Algal endosymbiosis in brown hydra: host/symbiont specificity

Host/symbiont specificity has been investigated in non-symbiotic and aposymbiotic brown and green hydra infected with various free-living and symbiotic species and strains of Chlorella and Chlorococcum. Morphology and ultrastructure of the symbioses obtained have been compared. Aposymbiotic Swiss Hydra viridis and Japanese H. magnipapillata served as controls. In two strains of H. attenuata stable hereditary symbioses were obtained with Chlorococcum isolated from H. magnipapillata. In one strain of H. vulgaris, in H. oligactis and in aposymbiotic H. viridis chlorococci persisted for more than a week. Eight species of free-living Chlorococcum, 10 symbiotic and 10 free-living strains of Chlorella disappeared from the brown hydra within 1-2 days. In H. magnipapillata there was a graded distribution of chlorococci along the polyps. In hypostomal cells there were greater than 30 algae/cell while in endodermal cells of the mid-section or peduncle less than 10 algae/cell were found. In H. attenuata the algal distribution was irregular, there were up to five chlorocci/cell, and up to 20 cells/hydra hosted algae. In the dark most cells of Chlorococcum disappeared from H. magnipapillata and aposymbiotic hydra were obtained. Chlorococcum is thus an obligate phototroph, and host-dependent heterotrophy is not required for the preservation of a symbiosis. The few chlorococci that survived in the dark seem to belong to a less-demanding physiological strain. In variance with known Chlorella/H. viridis endosymbioses the chlorococci in H. magnipapillata and H. attenuata were tightly enveloped in the vacuolar membrane of the hosting cells with no visible perialgal space. Chlorococcum reproduced in these vacuoles and up to eight daughter cells were found within the same vacuole. We suggest that the graded or scant distribution of chlorococci in the various brown hydra, their inability to live in H. viridis and the inability of the various chlorellae to live in brown hydra are the result of differences in nutrients available to the algae in the respective hosting cells. We conclude that host/symbiont specificity and the various forms of interrelations we show in green and brown hydra with chlorococci and chlorellae are based on nutritional-ecological factors. These interrelations demonstrate successive stages in the evolution of stable obligatoric symbioses from chance encounters of preadapted potential cosymbionts.