Observations on the Burrowing of Arenicola Marina (L.)

1966 ◽  
Vol 44 (1) ◽  
pp. 93-118
Author(s):  
E. R. TRUEMAN

1. Continuous recordings of the hydrostatic pressure in the coelom of Arenicola marina show a resting pressure of about 2 cm. of water in a non-burrowing worm. During burrowing a series of pressure peaks is produced and these gradually increase in amplitude up to 110 cm. as burrowing progresses. 2. The pressure peaks are of 2 sec. duration, occur at intervals of 5-7 sec., and for each there is a major contraction of the circular muscles followed by the shortening of the longitudinal muscles. The main power stroke in producing the high pressure is the contraction of the longitudinal muscles of most of the trunk segments. The sequence of muscular contractions and the phases of burrowing are considered. 3. The pressure is utilized at the anterior end of the worm both to aid passage through the sand and to anchor the head while the posterior segments are pulled into the burrow. 4. At maximum pressures the tension developed in the circular muscle of the body wall is estimated to be 3 kg./cm.2, while the resting pressure corresponds to less than 7% of this.

1954 ◽  
Vol s3-95 (30) ◽  
pp. 251-270
Author(s):  
G. P. WELLS

The mechanism of proboscis movement is analysed in detail in Arenicola marina L. and A. ecaudata Johnston, and discussed in relation to the properties of the hydrostatic skeleton. Proboscis activity is based on the following cycle of movements in both species. Stage I. The circular muscles of the body-wall and buccal mass contract; the head narrows and lengthens. Stage IIa. The circular muscles of the mouth and buccal mass relax; the gular membrane (or ‘first diaphragm’ of previous authors) contracts; the mouth opens and the buccal mass emerges. Stage IIb. The longitudinal muscles of the buccal mass and body-wall contract; the head shortens and widens and the pharynx emerges. Stage III. As Stage I. The two species differ anatomically and in their hydrostatic relationships. In ecaudata, the forward movement of body-fluid which extrudes and distends the proboscis is largely due to the contraction of the gular membrane and septal pouches. In marina, the essential mechanism is the relaxation of the oral region which allows the general coelomic pressure to extrude the proboscis. The gular membrane of marina contracts as that of ecaudata does, but its anatomy is different and it appears to be a degenerating structure as far as proboscis extrusion is concerned. Withdrawal of the proboscis may occur while the head is still shortening and widening in Stage IIb, or while it is lengthening and narrowing in Stage III. The proboscis is used both in feeding and in burrowing; in the latter case nothing enters through the mouth; the difference is largely caused by variation in the timing of withdrawal relative to the 3-stage cycle.


Author(s):  
G. P. Wells

Worms for dissection, or for museum preservation, should be prepared by the magnesium-formalin method, of which two modifications are given in the text.The body of Arenicola is differentiated into: (a) an achaetous ‘head’, comprising the prostomium and a small number (probably two) of subsequent segments, (b) a ‘trunk’, composed of a number, varying somewhat with the species, of chaetigerous segments, and (c) a ‘tail’, which may or may not be chaetigerous according to the species. The method of subdividing the body according to the distribution of the gills, so often met with in the literature, is misleading because it conceals the very fundamental differentiation between ‘head’ and ‘trunk’.The main layers of the body wall are described. There are grounds for supposing that the circular muscle layer plays a greater part than the longitudinal in the maintenance of a postural fluid pressure in active worms.


1969 ◽  
Vol 51 (1) ◽  
pp. 47-58
Author(s):  
M. K. SEYMOUR

1. Crawling movement and burrowing of Lumbricus terrestris (L.) have been studied by continuous recording of internal pressure, direct observation and analysis of cine film. Frequency of locomotory waves is from 5 to 20 per min. Timing of protrusion of setae and of backward slip of points d'appui in locomotion have been observed and recorded. 2. In normal locomotion elongation of segments by contraction of the circular muscles gives rise to a discrete pressure pulse; shortening, by contraction of the longitudinal muscles, may or may not do so, depending on the position of the segment in the worm and the relative extent of contraction of the longitudinal and circular muscles. 3. Consideration of crawling and burrowing pressure records emphasizes the importance of (a) the circular muscles in extension of the head end in crawling and in initial penetration of the soil, and (b) the longitudinal muscles during burrowing, in dilating the burrow and drawing in more posterior segments 4. Mean pressures at circular and longitudinal muscle contraction are 12 and 7 cm. H2O respectively. The highest pressure recorded was 75 cm. H2O and accompanied violent squirming with evident contraction of all the body wall muscles. Resting pressures, shown in the absence of organized movement, are low (mean 0.26 cm. H2O). In both resting and crawling negative pressures sometimes occur and these are considered in relation to the inherent stiffness of the body wall and to the septate condition. 5. Tension in the longitudinal and circular muscle layers of a worm developing 75 cm. H2O internal pressure are calculated to be 265 and 1323 g./cm2. respectively, demonstrating in this example that, relative to the circulars, the longitudinal muscles are understressed by a factor of 5. Mean locomotory L.M. and C.M. peak values yield tension values of only 25 and 212 g./cm. respectively, and these are clearly well within the worm's capacity.


Author(s):  
Hugh Y. Elder

The polychaete Polyphysia crassa (Oersted) is unusual in possessing a well-developed body-wall connective-tissue layer which exceeds the combined thickness of the circular and longitudinal muscles. Both collagenous and elastic fibres are present in this layer. The collagen is organized as a loose three-dimensional lattice allowing longitudinal, circumferential or radial distension of the body wall and, as in other soft-bodied invertebrates, serves the functions of providing a base on which the muscles can act and of imposing limits to the extensibility of the system. The elastic fibres are organized as an apparently randomly oriented meshwork of stout fibres around the circular muscle blocks and are attached to both the circular and longitudinal muscles. Columns of elastic fibres extend radially from the supramuscular coarse meshwork through the ‘holes’ in the collagen lattice to the epidermal basement membrane. As the elastic columns extend outwards the fibres become finer and more numerous and flute out to give a wide area attachment to the epidermal basement membrane. The radial elastic columns are cross-linked by tangential elastic fibres. Although at any point the main collagen and elasticfibre bundles are oriented at right angles to one another, collagen fibres are invariably associated with the elastic fibres and the two fibre types form a single functionally inte-grated system. Polyphysia lives in flocculent, sublittoral muds and the burrowing mechanism employed involves a pronounced direct peristaltic wave of simultaneous circular and longitudinal muscle contraction which necessitates considerable radial thickening of the body wall. The functions of the elastic fibres appear to be to oppose the radial distension of the body wall which the collagen lattice permits and to control the folding of the cuticle and epidermis and the return of the collagen system after the passage of a peristaltic wave.


1960 ◽  
Vol s3-101 (54) ◽  
pp. 149-176
Author(s):  
R. B. CLARK ◽  
M. E. CLARK

Nephtys lacks circular body-wall muscles. The chief antagonists of the longitudinal muscles are the dorso-ventral muscles of the intersegmental body-wall. The worm is restrained from widening when either set of muscles contracts by the combined influence of the ligaments, some of the extrinsic parapodial muscles, and possibly, to a limited extent, by the septal muscles. Although the septa are incomplete, they can and do form a barrier to the transmission of coelomic fluid from one segment to the next under certain conditions, particularly during eversion of the proboscis. Swimming is by undulatory movements of the body but the distal part of the parapodia execute a power-stroke produced chiefly by the contraction of the acicular muscles. It is suspected that the extrinsic parapodial muscles, all of which are inserted in the proximal half of the parapodium, serve to anchor the parapodial wall at the insertion of the acicular muscles and help to provide a rigid point of insertion for them. Burrowing is a cyclical process involving the violent eversion of the proboscis which makes a cavity in the sand. The worm is prevented from slipping backwards by the grip the widest segments have on the sides of the burrow. The proboscis is retracted and the worm crawls forward into the cavity it has made. The cycle is then repeated. Nephtys possesses a unique system of elastic ligaments of unusual structure. The anatomy of the system is described. The function of the ligaments appears to be to restrain the body-wall and parapodia from unnecessary and disadvantageous dilatations during changes of body-shape, and to serve as shock-absorbers against the high, transient, fluid pressures in the coelom, which are thought to accompany the impact of the proboscis against the sand when the worm is burrowing. From what is known of its habits, Nephtys is likely to undertake more burrowing than most other polychaetes.


1925 ◽  
Vol s2-69 (274) ◽  
pp. 245-290
Author(s):  
A. J. GROVE

During the sexual congress of L.terrestris, the co-operating worms become attached to one another in a head-to-tail position in such a way that segments 9-11 of one are opposed to the clitellum of the other, and vice versa. At these points the attachment between the worms is an intimate one, assisted by the secretion of the glands associated with the diverticula of the setal pores found in certain segments, and is reinforced by the mutual penetration of the setae into the opposed body-surfaces. There is also a slighter attachment between segment 26 of one and 15 of the other. Each worm is enclosed in a slime-tube composed of mucus secreted from the epidermis. The exchange of seminal fluid is a mutual one. The fluid issues from the apertures of the vasa deferentia in segment 15, and is conducted beneath the slime-tube in pit-like depressions in the seminal grooves, which extend from segment 15 to the clitellum on each side of the body, to the clitellum, where it accumulates in the space between the lateral surfaces of segments 9-11 of one worm and the clitellum of the other. Eventually it becomes aggregated into masses in the groove between segments 9 and 10, and 10 and 11, and passes thence into the spermathecae. The seminal groove and its pit-like depressions are brought into existence by special muscles lying in the lateral blocks of longitudinal muscles of the body-wall.


Parasitology ◽  
1971 ◽  
Vol 62 (1) ◽  
pp. 35-47 ◽  
Author(s):  
P. J. Whitfield

The mature egg and the acanthor of Moniliformis dubius have been redescribed with special emphasis on the features relevant to the locomotion of this larval acanthocephalan. The movements of acanthors have been analysed by the use of frame by frame study of filmed records of motile acanthors. Acanthors appear to use the same mode of locomotion for hatching, locomotion within the gut of the intermediate host and penetration of the host's gut wall. Movement is produced by a set of spiralled, longitudinal muscles in the body wall of the hind body and two rostellar retractor muscles. This musculature acts both directly on the body wall and indirectly by hydraulic effects via the hydrostatic skeleton of pseudocoelomic fluid. The spiny evertable rostellum and the backward facing spines of the hind body are the means whereby shape changes of the acanthor interact with the immediate environment to produce effective progression.I should like to thank Professor D. Arthur for the provision of laboratory facilities, Dr D. W. T. Crompton for the initial gift of eggs of M. dubius and Mr R. D. Reed for invaluable assistance with microcinematographic technique. The work was carried out during the tenure of a Nuffield Foundation Research Fellowship.


Parasitology ◽  
2003 ◽  
Vol 126 (3) ◽  
pp. 273-282 ◽  
Author(s):  
M. T. STEWART ◽  
A. MOUSLEY ◽  
B. KOUBKOVÁ ◽  
š. šEBELOVÁ ◽  
N. J. MARKS ◽  
...  

The major muscle systems of the metacercaria of the strigeid trematode, Apatemon cobitidis proterorhini have been examined using phalloidin as a site-specific probe for filamentous actin. Regional differences were evident in the organization of the body wall musculature of the forebody and hindbody, the former comprising outer circular, intermediate longitudinal and inner diagonal fibres, the latter having the inner diagonal fibres replaced with an extra layer of more widely spaced circular muscle. Three orientations of muscle fibres (equatorial, meridional, radial) were discernible in the oral sucker, acetabulum and paired lappets. Large longitudinal extensor and flexor muscles project into the hindbody where they connect to the body wall or end blindly. Innervation to the muscle systems of Apatemon was examined by immunocytochemistry, using antibodies to known myoactive substances: the flatworm FMRFamide-related neuropeptide (FaRP), GYIRFamide, and the biogenic amine, 5-hydroxytryptamine (5-HT). Strong immunostaining for both peptidergic and serotoninergic components was found in the central nervous system and confocal microscopic mapping of the distribution of these neuroactive substances revealed they occupied separate neuronal pathways. In the peripheral nervous system, GYIRFamide-immunoreactivity was extensive and, in particular, associated with the innervation of all attachment structures; serotoninergic fibres, on the other hand, were localized to the oral sucker and pharynx and to regions along the anterior margins of the forebody.


Parasitology ◽  
1996 ◽  
Vol 112 (5) ◽  
pp. 459-467 ◽  
Author(s):  
J. Schmidt ◽  
O. Bodor ◽  
L. Gohr ◽  
W. Kunz

SUMMARYParamyosin, although a widely distributed muscle component among invertebrates, has hitherto not clearly been shown to occur in the muscles of schistosomes. Instead, it has been reported to occur in the tegument. In the present study, a specific antibody reacting with each of 10 isoforms of paramyosin was used for light microscopical immunolocalization in sections of Schistosoma mansoni. Specimens were fixed by a new method to immobilize antigens with uranyl acetate–trehalose–methanol. In cercariae, schistosomula, and adults, the circular and longitudinal muscles of the body wall, the dorsoventral muscles and those surrounding the gut and the pharynx as well as the fast moving cross-striated muscles of the tail of cercariae intensely reacted with the antibody. However, neither immunohistologically nor on Western blots of isolated tegument, were indications found for the presence of paramyosin in the tegument. In vivo phosphorylation and binding of anti-phospho-tyrosine and anti-phospho-serine antibodies show phosphorylation of paramyosin which probably is responsible for the generation of the isoforms.


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