Page:Makers of British botany.djvu/332

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268
HARRY MARSHALL WARD

had no doubt that the bacteroids were the channel of supply. But he failed to get any proof that they could assimilate free nitrogen outside the plant. He suggested that their symbiosis might be an essential condition, and was obliged finally to leave it an open question whether the cells of the tubercles or the bacteroids were the active agents in nitrogen assimilation. He had already stated in 1887 that it is very probable that the bacteroids "may be of extreme importance in agriculture." But he was never satisfied with anything short of the strictest proof.

In 1890 Ward was invited to deliver the Croonian Lecture. He chose for his subject the relation between host and parasite in plant disease. He defined disease in its most generalised form as "the outcome of a want of balance in the struggle for existence." But the particular problem to which he addressed himself was the way in which the balance is turned when one organism is invaded by another. This is the most common type of disease in plants and a not infrequent one in animals. The first result reached was identical with that of Pasteur for the latter; the normal organism is intrinsically resistant to disease. It is an immediate inference that natural selection would make it so. Ward then discusses very clearly the physiological conditions of susceptibility, which he shows to be a deviation from the normal. He had already indicated this in the case of Entyloma. The epidemic phase is reached when the environment is unfavourable to the host but not so or even favourable to the parasite. He then attacks the more obscure case where there is no obvious susceptibility. This, he finds, resolves itself into a mere case of the struggle for existence: "a struggle between the hypha of the fungus and the cells of the host." It is more subtle in its operation but of the same order of ruthlessness as the ravages of a carnivore. Ward's account of the struggle is almost dramatic. The cellulose "outworks" are first broken down, as he had previously shown, by a secreted ferment. The "real tug of war" comes when the hypha is face to face with the ectoplasm. Its resistance is at once overcome by flooding it with a poison, probably oxalic acid.

War with attack and defence is a product of evolution. How did it come about in this particular case? Ward con-