONS

A number of interneurons integrate the

impulses from two abdominal segments.
None of these nor any of the single segment interneurons were found in the commissure. (A 32 might be considered as
the last of this series but its presence in
the commissure as C 3 is against this
view ) .
A 40. Homolateral dorsal aspect of
second and third abdominal segment. Location 83, medium in size, this fiber was
found 9 times.
A 38. Homolateral dorsal aspect of abdominal segments three and 4. This medium sized fiber was found 14 times.
Location 79.
A 58. Homolateral dorsal aspect of 4th
and 5th segments, medium in size. Location 85. Found 8 times.
A 57. Homolateral dorsal aspect of 5th
and uropod segments, medium in size,
location 85. It was found 8 times, close
to and also in the same preparation as
A 56.
A 34. This is the only heterolateral
dorsal hair fiber established for two segments, namely second and third, but it
might also react to segment 1. It is a
medium sized fiber located in area 80
and was found 10 times.
Unlike the interneurons for single segments, these two segment interneurons do
not run in bundles (fig. 8).
Interneurons reacting to three or more
dorsal segment hairs (figs. 10, 11). In
contrast to the single and two segment
fibers, many of these have also been found
in the commissure.
A 63. Homolateral dorsal hairs of segments 2, 3 and 4. Located in area 84,
medium in size, and found 5 times. It is
identical with C 45.
A 12. Homolateral dorsal hairs of abdominal segments 3, 4, and 5. Identical
with C 65. A large fiber in area 83, it was
found 16 times.
A 7. Homolateral dorsal hairs on abdominal segments 4, 5 and uropod segment, not on telson or uropod. It is identical with C 91. It is a large fiber located
in area 82, and found 17 times.
A 41. Homolateral dorsal hairs of abdominal segments l?,2-5th segment, not
on telson or uropods. Medium in size, it
is identical with C 71. It was found 9
times, location 79 (see fig. 12).

A 73. Homolateral dorsal hair fiber of

abdominal segments 4, 5 and uropod segment, uropod and telson. Identical with
C 56, medium in size, location 83. Found
4 times (see fig. 12).
In addition to these homolateral fibers,
some were found with a heterolateral and
a bilateral distribution :
A 54. Heterolateral dorsal hairs of all
abdominal segments, 3-5 for certain but
possibly also two and uropod segment.

A 71

Fig. 10 Cross section with location of interneurons covering three or more adjacent abdominal segments, dorsal hairs; 0 , bilateral interneurons.

HOMOLATERAL

HETEROLATERAL

I l l

54

Fig. 11 Dorsal view showing areas innervated

by interneurons responding to dorsal hair touch
of three adjacent abdominal segments.

220

HOMOLATERAL

C. A. G . WIERSMA AND G . M. HUGHES

BILATERAL

71

Fig. 12 Dorsal view showing areas innervated

by interneurons responding to more than three
adjacent abdominal segments.

There might be two rather similar fibers

in this area. Location 78, found 9 times,
mostly restricted to segments 2-5.
A 71. Bilateral interneuron for hairs
on abdominal segments 3, 4 and 5. Location in area 80 and found 7 times, of
medium size (see fig. 12).
A 64. Bilateral interneuron for dorsal
hairs of the whole abdomen. This medium sized fiber is most likely identical
with one of the several fibers in the commissure, each of which covers this area
in addition to others. Location 84, found
8 times, of which in a few cases the dorsal aspect of the uropods themselves did
not give a reaction.
Dorsal hairs from telson and uropods
(fig. 13). The interneurons specific for
these areas are separately treated because
it is difficult to compare the telson with
the abdominal segments and the function
of the uropods is specialized. Nevertheless some of these interneurons may be
members of one of the previous series.
A 3. Homolateral dorsal hair fiber of
telson and uropod, not on uropod segment.
This fiber is usually found in area 82.
However, in a number of cases a similar
fiber was found in more lateral areas, but
then never also in area 82. It seems possible, therefore, that A 3s localization is
more variable than that of other interneurons. It was found 21 times, of which

4 times in the same preparation as A 6.

It was located 11 times in the medial and
7 times in the lateral half of the cord and
twice near the midline. Like A 6 it is a
large and sensitive axon. It is identical
with C 19 (see fig. 9).
A 32. Homolateral dorsal hair fiber of
telson, uropod and uropod segment. Located quite consistently in area 80, near
the center of the connective. This fiber
cannot be confused with A 6, but could
with A 3. However, they have been found
twice in the same preparation. This fiber
is large, and was found 11 times. It is
identical in reaction to C 3 of the commissure fibers (see fig. 11).
A 6. This fiber may be called Prossers
fiber, as it reacts typically in the way
described in his experiments (Prosser,
35), which showed the relation between
touch of hairs on the fringe of the telson
or uropod and the firing of an interneuron.
It was found to fire for all those homolateral hairs which form the fringes of
the telson and the uropod, whereas in
some cases but not in others, it also fired
readily when the frontal edge of the anterior rim of the uropod was touched.
Of high frequency discharge and often
very sensitive, the fiber can also be made
to fire by bending the outer flap of the
endopodite of the uropod or the proximal
flap of the telson, which may be due to
the stretch of the primary hair fibers. It

Fig. 13 Cross section with locations of interneurons responding to hair touch on dorsal
uropod and telson areas. Note that A3 is represented twice (explanation see text).

22 1

CRAYFISH ABDOMINAL CORD NEURONS

is a large axon, in area 85, found 18

times, of which some were in 2/3 leads
and also 4/5 leads. Its characteristic difference from A 3 is that the latter does
not readily respond to touch of the fringe
hairs, but only of the dorsal hairs of these
parts.
A 37. Heterolateral uropod and telson
hair fiber, responds to both dorsal and
ventral aspects. Of medium size, its location is area 81. It was found 16 times,
in a few of which either the dorsal or the
ventral side was insensitive (see fig. 9).
A 17. Responds to hairs on the middle
dorsal aspect of the anterior part of the
telson. It is identical with C 15 of the
commissure, but responds most likely to
homolateral stimulation. The fiber is rather
large, located in area 83, and often gives
a tonic discharge which is inhibited by
touch of dorsal hairs on both uropods and
uropod segment, which was also true of
C 15. It was found 9 times (see fig. 7).
Summary. The outstanding feature of
this section is the observed abundance of
interneurons representing the dorsal abdominal hairs. The representation of
these hairs by interneurons is as great as
that of the claw. For instance, stimulation
of dorsal hairs on the 4th abdominal segment results in the excitation of 14 interneurons in the cord as against 11 for the
claw in the commissure. The presence of
so many interneurons does seem to indicate a complex set of reflexes, which differ
with the extent of the touch of the abdominal segments. But, besides the fact
that such reactions have so far not been
noticed, it seems highly unlikely that they
would serve any good purpose. Therefore,
the question of redundancy of information
on stimulation of single areas is here
especially poignant, and a further study
using whole animals should throw light
on this problem.
Interneurons responding to ventral
areas of the abdominal segments (fig. 14
15). Compared to the abundance of the
representation of the dorsal areas, few
interneurons integrate the stimulation of
ventral hairs.
A 55. Reacts to hairs on both pleural
plate and swimmerets of segments 2 and
3. It is a small fiber located in area 85,
and was found 7 times.

A 51. A large interneuron which is

fired by stimulation of the posterior pleural
plate hairs of the third segment and the
anterior ones of the 4th. It is thus restricted to the first root neural segment.
Location 84, found 9 times.
A 67. Responds only to stimulation of
the hairs on the third swimmerets, not
on pleural plate 1area. This is naturally
a descending fiber, which is rather small
A 59

52

Fig. 14 Cross section with locations of interneurons responding to hairs on the ventral aspects of the abdomen.

HOMOLATERAL

ASYMMETRIC

19

Fig. 15 Ventral view of abdomen indicating

areas for some of the interneurons responsive to
touch of hairs.

222

C . A. G. WIERSMA AND G. M. HUGHES

and located in area 76 and found 4 times.

In the same area an ascending interneuron for the 4th swimmeret hairs has been
found twice.
A 59. Responds to ventral hairs on the
homolateral uropod and telson surfaces.
Medium in size, located in area 77; it has
been found 5 times.
The absence of ascending fibers from
the more posterior swimmerets may indicate that this type of information is carried mainly in a caudal direction. The
following 4 fibers responded to stimulation
of ventral hairs on more than one segment.
A 13. Homolateral pleural plate fiber
of the abdominal segments. This large
fiber, located in area 85, is made to fire
exclusively by the hairs on the outer edge
of the pleural plates. Found 20 times.
It does not usually respond to the relatively few pleural hairs of the uropod segment, but in some cases did so.
A 19. Similar to A 13, but in addition
responds to stimulation of the other ventral hairs, including those of the swimmerets of abdominal segments 2-5. In
some instances, this medium sized fiber
also responded to stimulation of the heterolateral swimmeret and pleural plate 1
area of the third segment but never to any
others. It was found 20 times, on 5 of
which the heterolateral pleural plate response was present. Even so it is not yet
possible to decide whether there are two
fibers which differ in this one respect, or
whether the same fiber can have both
response types. It is unlikely that the
heterolateral pleural plate response is due
to an additional neighboring fiber with a
signal of the same strength.
A 28. A medium sized fiber which responds to the homolateral first root hair
areas of abdominal segments 2-5 and in
addition to hairs on the homolateral uropod, both ventral and dorsal, but not on
the telson or dorsal uropod segment. Located in area 84, it was found 13 times,
once in a whole preparation, when it also
responded to the thoracic leg hairs. It is
undoubtedly a fiber with a larger sensory
area than given here.
A 11. A bilateral fiber for swimmeret
and pleural plate I hairs, it responds to
stimulation of all abdominal swimmerets,
but not to uropods. A medium sized fiber

found 17 times in location 84. In anterior recordings it sometimes continues to

respond to all of its sensory areas, in other
cases only to those in front. Similarly in
posterior recordings responses to the
whole sensory field or only segments 4-5
may persist.
Interneurons activated by both dorsal
and ventral hairs (fig. 14). Two such
fibers were found, one homolateral and
the other bilateral.
A 46. Homolateral dorsal and ventral
hairs of abdominal segments one-uropod
segment, not on telson or uropod. Located in area 83, it was found 9 times,
and was of medium size.
A 52 responds rather tonically on stimulation of the whole of the dorsal and
ventral aspects of abdominal segments
3-5 and perhaps uropod segment. Located near the stretch receptor fibers in
area 78, it was found 8 times, and was
of medium size.
Summary. These results give the general impression that the ventral abdominal
surface is relatively unimportant as an
area for touch perception. This may be
related to the fact that this surface will
be constantly agitated by swimmeret movements, and often by touch of the bottom.
An exception is the pleural plate 1 area,
which is better represented, and would
also be less subject to stimulation by
swimmeret movements. Even so, the contrast between the dorsal and ventral sides
remains unexpectedly large. The fact that
a certain part of the ventral exoskeleton
was removed in order to expose the cord,
cannot have played a significant role in
this respect, as was shown by some control experiments in which the area removed was limited to fewer segments.
Some of these fibers may play a role
in the co-ordination of the metachronal
rhythm of the swimmerets.
Note that fibers responding to both dorsal and ventral hairs may be identical with
some of the commissure interneurons
which cover large body areas, but that
the evidence is insufficient to decide which
ones they might be.
Interneurons integrating proprioceptive
impulses (fig. 16). A single integrative
interneuron has been found for the abdominal stretch receptors, which is stim-

CRAYFISH ABDOMINAL CORD NEURONS

223

of the heterolateral uropod and was noticed 6 times. It is medium in size

(fig. 17).
A 70 is a large fiber which fires when
either uropod is spread. Located in area
79, and found 7 times (fig. 17).
For the anal valves two special fibers
70 have been found:
A 24
A 43. Homolateral anal valve fiber.
Small, in area 82 and found 6 times, it
A 35
often accompanied A 5, the asymmetric
joint fiber (fig. 17). A 72 is a much
larger fiber in area 76 for the heterolateral
anal valve, found 5 times. Both A 43 and
43 A 72 discharge when the valve is pulled
outward, and not on mere touch (fig. 17).
As mentioned previously there are two
A 2 0
I
\
A5
fibers for homolateral swimmerets, which
A 10
A 27
may be primary but fire only during active
Fig. 16 Cross section with location of interneurons responsive to joint movements; 8, asym- movement. A 35 is medium in size and
metric responding interneuron.
is located in area 81. It fires while the
5th swimmeret is in a backward position
ulated by the descending branches of the and is inhibited by stimulation of the
primary fibers in the 6th ganglion (Hughes dorsal hairs on the same segment. It
and Wiersma, '60a), the others integrate was found 8 times. A 31 similarly represwimmeret, uropod and anal valve ac- sents the 4th swimmeret and is a neightivity.
bor of A 35 in the same area 81, is meA 61 is medium in size, fires at fre- dium in size, and was obtained 16 times.
quencies lower than the tonic stretch receptors feeding into it and is located in
HOMOLATERAL HETEROLATERAL
area 77. It was found 4 times, whereas
once a similar fiber was found in the
same preparation for the phasic stretch
receptors.
A 20. Homolateral uropod, peripheral
joints fiber. This fiber may integrate the
sensory input from different joints of the
uropod and can also be activated by flexion of the proximal part of the telson.
However, such stimuli might elicit reflex
contraction of a muscle receptor organ,
hence the uncertainty about its nature,
primary sensory or interneuron, in this
and following cases in which such a mechanism might occur. Medium in size, this
fiber was found 14 times, and in most
cases gave a tonic discharge. Its location
is in area 84 (fig. 17).
A 68. A similar medium sized fiber,
but exclusively for the basal joint of the
homolateral uropod. It was found 5 times
in area 77, and might well be primary
(fig. 17).
A 69, also in area 77, is a partner of
Fig. 17 Ventral view indicating joints to
A 68, reacting to the same movements which different interneurons respond.

224

C. A. G . WIERSMA AND G . M . HUGHES

There are three interneurons which are

brought to discharge by movements of all
homolateral swimmerets.
A 27 is restricted to the basal joints of
the homolateral swimmerets, and in contradistinction to the two following fibers,
it is not excited by stimulation of the
anal valve. As in the two following fibers,
the discharge is tonic. Located in area 83,
medium in size, it was found 11 times.
In males this fiber does not seem to be
excited by the second swimmeret which
it is in females (fig. 17).
A 10 is a medium sized fiber in area 84,
which besides responding to swimmeret
joints responds to the basal joint of the
homolateral uropod and often the anal
valve. It is sometimes difficult to distinguish from the next fiber, when the latter
does not respond to the heterolateral
uropod as sometimes happens. Usually
though, the difference in location is sufficient to decide which fiber is being recorded from. It was found 13 times
(fig. 17).
A 5 is the only fiber which regularly
gave an asymmetric response, reacting to
all homolateral swimmerets, homolateral
uropod, to the heterolateral uropod and
both anal valves. It is large and very
characteristically located in the medial
ventral aspect of area 82. It is one of the
most readily found fibers, registered 26
times and noted many other times. It
appears to be identical with C 48, as was
practically proven in a whole animal preparation in which it also responded to stimulation of the homolateral thoracic legs.
Furthermore, this same fiber was prepared
also in area 70 of the commissure in this
preparation.
A 24 is a medium sized bilateral joint
fiber, identical with C 46, as found in a
whole animal preparation in which joint
movements of all head and body appendages made this fiber fire. It was found
5 times, two of them in a whole animal.
Location in area 81.
Interneurons integrating both hair and
joint monement (fig. 18). The three fibers showing this type of reaction all responded to the sensory area innervated
by the first root of a single abdominal
ganglion.

A75
Fig. 18 Cross section with location of the
interneurons with an integrative response to both
hair touch and joint movement.

A 33 is a medium-sized fiber in area 81

for the homolateral third swimmeret and
pleural plate 1. It was found 8 times.
A 75 is a similar fiber for the homolatera1 5th swimmeret and pleural plate
1. It was found 5 times in area 84.
A 66 is a similar fiber for the heterolateral third swimmeret and pleural plate
1,located in area 76 and found 5 times.
Similar fibers for other swimmerets are
indicated, but not yet established.
Summary. The number of interneurons from joint fibers is relatively small
in the cord as compared to the cornmissure. The approximate percentages of the
total number of interneurons are 20% for
the cord as against 40% for the commissure. This may be explained on the assumption that the swimmeret joints signal
their positions to integrative areas much
more by the primary sensory fibers than
do other appendages. Thus the mutual
co-ordination of the swimmerets, whose
only really significant motion for the animal as a whole may be the metachronal
beat, is largely regulated by the primary
fibers. The main driving of the waves
is done by interneurons (Hughes and
Wiersma, 60b), so that only smaller adjustments would have to be performed.
The uropods are better represented, and
i t is known that several of these interneurons reach the brain, indicating their
greater importance for general co-ordination of the body.

225

CRAYFISH ABDOMINAL CORD NEURONS

DISCUSSION

There are a number of differences between the results of this analysis of fibers
in the abdominal connective and that
obtained by similar methods for the circumesophageal commissure. Some of these
are due to experimental circumstances
but a more significant factor may be the
far greater degree of local traffic in the
cord than in the commissure. The many
primary sensory fibers found in the cord
are indicative of this difference. It is certain that in the commissure very few
descending primary sensory fibers are
present, whereas such fibers are abundant
in the cord. The situation is less clear
for ascending primary fibers since these
are represented in the commissure by the
many fibers from dorsal carapace hairs,
whereas others which might be present,
namely those from mouthparts would have
been overlooked because of the experimental limitations. In the cord it is quite
certain from both physiological and morphological evidence, that many of these
fibers are thin. Thus when the locations
of these bundles are superimposed on a

200p

150

100

50

microphotograph of a cross section of the

cord, there is a good correlation between
the position of such primary sensory fiber
bundles as A 9, A 18 or A 26 and the
presence of fine fibers in the same areas
of 84 and 82. It is strongly indicated,
therefore, that all really small fibers are
primary sensory ones (but not the converse, for the stretch receptor fibers are
large as may be other primary proprioceptor fibers). If this is so, and since
there are no indications that any interneuron is really small (below 10 cl), the total
percentage of the larger fibers accounted
for in the cord is quite considerable.
As can be seen in figure 19, out of a
total of about 1200 fibers only some 370
are more than 10 p in diameter. About
68 of these now have a known function so
that without considering any other fibers
20% would be accounted for, as against
only 7% on the same basis in the commissure. If all entities from which single
unit potentials were regularly obtained
were above 20 c1 in diameter, then 50%
of the population sampled by these methods will have been described physiologi-

50

IM)

150

Fig. 19 Relation between number of fibers (logarithmic Y-axis) and fiber diameters in
10 p classes as counted in the left and in the right cross section of the abdomnial connectives
of Procambarus clarkii on a photograph made by Dr. J. D. Robinson. Note the definite break
in size between the two giant fibers and the largest ordinary fibers. Also note that the
smallest fibers are more numerous than all other classes combined. Each half was counted
independently.

200p

226

C. A. G . WIERSMA AND G. M. HUGHES

cally, since there are about 140 fibers

in that class. This figure is surprisingly
high in relation to the other types of interneurons which must also be present.
These are the descending interneurons
from the thorax, the spontaneously active
fibers in the cord (which are fewer than
in the commissure, perhaps because all
those from the higher part of the central
nervous system are, of course, missing),
those reacting to other sensory modalities
exclusively, and finally those which
though reacting, have not yet been established. It is certain that a very high proportion of the entities that can be found
with the stimuli used has in fact been
found. For in our later experiments, the
number of fibers which could be designated became the great majority (90%
or more) of all the fibers reacting in these
preparations, a situation which was not
yet present in the commissure, where at
best one half of such fibers could be accounted for.
If all the interneurons are indeed larger
units, then any additional small fibers
which may be revealed by electromicroscopy (0.2-0.7 ~1 diameter . . J. H. McAlear,
personal communication) will not affect
the present issue. Larger numbers of small
fibers may well be expected because of the
considerable numbers of sensory fibers in
the first and second roots, most of which
may run either up or down or both ways
for varying distances after entering the
cord.
It is our view that the primary sensory
fibers of the first root inflow play an important part in the co-ordination of the
swimmeret movements (see Hughes and
Wiersma, '60b). But though this can account for the interganglionic pathways of
the first root fibers, it does not explain
that of the second root inflow, especially
of the dorsal hair fibers. In view of the
fact that for this particular area there
are also many interneurons transmitting
derived information, it is quite astonishing
that such extensive primary fiber branching should be present.
Though it is certain that many primary
fibers have a restricted path length in the
cord, the stretch receptor axons show that
such fibers can run the full length of the
central nervous system. At present, there
is no direct evidence that any others do

so. But if those fibers which fire only

during active backward contraction of individual swimmerets are from muscle
sense organs of the type described by
Alexandrowicz for the thoracic legs, finding them at distances as much as three
segments from their origin may indicate
that they, like the abdominal stretch receptors they resemble, also run for long
distances. They would then most likely
be of considerable importance in the general co-ordination of movements of the
body appendages. However, if they are
interneurons this reasoning would not be
valid.
It is of interest to compare the number
of fibers which have been found to react
identically in the cord and the commissure, with the number which might have
been expected from the previous work on
the commissure. In this paper a total of
16 have been shown to be identical out of
a possible total of 32. In addition there
were about 9 fibers found which either
could not definitely be shown to be identical or were found too few times to be
established in the cord. Thus by far the
major portion of expected fibers has been
accounted for. It remains uncertain
whether the other 7 are absent from the
cord because of secondary integration
(e.g., in the thoracic ganglia) or that
further investigation will show their
presence.
In the cord, as in the commissure, the
great majority of interneurons integrate
sensations of a single modality. Because
of the larger number of elements accounted for in the cord, the conclusion
that this predominance is a real feature of
the crayfish nervous system, is greatly
enhanced. The evidence presented here
for interneurons that integrate different
modalities depends on very few cases and
examples of fibers which have been found
relatively few times, but Kennedy and
Preston ('60) also found this type of interneuron at the cord level. It is to be
noted that in the commissure fibers integrating hair and proprioceptive sensation
have been encountered which, as in the
cord, integrate the sensations of only a
single appendage.
Tract formation may be briefly discussed on a comparative basis. As in the
commissure, the primary sensory fibers of

CRAYFISH ABDOMINAL CORD NEURONS

one modality form regular tracts, whether

they run long distances (stretch receptors)
or short ones (A 26). Furthermore, when
the same type of fiber runs both forward
and backward, tracts at about the same
location in the cross section are present,
which immediately adjoin tracts of similar sensory fibers from other segments.
Results in the cord, show that a tract
may also be formed by interneurons of
a certain type. In the commissure it has
already been noted that the interneurons
which reacted to stimulation of hairs on
the homolateral thoracic appendages, run
together in a tract. For axons responsive
to a sensory field in a single segment,
similar tracts have been noted in the
abdominal connective for two sets of homolateral interneurons reacting to dorsal
hairs (A 14, 15, 39, 62 and A 42, 49, 50,
56), as well as for heterolateral interneurons from these areas (A 22, 24, 30,
65).
On the other hand interneurons representing two or more segments, though
similarly related among themselves, do
not show this type of tract formation.
Possibly this indicates that the impulses
from them are not integrated with each
other at a higher level, whereas those in
tracts are.
The distribution of the fibers found in
both commissure and cord can be compared. Their relative locations generally
show a fair correspondence, but in a few
cases fibers are in quite different quadrants. Whether or not this is because a
mistake has been made or that such fibers
do shift in relative position, is not certain, but the latter possibility appears the
more likely at present.
With regard to the nature of the sensory representation, here as in the commissure all primary sensory fibers appear
to be homolateral. Homolateral interneurons are again in the great majority, but
bilateral and heterolateral representation
is present. Of 92 interneurons of the commissure, 63 were homolateral, 10 heterolateral, 16 bilateral and 3 asymmetric. In
the cord these figures were of 57 interneurons: homolateral 39 (38), heterolatera1 10, bilateral 7 and asymmetric 1 (2).
Thus the total percentage of homolateral
interneurons is about the same (68% ) in

227

both locations. This contradicts Prosser's

belief ('35) that contralateral representation would be absent or slight in the cord.
SUMMARY

1. The electrical activity of single units

has been recorded from small bundles obtained by splitting the connectives between abdominal ganglia of the crayfish.
Responses to mechanical stimulation of
the abdomen, either to hairs or by movement of joints were investigated and the
location of the responding units in the
cross section noted.
2. An account is given of the location
and properties of 75 such entities in the
connective between the third and 4th abdominal ganglia, which were found a sufficient number of times to justify their
establishment.
3. Several of these represent bundles of
primary sensory fibers from a single area
of the body. They may pass up or down
the cord or in both directions from the
ganglion which they enter. All of these
remain on the same side (homolateral)
but vary in the extent of their spread.
This is maximal for fibers from the abdominal stretch receptors (SR1 and SR2),
which send one branch to the brain and
another to the last abdominal ganglion.
4. Hair fibers entering through first
roots have very short intracentral pathways but those from first root proprioceptors and second root hair fibers ascend
across at least three interganglionic
spaces.
5. Interneurons integrate inputs from
single segments or homologous areas of
several adjacent segments which may be
homolater a1, he terolater a1, bil ater a1, or
asymmetrical in their distribution with respect to the leading off position.
6. Many have properties identical with
units identified in the circumesophageal
commissure but interneurons responsive
to dorsal hairs on a single or two segments
are a conspicuous exception, for no such
fibers have been established in the commissure.
7. The great multiplicity of the central
representation of a given peripheral field
is one of the most striking features of this
study. Its significance will only be fully
understood when the efferent connections
of these interneurons are investigated.

228

C . A. G . WIERSMA AND G.

LITERATURE CITED
Alexandrowicz, J. S. 1951 Muscle receptor organs in the abdomen of Homarus vulgaris and
Palinurus vulgaris. Quart. J. Micr. Sci., 92:
163-199.
1958 Further observations on proprioceptors in Crustacea and a hypothesis about
their function. J. Mar. Biol. Assoc. U. K., 37:
379-396.
Allen, E. D. 1894 Studies on the nervous system of Crustacea. I. Some nerve elements of
the embryonic lobster. Quart. J. Micr. Sci., 36:
461-482.
Hughes, G. M.,and C. A. G. Wiersma 1960a
Neuronal pathways and synaptic connexions in
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Biol., 37: 291-307.
1960b The co-ordination of swimmeret
movements in the crayfish, Procambarus clarkii
(Girard). Ibid., 37: 657-670.

M. HUGHES

Kennedy, D., and J. B. Preston 1960 Activity

patterns of interneurons in the caudal ganglion
of the crayikh. J. Gen. Physiol., 43: 655-670.
Prosser, C. L. 1935 Action potentials in the
nervous system of the crayfish. 111. Central
responses to proprioceptive and tactile stimulation. J. Comp. Neur., 62: 495-505.
Retzius, G. 1890 Zur Kenntniss des Nervensystems der Crustaceen. Biol. Untersuch, N. F.,
I: 1-50. Central. Druck., Stockholm.
Van Harreveld, A. 1936 A physiological solution for fresh-water crustaceans. Proc. SOC.
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Wiersma, C. A. G. 1958 On the functional
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Girard. J. Comp. Neur., 110: 421471.
Wiersma, C. A. G., and R. L. C. Pilgrim 1961
Thoracic stretch receptors in crayfish and rocklobster. Comp. Biochem. Physiol., 2: 51-64.