Functional Morphology of Perihelia conradi Relative to Shell-Penetration

The pholad, Penitella conradi, is found along the Californiacoast in the calcareous shell of the abalone, Haliotis rufescens.These pholads penetrate the abalone shell, and when they breakthrough the inside of the shell they cease boring, secrete acallum, and then become sexually mature. The normal adult isa stenomorphic form,defined by Bartsch as an animal whose sexualmaturity is induced by over-crowding or insufficient substratumin which to bore. In the case of P. conradi, sexual maturityis always induced by the spatial limits of the substratum, thatis, the relatively thin abalone shell. The role of mechanical abrasion by the valves of P. conradiis minor. Experiments indicate that the teeth of P. conradiare worn at a greater rate than the polished shell of the abalone. The boring process in P. conradi proceeds mainly by chemicaldissolution of the calcareous substrate. The pathway of thesolvents is unknown. It may be through the organic matrix, orthe solvent may react directly with the crystals. Mechanicalabrasion helps to remove loosened crystals and/or organic matrixwhich are then carried to the exterior by the ciliary currentsflowing in through the pedal gape and out through the exhalentsiphon.     

THE ADDUCTOR AND RETRACTOR MUSCLES OF THE VELIGER OF PECTEN MAXIMUS (L.) (BIVALVIA)

In Pecten maximus (L.), retractor and adductor muscles becomefunctional in the early veliger larva. The twelve-day-old veligerhas four pairs of velar retractors, three pairs of retractorsattached to the posterior body wall and an anterior adductor.The pediveliger has in addition, pedal retractor muscles anda posterior adductor. The retractors consist of striated muscle:the adductors have both smooth and striated portions. The retractorsattach near the hinge, branch to a greater or lesser extent,then attach to specific areas of the velum, posterior body walland foot. Some features of the branching and of the dispositionof points of attachment form a pattern which exhibits mirrorsymmetry about the plane between the two shell valves. Thispattern is characteristic of the species. It is deduced thatretraction and protraction of the velum result from co-ordinatedsequences of muscle contractions. ^*Present address: Forest Products Research Centre, P.O. Box1358, Boroko, Papua New Guinea. (Received 15 June 1984;     

Evolution of the molluscan body plan: the case of the anterior adductor muscle of bivalves

The separated shell plates with the rearranged musculature (adductor muscle) is a novelty for bivalves. Despite its importance in the bivalve bodyplan, the development of the anterior adductor muscle remains unresolved. In this study, we investigate the myogenesis of the bivalve species Septifer virgatus to reveal the developmental origin of the larval muscles in bivalves, focusing on the anterior adductor muscle. We observed that larval retractor muscles are differentiated from the ectomesoderm in bivalves, and that the anterior adductor muscles are derived from primordial larval retractor muscles via segregation of the myoblast during the veliger larval stage. Through the comparative study of myogenesis in bivalves and its related taxa, gastropods, we found that both species possess myoblasts that emerge bilaterally and later meet dorsally. We hypothesize that these myoblasts, which are a major component of the main larval retractor in limpets, are homologous to the anterior adductor muscle in bivalves. These observations imply that the anterior adductor muscle of bivalves evolved as a novel muscle by modifying the attachment sites of an existing muscle.     

Ecological Aspects of Some Coral-Boring Gastropods and Bivalves of the Northwestern Red Sea

More than 17 molluscan species were obtained from burrows incoral substrata at Al-Ghardaga (Hurghada on maps) on the RedSea coast, six of which in particular bore into livingcolonies.The species reported in this paper belong to the families Mytilidae,Coralliophilidae, and Gastrochaenidae. The direction of boringin living corals is to the outside, the borers keeping pacewith the growing coral layer to maintain their burrows open.Coral growth is generally of a higher rate than that of borers,and burrows are accordingly mostly much larger than their inhabitants.There is evidence in such cases that burrows form initiallyby growth of coral around the settling young. Boring of Lithophagaspecies is mostly due to the abrasive action of the shell whichmoves straight and posteroventrally without any rotation. Incoralliophilids,boring is also executed mechanically by the turning movementsof the shell. Boring in dead coral is directed inwards, andburrows are nearly as large as the borers. The latter avoidthe blocking of their burrows (e.g., by a living coral incrustation)either by great siphonal extension (Rocellaria) or the freeends of the shell may be strengthened to maintain the capabilityof boring in the opposite direction (Lithophaga laevigata).Both L. luevigata and Modiola chmamomeus bore mainly mechanicallyby the rocking movements of the shell. Chemical boring is stilla possibility,particularly in the posterior narrow region ofburrows of Modiola lodging the extended pallial siphons whichare deprived of any effective mechanical devices for boring.Therole of boring algae in rarefying bored coral material hasalso to be included as an indirect chemical factor.     

EVIDENCE FOR CHEMICAL BORING IN PETRICOLA LAPICIDA (GMELIN, 1791) (BIVALVIA: PETRICOLIDAE)

Specimens of the dead coral-boring bivalve Petri-cola lapicidahave been obtained from Thailand and Jamaica. Although formerlyconcluded to be a mechanical borer, examination of the burrowand the shell strongly suggests chemical boring. Two glandslocated in the inner mantle folds around the antero-ventr'alpedal gape are thought to be involved in this, although onemay secrete the calcareous material cemented to the posteriorshell margin Less specialised petricolids are mechanical borers of stiffmuds, shales and calcareous rocks. A few are nestlers, e.g.,Claudiconcha. As has been recently suggested for other familiesof borers, the Petricolidae constitute another example of theevolution of a specialised chemical borer from a less specialisedmechanically-boring ancestor (Received 20 July 1987;     

DISTRIBUTION, HABITAT AND MORPHOLOGY OF THE CARIBBEAN CORAL- AND ROCK-BORING BIVALVE,LITHOPHAGA BISULCATA (d'ORBIGNY) (MYTILIDAE: LITHOPHAGINAE)

Lithophaga bisulcata is the most common Caribbean and Atlanticlithophagine and is the only species of the genus known to occurfrequently in both living and dead coral. The abundance in livingcorals is non-random and variable. Most common hosts are Siderastereasiderea and Stephanocoenia michelini. The bivalves are moreabundant in their preferred hosts than in dead coral. Individualsfrom the two habitats are indistinguishable in shell shape,musculature and size of boring and posterior pallia! glands,indicating a single population. Boreholes differ in the twohabitats with respect to size and lining. Linings are formedat the "inactive" end of the burrow; therefore living coralinhabitants line the anterior end of the burrow and dead coralborers line the posterior end. Recruitment rates are unknownin dead coral but were very low in living coral (Received 9 June 1987;     

Possible Boring Structures of Sipunculids

At the present time there is no experimental evidence whichlinks the supposed boring activities of sipunculids to a specificorgan or structure. Structures which have been speculativelyassociated in the literature with boring are: hooks and spinesof the introvert, cuticular papillae with associated epidermalglands, anterior and posterior horny shields, and anterior calcareousshields. In this review these structures are described as theyoccur in five representative species of sipunculids collectedby the author from calcareous rock in the Indian Ocean or theCaribbean Sea. The five species are: Pliascolosoma antillarumGrube and Oersted, Phascolosoma dentigerum (Selenka and de Man), Paraspidosiphon steenstrupi (Diesing), Lithacrosiphon gurjanovaeMurina, and Cloeosiphon aspergillum (Quatrefages). Localitiesof collections are cited, habitats and burrows are described,and the behavior of the animals as observed in the field andlaboratory is noted. In view of the morphology of the possibleboring structures and in light of observations on habitats andbehavior, the possible roles of the structures in boring activitiesare discussed. Highly organized horny shields are present at the anterior andposterior extremities of thetrunk or Paraspidosiphon steenstrupi,whereas anterior calcareous shields are characteristic of Cloeosiphonaspergillum and Lithacrosiphon gurjanovae. Papillae and epidermalglands are present in all five of the species but these aremost highly developed in Phascolosoma dentigerum and P. antillarum.Of the species considered, only P. antillarum lacks hooks onthe introvert. Because of the position of the animal within the rock with anteriorend directed toward themouth of the burrow, it is assumed thatthe anterior shields and the hooks of the introvert play nosignificant role in the formation of the burrow. However, therigid papillae of the trunk and the thickened posterior shield,if rubbed against the wall of the burrow, presumably could beutilized in the mechanical attrition of the more friable rock,whereas the secretory products of the numerous epidermal glandsmight be implicated in the chemical dissolution of the hardersubstrates.     

Comparative anatomy of the cheek muscles within the Centromochlinae subfamily (Ostariophysi, Siluriformes, Auchenipteridae)

Glanidium melanopterum Miranda Ribeiro, a typical representative of the subfamily Centromochlinae (Siluriformes: Auchenipteridae), is herein described myologically and compared to other representative species within the group, Glanidium ribeiroi, G. leopardum, Tatia neivai, T. intermedia, T. creutzbergi, Centromochlus heckelii, and C. existimatus. The structure of seven pairs of striated cephalic muscles was compared anatomically: adductor mandibulae, levator arcus palatini, dilatator operculi, adductor arcus palatini, extensor tentaculi, retractor tentaculi, and levator operculi. We observed broad adductor mandibulae muscles in both Glanidium and Tatia, catfishes with depressed heads and smaller eyes. Similarities between muscles were observed: the presence of a large aponeurotic insertion for the levator arcus palatini muscle; an adductor arcus palatini muscle whose origin spread over the orbitosphenoid, pterosphenoid, and parasphenoid; and the extensor tentaculi muscle broadly attached to the autopalatine. There is no retractor tentaculi muscle in either the Glanidium or Tatia species. On the other hand, in Centromochlus, with forms having large eyes and the tallest head, the adductor mandibulae muscles are slim; there is a thin aponeurotic or muscular insertion for the levator arcus palatini muscle; the adductor arcus palatini muscle originates from a single osseous process, forming a keel on the parasphenoid; the extensor tentaculi muscle is loosely attached to the autopalatine, permitting exclusive rotating and sliding movements between this bone and the maxillary. The retractor tentaculi muscle is connected to the maxilla through a single tendon, so that both extensor and retractor tentaculi muscles contribute to a wide array of movements of the maxillary barbels. A discussion on the differences in autopalatine-maxillary movements among the analyzed groups is given.     

PREDATION ON THE BIVALVE CHOROMYTILUS MERIDIONALIS (KR.) BY THE GASTROPOD NATICA (TECTONATICA) TECTA ANTON

The effects of a population of the boring gastropod Natica tectaon the bivalve Choromytilus meridionalis were investigated atBailey's Cottage, False Bay, South Africa. In July 1979 theN. tecta density on the mussel bed averaged 69 m^–2 andthe population consisted mainly of reproductively mature individualsbetween 20–33 mm shell width. Laboratory experiments on N. tecta showed that prey size selectionis an increasing function of predator size. The prey size rangetaken by large N. tecta is also greater than that taken by smallindividuals. The position of the borehole on the mussel shellis a function of the way in which the shell is held by the footduring the boring process. Consumption rates measured in thelaboratory showed an increase from approximately 1 kJ per weekin 18 mm N. tecta to 4.5 kJ per week in 28 mm individuals. Populationconsumption in the field was calculated as 663 kJ m^–2month^–1. It was estimated that at this rate the standingcrop of mussels in the pool would be eliminated within 10 months.Field measurements showed significant depletion after 6 months. New spat settlement of mussels occur every 4–6 years.The growth curve shows that after one year the population meansize exceeds 30 mm shell length, which is beyond the prey selectionsize range of small N. tecta. It was concluded that at the timeof a new mussel settlement a niche is provided for the simultaneoussettlement and growth of juvenile N. tecta in high densities.However, within one year the increase in prey size, togetherwith depletion due to over-exploitation, limits population growthand density in N. tecta. (Received 14 March 1980;     

How the 'forceps' of Lithophaga aristata (Bivalvia: Mytiloidea) are formed

The calcareous substratum borer, Lithopaga aristata Dillwyn, 1817, secretes posterior incrustations that take the form of interlocking 'forceps'. These are secreted, initially, as asymmetrical ridges by similarly asymmetrical dorsal and ventral glands in the left and right middle folds of the posterior mantle lobes. In the adult, the secretions are more uniformly spread over the posterior shell margins, concealing the juvenile ridges.
Opening and closing of the valves smooths the outer surfaces of the 'forceps' against the burrow wall, which also comes to be lined with calcium carbonate that is reciprocally smoothed, so that they occlude the borehole more effectively. Extension and retraction of the siphons probably smooths the inner surfaces of the 'forceps' which are sharpened by abrasion, one against the other, during valve opening and closing. The pair of inequilateral spikes so produced, project from the burrow aperture, occluding it, but probably, more importantly, distancing aperturally attacking predators from the true posterior shell margin. Interlocking of the 'forceps' and their sharpness may further deter would-be predators.     

Alternative predation tactics of a tropical naticid gastropod

Aquarium observations of naticid gastropods from Hong Kong show that different species attack their bivalve prey in different ways. Natica gualteriana and Glossaulax didyma appeared always to use conventional modes of boring, i.e., through one shell valve, before consuming the prey, but some larger prey of C. didyma with incomplete borings were consumed after having apparently suffocated before boring was complete. In contrast, Polinices tumidus prey may be side-bored, edge-bored (i.e., through the commisure of the valves) or suffocated and consumed without boring. The frequency of each of these modes of attack vary with different prey species. Non-boring prédation, in aquarium experiments, accounted for 14.7–54.9% of attacks with different species of prey. Suffocated prey were found to be enwrapped in a thick, viscous coat of mucus, which in partially consumed prey showed a round hole overlying the ventral shell gape marking the entrance hole made by the proboscis. The observations reveal considerable flexibility in predation behaviour in this tropical naticid and have important implications in the interpretation of naticid prédation rates in recent and fossil dead shell assemblages.     

Comparative Ultrastructure and Organization of the Prismatic Region of Two Bivalves and its Possible Relation to the Chemical Mechanism of Boring

The prismatic region of two bivalve molluscs exemplifies, inits structure and organization, one of the types of differentiatedcalcareous substrates through which boring organisms must penetrate. The oriented inorganic crystals, separated from one anotherby intercrystalline "spaces", are structurally organized intowell defined prisms. The prisms of each bivalve vary in shapeand size and are delineated from one another by electron-lucent,non-calcified regions. The demineralized organic matrix is also structurally organizedinto prisms, delineated from one another by prism sheaths, andan intraprismatic matrix structurally organized into closelypacked sheet-like compartments and subcompartments in whichthe inorganic crystals are deposited. The non-mineralized intercrystalline "spaces" between the individualinorganic crystals of the same or adjacent rows in a mineralizedsection are occupied by the walls of the intraprismalic sheet-likecompartments. Similarly, the non-calcified electronlucent regionsdelineating one mineralized prism from the next are occupiedby the thick prism sheaths. These portions of the organic matrixwhich fail to mineralize completely undoubtedly provide readypathways for the passage of solutes and solvents through thesetissues of highly ordered, densely packed, inorganic crystals.Moreover, the framework of the organic matrix, which fails tomineralize in these heavily calcified, molluscan substrates,may provide the primary, not the secondary source of chemicalattack during boring, for once the sheaths and compartmentssurrounding the crystals are broken down or solubilized, thecrystals are themselves loosened and freed for mechanical removalby shell-penetrating organisms.     

Boring Mechanism of Polydora websteri Inhabiting Crassostrea virginica

The boring mechanisms of species of polydorid polychaetes arelittle understood due to lack of experimental evidence and directobservations. In the present studies the boring mechanism ofadults and metamorphosing larvae of Polydora websteri was investigatedby (1) inducing adults and larvae to settle against test substrates,(2) observing behavior in natural burrows and in "artificialblisters" composed of transparent "Pliobond" films surroundingIceland spar substrates, (3) removing the giant setae of wormspriorto tests of boring, (4) applying the giant setae to substrates,and by (5) testing for production of acid. All the layers of oyster shell, including conchiolin, were bored.Calcareous substrates andIceland spar were penetrated rapidlyby adults without the assistance of the giant setae. Nor werethese organs essential to the boring of a larva. A characteristictype of behavior involving close contact with the substrateduring backwards and forwards movements and periods of immobilityalways preceded boring. The worms produced acid, probably somecommon product of metabolism, which can account for these results.     

Functional morphology and palaeontological significance of the conchiolin layers in corbulid pelecypods

The Corbulidae, which today are slow, cumbersome, very shallow burrowers, developed special morphological features by which they obtained an outstanding capability to withstand the physical and biological stresses characteristic of their preferred habitat. These features are: an inequivalve, globose shape, thick shells, and conchiolin layers (at least one) embedded within their valves in a unique way. These features enable the corbulids to close their valves tightly during the unfavourable environmental conditions (e.g. low salinity, low oxygen content) which may prevail in the marginal marine regions inhabited by several corbulid species. The conchiolin layers act as a barrier preventing all chemically boring organisms from penetrating into the bivalve shell, or shell dissolution by sea water undersaturated with respect to calcium carbonate. The layered conchiolin weakens the shell mechanically, however, especially during fossilization, when the conchiolin is decomposed. The valve splits apart into two shells so completely different in appearance that they may be attributed to different taxa. The conchiolin layers are therefore of great ecological and palaeontological significance. The nature of these conchiolin layers in Corbula (Varicorbula) gibba (Olivi) is described and illustrated and their functional significance discussed in relation to other living and fossil corbulid species.     

Torsion in Patella caerulea (Mollusca, Patellogastropoda): ontogenetic process, timing, and mechanisms

Abstract. Torsion is a process in gastropod ontogenesis where the visceral body portion rotates 180° relative to the head/foot region. We investigated this process in the limpet Patella caerulea by using light microscopy of living larvae, as well as scanning electron microscopy (SEM) of larvae fixed during the torsion process. The completion of the 180° twist takes considerably less time in larvae of Patella caerulea than previously described for other basal gastropod species. At a rearing temperature of 20–22°C, individuals complete ontogenetic torsion in ?2 h. Furthermore, the whole process is monophasic, i.e., carried out at a constant speed, without any evidence of distinct ‘fast” or ‘slow” phases. Both larval shell muscles—the main and the accessory larval retractor—are already fully contractile before the onset of torsion. During the torsion process both retractors perform cramp‐like contractions at ～30 s intervals, which are followed by hydraulic movements of the foot. However, retraction into the embryonic shell occurs only after torsion is completed. The formation of the larval operculum is entirely in‐dependent from ontogenetic torsion and starts before the onset of rotation, as does the mineralization of the embryonic shell. The reported variability regarding the timing (mono‐ versus biphasic; duration) of torsion in basal gastropod species precludes any attempt to interpret these data phylogenetically. The present findings indicate that the torsion process in Patella caerulea, and probably generally in basal gastropods, is primarily caused by contraction of the larval shell muscles in combination with hydraulic activities. In contrast, the adult shell musculature, which is independently formed after torsion is completed, does not contribute to ontogenetic torsion in any way. Thus, fossil data relying on muscle scars of adult shell muscles alone appear inappropriate to prove torted or untorted conditions in early Paleozoic univalved molluses. Therefore, we argue that paleontological studies dealing with gastropod phylogeny require data other than those based on fossilized attachment sites of adult shell muscles.     

Development of the musculature in the limpet Patella (Mollusca, Patellogastropoda)

 Whole-mount technique using fluorescent-labelled phalloidin for actin staining and confocal laser scanning microscopy as well as semi-thin serial sectioning, scanning and transmission electron microscopy were applied to investigate the ontogeny of the various muscular systems during larval development in the limpets Patella vulgata L. and P. caerulea L. In contrast to earlier studies, which described a single or two larval shell muscles, the pretorsional trochophore-like larva shows no less than four different muscle systems, namely the asymmetrical main head/foot larval retractor muscle, an accessory larval retractor with distinct insertion area, a circular prototroch/velar system, and a plexus-like pedal muscle system. In both Patella species only posttorsional larvae are able to retract into the shell and to close the aperture by means of the operculum. Shortly after torsion the two adult shell muscles originate independently in lateral positions, starting with two fine muscle fibres which insert at the operculum and laterally at the shell. During late larval development the main larval retractor and the accessory larval retractor become reduced and the velar muscle system is shed. In contrast, the paired adult shell muscles and the pedal muscle plexus increase in volume, and a new mantle musculature, the tentacular muscle system, and the buccal musculature arise. Because the adult shell muscles are entirely independent from the various larval muscular systems, several current hypotheses on the ontogeny and phylogeny of the early gastropod muscle system have to be reconsidered. Received: 23 June 1998 / Accepted: 25 November 1998     

Developmental dynamics of myogenesis in the shipworm <Emphasis Type="Italic">Lyrodus pedicellatus</Emphasis> (Mollusca: Bivalvia)

Background

The shipworm Lyrodus pedicellatus is a wood-boring bivalve with an unusual vermiform body. Although its larvae are brooded, they retain the general appearance of a typical bivalve veliger-type larva. Here, we describe myogenesis of L. pedicellatus revealed by filamentous actin labelling and discuss the data in a comparative framework in order to test for homologous structures that might be part of the bivalve (larval) muscular ground pattern.

Results

Five major muscle systems were identified: a velum retractor, foot retractor, larval retractor, a distinct mantle musculature and an adductor system. For a short period of larval life, an additional ventral larval retractor is present. Early in development, a velum muscle ring and an oral velum musculature emerge. In late stages the lateral and dorsal mantle musculature, paired finger-shaped muscles, an accessory adductor and a pedal plexus are formed. Similar to other bivalve larvae, L. pedicellatus exhibits three velum retractor muscles, but in contrast to other species, one of them disappears in early stages of L. pedicellatus. The remaining two velum retractors are considerably remodelled during late larval development and are most likely incorporated into the elaborate mantle musculature of the adult.

Conclusions

To our knowledge, this is the first account of any larval retractor system that might contribute to the adult bodyplan of a (conchiferan) mollusk. A comparative analysis shows that a pedal plexus, adductors, a larval velum ring, velum retractors and a ventral larval retractor are commonly found among bivalve larvae, and thus most likely belong to the ground pattern of the bivalve larval musculature.

     

The Fossil Record of Shell Boring by Snails

The predatory boring habit common to many recent snails probablyarose first in the Polinicinae (Naticacea) during Upper Cretaceous(Cenomanian) times (100 million years B.P.) . In the fossilrecord the frequency of bored shells increasesgreatly in rocksof latest Cretaceous age and becomes more widespread duringearly Tertiary times coincident with the major diversificationof the primary groups of boring snails. The borings in these Cretaceous and Tertiary shells show thesame characteristics of preference of penetration in one pelecypodvalve rather than the other or in position of the boring siteon the shell that are found in recent shell assemblages. Borings in Paleozoic brachiopod shells (230–550 millionyears old) that have previously been attributed to gastropodpredation are herein attributed to other but unknown boringorganisms. In part these borings are not accepted as evidence of Paleozoicgastropod predation because it necessitates: (1) Postulationof the separate development of a boring habit ith its concomitantdevelopment of an accessory boring organ in a groupwhose descendantsare all herbivores, and (2) The development of such a habithundreds of millions of years before the appearance of any relativesof present day borers.     

Ontogenetic torsion in two basal gastropods occurs without shell attachments for larval retractor muscles

Results of this study on two species of vetigastropods contradict the long-standing hypothesis, originally proposed by Garstang (1929), that the larval retractor muscles power the morphogenetic movement of ontogenetic torsion in all basal gastropods. In the trochid Calliostoma ligatum and the keyhole limpet Diodora aspera, the main and accessory larval retractor muscles failed to establish attachments onto the protoconch (larval shell) when the antibiotics streptomycin sulfate and penicillin G were added to cultures soon after fertilization. Defects in protoconch mineralization were also observed. Despite these abnormalities, developing larvae of these species accomplished complete or almost complete ontogenetic torsion, a process in which the head and foot rotate by 180 degrees relative to the protoconch and visceral mass. Analysis by using phalloidin-fluorophore conjugate and transmission electron microscopy showed that myofilaments differentiated within myocytes of the larval retractor muscles and adherens-like junctions formed between muscle and mantle epithelial cells in both normal and abnormal larvae. However, in abnormal larvae, apical microvilli of mantle cells that were connected to the base of the larval retractor muscles failed to associate with an extracellular matrix that normally anchors the microvilli to the mineralized protoconch. If morphogenesis among extant, basal gastropods preserves the original developmental alteration that created gastropod torsion, as proposed by Garstang (1929), then the alteration involved something other than the larval retractor muscles. Alternatively, the developmental process of torsion has evolved subsequent to its origin in at least some basal gastropod clades so that the original alteration is no longer preserved in these clades.     

Neural mechanisms governing distribution of cardiac output in an isopod crustacean,Bathynomus doederleini: reflexes controlling the cardioarterial valves

In Bathynomus doederleini all of the cardioarterial valves located at the origin of the lateral arteries are dilated by impulses of lateral cardiac nerves. Tactile stimuli applied to sensillar setae depress impulse activities of the 1st and 5th lateral cardiac nerves. The 1st lateral cardiac nerve controls the valve of the lateral artery which runs to the walking-legs and viscera. The 5th lateral cardiac nerve controls the valve of the lateral artery which runs to the swimmeret muscles. The response indicates that tactile receptor reflexes bring about decreased haemolymph flow to the organs. Augmented swimmeret movements were always accompanied by an increased firing rate in the 5th lateral cardiac nerve. Artificial full protraction of swimmerets simultaneously induced excitation of the 5th lateral cardiac nerve and inhibition of the 1st lateral cardiac nerve. The excitation corresponds to an increase in haemolymph flow to the swimmerets, and the inhibition a decrease in haemolymph flow to walking-legs and viscera. Three kinds of mechanoproprioceptors which were activated by swimmeret movements were found. Two of the mechanoproprioceptors are located at the base of the basipodite. The other mechanoproprioceptor supplies processes to a nerve to the retractor muscles. Activation of three kinds of mechanoproprioceptors, induced by artificial swimmeret protraction, triggered lateral cardiac nerve reflex responses.Abbreviations LA lateral artery - LCN lateral cardiac nerve - RMN nerve to retractor muscles - StR stretch receptor