SINDERMANX: INTERNAL DEFENSES OF CRUSTACEA 



Caiitacuzene (1923b) also examined the in- 

 ternal defenses of the hermit crab, Eiii)a(/nnis 

 prideauxii. which he had reiiorted earlier to 

 possess strong hemolysins and strong' antibac- 

 terial agglutinins. Injected gram-negative bacilli 

 were entrapped and immobilized on the cell sur- 

 faces of the branchial lacunae, and then phago- 

 cytized, as was the case with Maia described 

 ]ireviously. 



Cantacuzene (1923b) also reported that the 

 sera of crabs, Carcinus maenas, infected by the 

 rhizocephalan SacrnUna, contained a factor ab- 

 sent from normal crabs. Using a standard com- 

 plement fixation test, with extract of the rhizo- 

 cephalan as antigen and with ci-ab serum, 

 Cantacuzene (1925b) was able to demonstrate 

 an antibody-like response in parasitized crabs. 

 Sheep cells were lysed in tubes with normal crab 

 serum but not in those containing parasitized 

 crab serum. Using a fine suspension of Sarcu- 

 lina. Cantacuzene found precipitating and agglu- 

 tinating activity in the serum of parasitized 

 crabs. The activity was not consistent, however, 

 in that some sacculinized crabs lacked it. In a 

 concurrent study. Levy (1923) found that macer- 

 ated sacculinids wei-e toxic when injected into 

 crabs, but that no antitoxic activity could be dem- 

 onstrated in ]iarasitized crabs. Both groups, 

 normal and parasitized, died at about the same 

 rate. 



In an earlier study Cantacuzene (1913) re- 

 ported that inoculation of the sacculinid parasite 

 with gram-negative bacteria resulted in septi- 

 cemia and death of the parasite within 1 week, 

 and infection of the crab host by 10 days after 

 inoculation. At about 5 days after inoculation, 

 the crab hemolymph became incoagulable, and 

 agglutinins appeared against the bacteria inoc- 

 ulated into the parasite. The antibacterial 

 agglutinins were not present in sacculinized un- 

 inoculated control crabs. 



Appreciable evidence for some degree of spe- 

 cificity of the natui'al agglutinins of Crustacea 

 was accumulated by Cantacuzene. An agglutinin 

 in Maia against mammalian red cells could be 

 absorbed from crab serum by certain gram-jjos- 

 itive bacteria but did not agglutinate them, nor 

 did it agglutinate cholera vibrios. It did, how- 

 ever, strongly agglutinate typhoid bacilli. 



Agglutinins against vei-tebrate erythrocytes and 

 certain bacteria were found in sera of Homarus 

 vulgaris, Eupagurus prideauxii, and E. bernhar- 

 dus, but the same antigens were not agglutinated 

 by sera of Cancer pagurus, Carcinus maenas, 

 Portunus puber, or Galathea punctata. The 

 serum of Eupagurus prideauxii agglutinated 

 mammalian red blood cells and amebocytes of 

 Maia and Buccinum, but did not agglutinate 

 the coelomic cells of sipunculids or ascidians. 

 And so Cantacuzene set the scene, by the 

 early 1930's, for the continuation of broad and 

 elaborate studies of humoral internal defenses 

 of marine invertebrates — particularly the Crus- 

 tacea — but the stage, with only a few notable ex- 

 ceptions, remained curiously empty and dark 

 until the mid-19.50's. Although research during 

 the last decade emphasized species other than 

 those studied by Cantacuzene (except for the 

 work of F. Bang) , it reinforced many of Can- 

 tacuzene's findings: that natural agglutinins 

 and lysins, with some specificity, occur in Crus- 

 tacea and other invertebrates, and that respon- 

 ses to foreign antigens can be induced in selected 

 invertebrate.? — responses which are only par- 

 tially specific. The increased precision and 

 quantitation of tests, and the careful attention 

 to controls, have improved the quality of the 

 newer data, but have neither provided new con- 

 cepts nor modified the genei-al conclusions of 

 Cantacuzene. The more recent literature on 

 humoral defenses of Crustacea will be summar- 

 ized by general categories in the following 

 sections. 



Bactericidal Systems 



Natural bactericidins have been reported from 

 a number of marine invertebrates (Bang, 

 1967b). Recently, increase in titers of bacteri- 

 cidal activity after inoculation with Formalin- 

 killed bacteria was noted in West Indian spiny 

 lobsters, Panidirus argus, and American lob- 

 sters, Homarus americanus (Evans et al, 1968; 

 Acton, Weinheimer, and Evans, 1969). A bac- 

 tericidal assay system described by Schwab and 

 Reeves (1966) was used to quantitate the degree 

 of response. In experiments with American 

 lobsters held in seawater at 5° C, the peak of 



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