
Autotomy.
Autotomy occurs in some holothurians as a method of transverse fission into two parts, both of which survive. This has been shown by Dalyell (1851) in Cucumaria lactea (suggested to be really C. planci by Deichmann (1922); in C. planci (Chadwick, 1891; Monticelli, 1896): in Holothuria parvula (syn. H. captiva, Actinopyga parvula by Crozier, 1914, 1917; Deichmann, 1922; Kille, 1936; 1937): in Actinopyga difficilis Deichmann (1922): and in Holothuria surinamensis Crozier (1917).
Autotomy in the sense of explusion of viscera could be deduced as occurring in natural conditions from Minchin's (1892) observation that the viscera of Holothuria were frequently fished up by fishermen. He states that evisceration thus seems to be almost a normal habit. It is stated by Bertolini (1932a) that autotomy is a normal and periodic occurrence in Stichopus regalus. She found that of 119 specimens examined during September and October, 110 were regenerating and 7 had recently eviscerated, while of 50 animals collected during spring none was in a state of regeneration. Kille (1936) found specimens of Holothuria floridana already in a state of regeneration when they were collected from their natural environment.
A number of other holothurians have been found capable of autotomy as a result of certain chemical and physical stimuli, but with no definite evidence that the process occurs in these species in natural conditions. In Thyone briareus autotomy was induced by the injection of chemicals of which strychnine was most satisfactory (7 out of 20), Pearse (1909); by allowing the animal to stand in stagnant sea-water and alternating this with running water containing much oxygen which proved effective in 65 per cent. of the animals treated, Scott (1914); by the use of dilute ammonia, Kille (1931); and by electrical stimuli, Kille (1935). Both the latter methods were almost invariably successful. The injection of various chemicals induced autotomy in Holothuria atra, H. sanguinolenta, and Stichopus chloronotus, and it was induced in Synapta maculata by placing specimens in a small volume of sea-water, Domantay (1931). There appears to be no evidence that autotomy occurs in any of these species in natural conditions. On the contarary Kille (1935) found that among some 400 medium-sized specimens of Thyone briareus which were examined, there was none in a natural state of regeneration. This indicates that autovisceration had not occurred in nature in any of these specimens. Of 420 specimens of Stichopus mollis examined during all seasons of the year, none showed any trace of regeneration, and the only specimens which were eviscerate on collection were 6 cast up after storms,

and some specimens collected by trawling. All the eviscerate specimens showed clearly by the state of the mesentery edges that autotomy had just occurred, and can be presumed to be due to the severe mechanical effect of wave action in one case and of trawling in the other. Autovisceration could, however, be artificially induced in all specimens tested, which included a size range from 8 gm. to 300 gm. Although there is ample evidence to show the widespread ability of holothurians to autotomise certain organs under special conditions. there are fewer recorded cases of the occurrence of this process under natural conditions. Each stimulus varies greatly in effectiveness from species to species, e.g., the methods used by Kille (1931 and 1935) for inducing autotomy in Thyone were found quite ineffective for Holothuria parvula Kille (1937). The use of dilute ammonia was found unsatisfactory for S. mollis, whereas the injection of distilled water was effective in all specimens of this species. In view of the special conditions sometimes needed to induce autotomy in holothurians, it is surprising to find the property apparently present in all specimens when suitable stimuli are used.
The organs expelled by different holothurians vary among genera but are very constant within members of the genus. In Thyone the entire length of the alimentary canal, and its haemal vessels, the introvert and the lantern with its associated structures are autotomised, Kille (1935). The respiratory trees and the gonads are retained. Phyllophorus magnus under some conditions autotomises the whole anterior part together with the tentacles, mouth, calcareous ring and other portions “snap off from the body,” Domantay (1931). Synapta appears to have less clearly defined points of autotomy and responds to inducing stimuli by constriction of the body into fragments. Species of Holothuria lose the part of the alimentary canal between the oesophagus and the cloaca, the rete mirabile and the left respiratory tree (Semper, 1861; Kille, 1936). This differs from Stichopus mainly in the retention of the right respiratory tree. In Stichopus mollis branches of gonads are also frequently expelled. Although there is a considerable variation in the organs expelled by different genera of holothurians, there is very great constancy in the points of rupture and in the organs expelled in each genus. There appear to be constant breaking points present within each genus. Lukas (1905), in a work not available to the writer, is stated by Pearse (1909) to interpret the different results as indicating that the place where self-mutilation takes place in an animal is usually determined by certain structural characteristics of the animal concerned. There is, however, insufficient data on the morphology of the regions at which rupture occurs, to show the reason for separation at these points. Histological examination of the regions of rupture in S. mollis did not show any distinct discontinuity or constriction of any of the layers in the alimentary canal. It is understandable that the point of junction of respiratory trees and rectum with the cloaca might be structurally weaker than other regions of the alimentary canal, but the reason for the constancy in position of the rupture in the oesophagus is not clear. All specimens of Stichopus mollis examined showed identical points of rupture.

It seems rather surprising, in view of the widespread occurrence of autotomy in Holothurians and in the constancy of breaking points in each genus, that possible utility of the process in suggested only in certain species. Those which undergo binary fission probably use the process as a method of asexual reproduction. Species which show regeneration in natural conditions must be capable of responding to some inducing stimulus in the environment, and it is possible that this stimulus is predator attack. The process has been suggested as protective by Minchin (1892) for Holothuria and by Domantay (1931) for the genera Holothuria, Stichopus, Thelenota and Actinopyga, which are stated to react to irritation or strong stimulation by ejecting their viscera. A series of experiments was performed on Stichopus mollis to determine its rate of response to the type of mechanical stimuli likely to be involved in predator attack. It was found that violent agitation accompanied by extensive pricking with tacks projecting from plates which held the specimens was effective only after ten or more minutes. Crushing, although effective after prolonged action, was also very slow in inducing autotomy. The treatments used were much more severe than would be expected from a predator, yet the response was far too slow for autotomy to be of any value in diverting the attention of a possible predator to the expelled viscera. There is also no evidence of any fish or other organism which preys on S. mollis. It is difficult to visualise a protective function for the process in this species.
There is much evidence, as previously mentioned, to show that autotomy can be induced casily in many holothurians by certain chemical and physical stimuli. It is difficult to relate the response to these stimuli with any conditions which would be encountered by holothurians in their natural environment. A number of holothurians, including S. mollis, undergo autotomy when the sea-water becomes, sufficiently soul. Experiments on S. mollis were carried out to determine the relative effects of oxygen deficiency and that of the accumulation of excretory products in inducing autotomy. It was sound that specimens in previously boiled and cooled seawater would not undergo autotomy any sooner than those kept in a similar volume of fresh sea-water. Of eight specimens kept separately in sea-water in which the stirred-up faeces of Stichopus mollis was added, six eviscerated within three hours, while the eight control specimens took 15–20 hours. Two specimens died without undergoing autotomy. This shows that in S. mollis the accumulation of excretory products in the main factor in inducing autotomy in specimens contained in a small volume of sea-water. It has been suggested (Pearse, 1909; Domantay, 1931) that the ejection of the visceral organs usually decreases the total amount of metabolism in holothurians, and thus they are able to survive until better environmental conditions again arise. S. mollis was found to be even more sensitive to foulness of the water after autotomy than before the process. Specimens which had eviscerated and were not immediately transferred to fresh water showed a high mortality rate. Specimens which remained intact under the same conditions showed no after-effects. Those which eviscerated and were quickly transferred to fresh sea-water showed a higher survival rate than

those left in the fouled sea-water for a longer time. Even if autotomy could be a response to certain adverse environmental conditions, the survival of S. mollis is dependent on rapid transference to normal conditions. The habitat of S. mollis in sheltered regions below tide levels in any case presents no opportunities for the accumulation of excretory products. It is therefore difficult to see how autotomy could be of benefit to S. mollis. as an adaptation against adverse environmental conditions of a chemical nature.
It has been suggested that autotomy in Synapta is pathological, and that other species in which autotomy is due to foulness of the water also employ “pathologic autotomy” (Domantay, 1931). While the lack of evidence of autotomy in natural conditions in many species including Stichopus mollis supports this view, the fact that it can be artificially induced, with quite constant points of rupture in all specimens of S. mollis, shows that the potentiality for the process is universally present in this species. For that reason the term “pathologic” can hardly be used in its strict sense. The process appears to be a normal response to certain stimuli, which rarely occur in natural conditions, and there is at present no satisfactory hypothesis to suggest a possible utility of the process to this species.
Except for holothurians in which autotomy is a process of self-division and multiplication, there is at present insufficient evidence to demonstrate clearly the function of this process in any of the holothurians known to have this property.
