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Participation of a sialic acid-specific lectin from freshwater prawnMacrobrachium rosenbergii hemocytes in the recognition of non-self cells

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JEZ 842
THE JOURNAL OF EXPERIMENTAL ZOOLOGY 279:243–253 (1997)
Muscle Receptor Organs of the Crayfish, Cherax
destructor: Organisation of Central Projections of
Stretch Receptor Neurons
DAVID L. MACMILLAN* AND PAUL J. VESCOVI
Department of Zoology, University of Melbourne, Parkville Vic. 3052,
Australia
ABSTRACT
The stretch receptor neurons of the abdominal muscle receptor organs of crayfish
enter the ventral nerve cord and branch to send axons to the brain and to the last abdominal
ganglion. They mediate local reflexes in the ganglion of entry and adjacent ganglia but evidence
from previous cobalt filling and histology suggested that local branching is limited and variable
and does not accord closely with physiological evidence. We examined the fine structure of the
branching in a large number of preparations from large animals of the crayfish Cherax destructor,
which is also a large species, to determine whether the increased resolution afford by size would
permit us to detect previously undetected patterns. We found some predictable features in the
pattern of fine branching in the ganglion of entry and adjacent ganglia that could explain some of
the apparent anomalies. We also examined the relative positions of the hook-shaped projections
from different segments where they terminate in the last abdominal ganglion by differentially
staining the stretch receptors with Co++ and Ni++. We found evidence for somatotopic organisation
in the longitudinal position of the endings. J. Exp. Zool. 279:243–253, 1997. © 1997 Wiley-Liss, Inc.
Allen (1864) first described some simple sensory
cells which were later named muscle receptor organs (MROs) by Alexandrowicz (’51) in his studies
on the abdomen of the lobsters Homarus vulgaris
and Palinurus vulgaris. Since Alexandrowicz’s description there have been numerous studies carried out on the microanatomy, neurophysiology,
and neuropharmacology of the MRO. MROs have
since been identified in more than ten species of
decapod crustacean (Pilgrim, ’60; Fields, ’76). Each
MRO consists of a stretch receptor neuron (SR)
with its dendrites embedded in a thin muscle
bundle in parallel with the dorsal extensor musculature. They are found in pairs on either side
of the midline in each abdominal segment. The
MROs that make up a pair have distinct anatomical and physiological characteristics and respond
to different features of abdominal movement. The
medial organ has a SR with a broad dendritic area
and responds tonically to maintained stretch
(MRO1 and SR1). The lateral receptor organ has
a shorter, thinner muscle and its conically-shaped
SR responds phasically to maintained stretch
(MRO2 and SR2).
There is a considerable amount of evidence from
extracellular studies that the SRs are involved
in a number of local reflexes (Eckert, ’61; Fields
and Kennedy, ’65; Fields, ’66; Fields et al., ’67;
© 1997 WILEY-LISS, INC.
Sokolove, ’73; Nja and Walloe, ’75) involving the
ganglion of entry and adjacent ganglia. These appear to be present in the range of crayfish species
studied (for review see Fields, ’76) and some of
them have been confirmed and further investigated with intracellular techniques (Hausknecht,
’96, and in preparation). One might have predicted, therefore, that there would be clear and
consistent patterns in the branching and projections of the SRs in these ganglia. Histological studies of the abdominal ganglia of Procambarus
clarkii failed to find substantial branches (Liese
et al., ’87; Skinner, ’85a,b) or stereotyped projection patterns. Bastiani and Mulloney (’88a) also
examined this aspect of the SRs in their study of
the morphology of the projections anteriorly and
posteriorly in the cord. They stained the SRs by
cobalt infusion into the cut peripheral ends of the
axons and noted a number of small branches in
the ganglion of entry and in other ganglia through
which the axons passed but could find no consistency in the position or shape of those branches.
While this result was suggestive, it was not con-
*Correspondence to: Dr. D.L. Macmillan, Department of Zoology,
University of Melbourne, Parkville, Vic. 3052, Australia. E-mail:
zoomac@ariel.ucs.unimelb.edu.au
Received 15 November 1996; revision accepted 10 June 1997.
244
D.L. MACMILLAN AND P.J. VESCOVI
clusive because this aspect of the work was subordinate to the main aim of the project which was
to visualise the terminations. To achieve this they
used small animals (2–3 cm) which meant that
details of the fine branching in the ganglia in between were difficult to determine and may not represent the mature condition. One of the objectives
of the experiments described here was to use large
animals and target the local ganglia specifically
so that all details of even the finest branches
would be apparent following intensification of the
cobalt filled preparations. The SRs from each
MRO pair enter the ventral nerve cord in the second root of the next anterior segmental ganglion.
Bastiani and Mulloney (’88a) demonstrated that
Alexandrowicz’s (’51) conjecture about the possible
fate of the SR axons was correct. Upon entering
the ganglion, the sensory axons bifurcate and run
in opposite directions. They demonstrated further
that one branch travels all the way to the brain
and the other to the last abdominal ganglion, A6.
A6 is the major integrating centre for the motor
system controlling the various appendages that
comprise the tailfan (Dumont and Wine, ’87a,b).
The sensory axons of the SRs travel the entire
length of the body, passing through every ganglion
and running together in a single bundle in the
dorsal medial tract (DMT) (Wiersma and Hughes,
’61). The sensory axons from all the abdominal
muscle receptors terminate in the same dorsal
area of A6 with a characteristic, hooked structure
that is symmetrically disposed across the midline.
The hook is formed near the posterior end of the
interconnective fissure where the axons turn medially across the midline in commissures A6 DCII
or A7 DCII (terminology from Kondoh and Hisada,
’86) and, after crossing the midline, run anteriorly
to the end of the core of A6.
This labelled-line input and extensive branching in A6 suggests that MRO input plays some
important role in the tailfan control. The tailfan
is a highly specialised terminal structure in the
crayfish, formed by a fusion of the telson and the
paired appendages of the sixth abdominal segment, the uropods. Many of the neurons in the
terminal ganglion and muscles of the tailfan have
been characterised morphologically and physiologically and some of their functional relationships have
been determined (Larimer and Kennedy, ’69; Wine
and Krasne, ’82; Hisada et al., ’84a,b; Wine, ’84;
Dumont and Wine, ’87a,b; Kondoh and Hisada,
’87; Takahata and Hisada, ’85; Vescovi et al., ’97).
Bastiani and Mulloney (’88a) recorded intracellularly from a number of neurons of different
classes in A6 that were postsynaptic to SR axons
in P. clarkii. They found local interneurons, plurisegmental interneurons and motor neurons that
responded to SR activation. Vescovi et al. (’97)
identified some of those neurons in Cherax destructor. Bastiani and Mulloney (’88b) also found
that SR input from posterior abdominal segments
was more likely to cause larger EPSPs in unidentified terminal ganglion neurons than input from
anterior segments of P. clarkii and demonstrated
evidence of an anterior-posterior gradient in the
strength of the response. Vescovi et al. (’97) found
that similar gradients occur in C. destructor, but
not in any of the neurons they identified. Studies
in insects have demonstrated some highly ordered
projections from mechanosensory afferents, with
somatotopic relationships between the location of
the sensory structure and the projection in the
central nervous system (Teugels and Ghysen, ’83;
Johnson and Murphey, ’85; Kent and Levine, ’88;
Murphey et al., ’89). We hypothesized that the
physiological gradients in some postsynaptic cells
in both P. clarkii and C. destructor could be related to some kind of somatotopic order of the SR
endings in A6, perhaps echoing the topological
organisation of hair projections from different tailfan nerves into the last abdominal ganglion described by Kondoh and Hisada (’87). The search
for evidence for such organisation was a second
objective of this study.
MATERIALS AND METHODS
Australian freshwater crayfish, Cherax destructor, from the Murray River system were obtained
from a commercial supplier and kept indoors at
22°C in polystyrene aquaria. They remained active and in good condition for long periods when
provided with shelters, normal light-dark cycles
and a change of water following a weekly feeding
with dry-pellet cat food. Seventy five inter-moult
specimens of both sexes were used in the experiments. Specimens with a carapace length of 6–10
cm were used. Animals were anaesthetised by immersion in ice for ten minutes and the brain destroyed. Specimens were immobilised and dissected
to expose the nerves to be stained.
Details of SR morphology were visualised by infusing CoCl2 intracellularly through the cut ends
of the appropriate axons for periods of up to 24
hours. Some of the infusions were done in whole
preparations, others were done in vivo by dissecting nerves and ganglia free and pinning the preparations in a Sylgard-lined (Dow-Corning, Midland,
MI) dish flooded with physiological saline (modi-
CRAYFISH ABDOMINAL STRETCH RECEPTOR PROJECTIONS
fied van Harreveld’s (’36) solution: 11.68 g/l NaCl;
0.40 g/l KCl; 0.55 g/l MgCl2.6H2O; 1.51 g/l Trisma;
made up in distilled H2O, pH = 7.4; developed by
Pasztor and Macmillan, ’90). In most cases, 1.5%
CoCl2 was used but a range of concentrations up
to 5% CoCl2 solution was used to reduce the probability of missing details of fine axonal and dendritic branches because of factors associated with
ionic concentration and penetration. The time and
temperature were varied to achieve a balance that
best suited the part of the neuronal morphology
under examination. Once determined, a particular regime was used over and over to examine any
area in detail because typically a regime designed
for one purpose was unsuitable for others e.g., a
regime to show the detail of the endings in A6
produces an overstained preparation in the ganglion of entry. A typical filling regime would be
10–12 hours at 4°C. The cobalt ions were precipitated with 0.1% ammonium sulphide (four drops
per 10 ml) and left for 10 minutes. The tissue was
then washed in several changes of saline over a
period of 30 minutes and fixed in Bouins for 2
hours. The preparation was dehydrated and
cleared and mounted in methyl salicylate. Some
preparations were also silver intensified using the
Timm’s method (Bacon and Altman, ’77; Altman
and Tyrer, ’80; Altman, ’81) before being cleared
and mounted. The mounted preparations were
drawn using a drawing tube attached to a Zeiss
compound microscope (Thornwood, NY). To improve our ability to differentiate between SRs from
different segments, in some experiments we used
NiCl2 to fill the SR from one segment and CoCl2
to fill from the other. The infused ions were then
precipitated with Rubeanic Acid to obtain fills with
colour differentiation (Altman, ’81; Jones and
Page, ’83). These preparations were always drawn
before and after intensification because the colour
difference disappears following the silver intensification process.
RESULTS
The general morphology of the abdominal MROs
in C. destructor is essentially the same as that
described in other species (Alexandrowicz, ’51;
Macmillan and Field, ’94; Pilgrim, ’60, ’64). Upon
entering the abdominal ganglion the SRs bifurcate and send a branch to the head and a branch
to A6 as in P. clarkii (Bastiani and Mulloney, ’88a).
In all cases (n = 72), the tonic and phasic SR filled
simultaneously and there were no obvious differences in the point of bifurcation, although the phasic SR usually has a slightly larger diameter than
245
the tonic SR. To determine whether there was an
as yet undisclosed pattern to the fine branching
in the ganglion of entry we conducted a number
of fills (n = 24) of the second nerve of the third abdominal ganglion (A3) in large animals. As in P.
clarkii (Bastiani and Mulloney, ’88a), no branches
are found in consistent positions in C. destructor
and all branches are short and fine. Because of
the larger size of our preparations, we are able to
provide further details of the branching. In C. destructor, the point of bifurcation of the two axons
is invariably very close and there are numerous
fine branches from both axons (Fig. 1). The
branches are all short and run perpendicular to
the main axon with very little branching and are
found along the axons throughout the whole central region of the ganglion from the point of entry
of the first nerve to slightly posterior to the entry
of the second nerve.
We also used large animals (n = 24) to examine
the branching patterns of the SRs in the adjacent
anterior (A2) and posterior (A4) ganglia. There are
always some branches in both the anterior and
posterior ganglion but they occur over a shorter
length of the axons in the central region of the
ganglia. It should be noted in respect of this last
observation, however, that in C. destructor the distance between exit points of the first and second
nerves is commonly greater in A3 than in A2 or
A4. In the anterior ganglion the branches are
similar to those in the ganglion of entry but
there are far fewer of them. In the posterior
ganglion there are also fewer branches but they
commonly give rise to long transverse secondary branches running parallel and in close lateral proximity to the main axon (Fig. 2B). In
many of the experiments designed to reveal SR
branching in the posterior ganglion we also
filled an accessory neuron (Fig. 2B) (Alexandrowicz, ’51, ’67; Wine and Hagiwara, ’77).
The projections of the SRs into the last abdominal segment (A6) in C. destructor (Macmillan and Field, ’94) are closely similar in
position and form to those described in P. clarkii
(Bastiani and Mulloney, ’88a). The hook-shaped
endings of the phasic and tonic units from the
same segment are almost identical and in some
preparations are intertwined (Fig. 3A). We filled
pairs of SRs entering in different nerves of segments A3, A4, and A5 simultaneously to study
the spatial relationship between them in A6 (n
= 48). Simultaneous SR fills from both sides of
segments A3, A4, and A5 showed that the longitudinal position of the hooked part of the pro-
246
D.L. MACMILLAN AND P.J. VESCOVI
Fig. 1. Two examples of branching pattern of the SR axons in the ganglion of entry. Cobalt backfill of SRs into abdominal ganglion A3. The axons pass medially into the
ganglion and then bifurcate to send a branch anteriorly to
the brain and also posteriorly to the last abdominal ganglion,
A6. The phasic and tonic axons branch at the same point.
Both axons give rise to many short fine branches in the region between the first and second nerves but no single branch
is consistent in position or shape.
jection in A6 is always the same so that the pairs
from the same segments are bilaterally symmetrical in this regard (Fig. 3B). Most of the secondary branching also matches closely although the
finest branches show differences. In some preparations in which we filled the second nerves of
A5, we stained a neuron with a contralateral cell
body in A6 (Fig. 3C).
To investigate the relationship between the A6
projections of the SRs from different segments we
filled SRs from different segments simultaneously
with Co++ and Ni++. Although we were technically
able to fill more than two segments simultaneously, we found that we could not interpret the
branching patterns in A6 unambiguously because
of the tangle of projections. We also concentrated
CRAYFISH ABDOMINAL STRETCH RECEPTOR PROJECTIONS
247
Fig. 2. Examples of branching patterns of the SR axons
in the adjacent ganglia. A: Branching pattern in A4 of cobalt
filled SRs entering second nerve of A3. There are typically
fewer branches but some of these give rise to long transverse
secondary branches that lie parallel and close to the main
axons. B: In some preparations we filled accessory neurons
as well as SRs. Branching pattern and accessory neuron in
A4 stained in a fill of the ipsilateral second nerve of A3.
on contralateral pairs because we found the preparations were easier to interpret than ipsilateral
ones and on segments A3, A4, and A5 because to
fill from A1 or A2 it was necessary to use such
small animals that the relationship in A6 could
not be determined with any reliability. The results
reported here are based on 23 successful fills of
combinations of SRs from segments 3 and 4, 4
and 5, and 3 and 5. In all cases examined the
longitudinal position of the transverse part of the
SR hook in A6 reflected the antero-posterior order of the segments of SR origin. The transverse
branches from adjacent segments are 5–10 µ apart
longitudinally (Figs. 4, 5A) and simultaneous fills
of A3 and A5 express a multiple of this spatial
order (Fig. 5B).
248
D.L. MACMILLAN AND P.J. VESCOVI
Fig. 3. Projections of filled SR axons in the last abdominal ganglion (A6). A: Example of the hook-shaped endings of
the phasic and tonic units that enter the ipsilateral second
nerve of A5. Note the closely similar branching pattern in
the phasic and tonic SRs. B: Example of a simultaneous fill
of both pairs of SRs from A5. The transverse branch that
forms the hook is always at the same longitudinal position in
contralateral pairs. In some preparations we stained a neuron with a cell body in A6 when we filled the second nerves
of A5.
CRAYFISH ABDOMINAL STRETCH RECEPTOR PROJECTIONS
249
Fig. 4. Two examples of pairs of filled SR axons from contralateral sides of the adjacent
segments A4 and A5.
DISCUSSION
Early extracellular recordings from abdominal
extensor motor neurons indicated, and later intracellular studies confirmed, that the SRs excite
the motor neurons in the ganglion of entry more
strongly than they excite those in other ganglia
(Eckert, ’61; Wine. ’77; Wine and Hagiwara, ’77;
Hausknecht, ’96 and in preparation). The detailed
250
D.L. MACMILLAN AND P.J. VESCOVI
Fig. 5. Examples of pairs of filled SR axons from contralateral sides of the (A) the adjacent A3 and A4 segments and (B) the A3 and A5 segments which are two segments apart.
CRAYFISH ABDOMINAL STRETCH RECEPTOR PROJECTIONS
information provided to us by Hausknecht (’96)
from her thesis about the morphology of the extensor motor neurons in C. destructor, shows that
most of them, including number 2 (for numbering
details see Kennedy and Takeda, ’65; Fields, ’66;
Sokolove and Tatton, ’75), which responds reflexly,
have long side branches running parallel to the
SR axon through the ganglionic regions close to
the SR axons where the SR short branches lie
(Liese et al., ’87). These branches are thus essentially perpendicular to the short SR branches and
there is a considerable longitudinal overlap where
connections between them or their associated neurons could occur. It may be this aspect of the relationship that explains why neurons that have
been shown to have a close functional relationship (some are probably monosynaptic) do not
have a stereotyped morphological one. It is possible that the precise contact points of the SRs or
their associated neurons on the dendritic tree of
the motor neurons are not important so long as
there are sufficient at the right distance from the
integrating sites. The difference in the number of
fine branches found in this study of C. destructor
in comparison with the study on P. clarkii (Bastiani and Mulloney, ’88a) could reflect species difference or perhaps a difference in the maturity of
the animals used.
Given the physiological background to the SR
motor neuron relationship, our finding that there
are always more fine SR branches in the ganglion
of entry would not be surprising except that a detailed histological study by Liese et al. (’87) found
that the total length of axonal branches from SR
axons was always greatest in the adjacent posterior ganglion. While this could be a species difference between Pacifastacus leniusculus, which they
used, and C. destructor, which we used, some of
the other possible explanations could prove more
interesting. There are typically long secondary
parallel branches in the posterior ganglion of C.
destructor and in any measurement of the total
length of SR branches these might well compensate for any effect that a greater number of short
branches in the ganglion of entry might have.
From morphological analyses in other species
(Liese et al., ’87) and intracellular dye fills of
single neurons in C. destructor (Hausknecht, ’96
and in preparation) we conclude that the accessory neuron that we filled was Accessory 1 (Acc1). We found, like Liese et al. (’87) in their study
of the Acc-1 projections, that the dendritic tree in
the ganglion with the soma is not extensive. In
C. destructor, however, as in the case of the ex-
251
tensor motor neurons, there are again long secondary branches from the main neurite that run
parallel to the long axis of the cord in regions
where the short and long parallel SR branches are
found. In C. destructor the dendritic tree is more
extensive in the ganglion of entry and there are
even more parallel branches (Hausknecht, ’96 and
in preparation) which might be expected given the
strong intra and interganglionic reflex connections
between the SRs and the accessory neurons.
In a number of preparations in which we filled
the second nerve of A5 we stained a neuron with
a soma in A6 (n = 16). The morphology of the neuron is consistent with its being an accessory and,
based on Hausknecht’s (’96 and in preparation)
individual staining of all four accessory neurons
in more anterior ganglia of C. destructor, it resembles Acc-2 most closely. The result was completely unexpected because no accessory neurons
have been found associated with the A5 nerves in
other species in spite of specific searching. Larimer
and Kennedy (’69b) could not detect physiological
evidence for their presence when they looked for
an MRO-accessory reflex associated with the SR
entering A5 in P. clarkii. Wine and Hagiwara (’77)
confirmed that conclusion when they filled the second nerve of A5 stating that “Larimer and Kennedy
concluded that the accessory efferent neuron is
missing in the terminal segment; our anatomical
evidence is consistent with that conclusion.” It is
possible that we have described a species difference, it is also possible that it is a very difficult
neuron to fill from A5 in P. clarkii. It did not fill
in some C. destructor preparations, even ones in
which the SRs projections filled well in the same
time over the same distance. The assumption is
that if this is indeed an accessory neuron, it is an
accessory from the ganglion posterior to A5. But
because A6 is a fused ganglion (Dumont and Wine,
’87a), the simplest supposition would be that it
was originally associated with the anterior primordial ganglion. If this is so, it is surprising that
the soma of the neuron lies so posteriorly in A6,
almost certainly in the domain of the second primordial ganglion. Dumont and Wine (’87a,b; P.
clarkii) and Vescovi and Macmillan (this volume;
C. destructor) both found that the relative position of the motor neurons associated with the two
primordial ganglia are generally maintained. This
anomaly warrants further investigation for what
it may reveal about the evolution of both A6 and
the MRO-accessory complex.
Somatotopic organisation of mechanosensory
projections has been described in a number of situ-
252
D.L. MACMILLAN AND P.J. VESCOVI
ations in insects (e.g., Murphey et al., ’80, ’89;
Johnson and Murphey, ’85). Kondoh and Hisada
(’87) also showed that the projections of the different nerves associated with the last abdominal
ganglion in P. clarkii are topologically organised
but such organisation remains to be documented
at the single cell level in crustaceans. Liese et al.
(’87) looked for evidence for its presence in their
detailed study of sensory projections in crayfish
abdominal ganglia but found none. Our data suggest that the segment of origin of each SR may
determine where its terminal ending is positioned
in A6. The strength of that case is dependent on
a number of considerations. First, we were limited by technical considerations to pairs of neurons, so the argument is based on the finding that
in no case was the projection of an SR from the
more anterior segment ever found posterior to its
filled pair. Second, our assumption of longitudinal position is based on the position of the transverse part of the hook and there is no a priori
reason for arguing that this is representative of
the functional position of the neuron. We analysed other structural features such as prominent
branches, the length between branches, and the
number of secondary branches to see if we could
detect any other markers for longitudinal position
but the variation in the level of secondary branching (also remarked by Bastiani and Mulloney, ’88a)
obscured any order that might be present. Third,
we found analysing bilateral pairs improved our
ability to compare the neurons but this means that
any interpretation of the relative longitudinal position of two filled neurons from different segments will depend to an extent on the relationship
between bilateral homologues as well. We are reasonably confident that this issue does not put the
conclusion about the order at risk because all bilateral homologues that we filled crossed the midline at the same point and where we did fill
ipsilateral pairs of cells from different segments,
the outcome was the same as that predicted by
the rest of our results. Fourth, to see the relationship of the neurons in A6 clearly we were limited to large animals and because of this were only
able to fill reliably from A3 back so we cannot be
sure that the apparent order we found applies to
the rest of the abdominal segments. Even with
these caveats, the evidence appears to point to
somatotopic organisation, but even if only three
of the ganglia are involved, the result remains interesting, particularly in light of the physiological evidence of gradients in the strength of SR
synapses in A6 (Bastiani and Mulloney, ’88b; Vescovi et al., ’97).
ACKNOWLEDGMENTS
Supported by a grant from the Australian Research Council to D.L.M. P.J.V. was the holder of
an Australian Post-Graduate Research Award.
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