Comparatively little was known of the life history of any Schizomycete. Ward therefore made a detailed and exhaustive study of that of Bacillus ramosus, the Wurzel bacillus of German authors, which is common in Thames water, and bears a superficial resemblance to the anthrax bacillus, but is innocuous. It proved convenient for study, as it ran through its entire life history in from thirty to sixty hours at ordinary temperature. It forms long filaments, the growth of which Ward was able to measure under the microscope with great precision. On plotting out his measurements he obtained a regular curve, from which he found that, under constant conditions, the filament doubled itself in equal times. This he called "the law of doubling." It is the same as the so-called "law of compound interest," and leads to the expression of the growing quantity as an exponential function of the time, so that the time is proportional to the logarithm of that quantity. This relation has, of course, long been familiar in chemical reactions, but, as far as I know, Ward was the first to detect it in any vital process in a plant. This, which was in 1895, has, I think, been overlooked. Stefanowska has since, in 1904, obtained a logarithmic curve for the early period of the growth of maize, which doubles its weight every ten days, and the subject has since been pursued by Chodat and others.
In speculating on the cause of the destructive action of light on bacteria, Ward adopted the view of his friend Elfving, that it inhibited metabolic processes necessary to nutrition. He suggests that the "constructed metabolites" at the moment of assimilation are in a highly unstable condition, and liable to destruction by oxidation promoted by light. He points to the fact that plant structures are frequently provided with colour screens, which would cut off the blue-violet rays and check their action in promoting the rapid oxidation of reserve materials, and he quotes the suggestion of Elfving that chlorophyll itself may serve as such a screen against "destructive metabolic action in synthesis." Ward seems to have attributed little importance to the fact that substantially the same view had long before been put forward by Pringsheim, though received with little favour. His own view that when red and orange predominate in the screens their effect is protective, has since afforded a probable explanation of the colouration of young foliage, especially in the tropics.
It can hardly be doubted that the upshot of Ward's laborious investigations has had a powerful influence in deciding the policy of the future water supply of London. If we hear nothing now of obtaining it from Wales, it is because we know that even polluted flood-water if exposed in large reservoirs will rid itself of its bacterial contamination, partly, as was known already, by subsidence, but most effectually, as shown by Ward, by the destruction of its most deleterious constituents by the direct action of sunlight.
In 1895, Ward was called to the Chair of Botany at Cambridge. He was supported by a distinguished body of fellow-workers, and developed a flourishing school, in which every branch of the science found its scope. The University erected for it an institute which is probably the best equipped in the country, and in March, 1904, I had the pleasure of seeing Ward receive the King and Queen at its inauguration.
During the later years of Ward's life he returned to the study of the Uredineae. The scourge of wheat perhaps from the dawn of agriculture has been "Rust,"
"Ut mala culmos esset rubigo ... intereunt segetes";
and the loss inflicted by it throughout the world is probably not calculable. But the history of the Ceylon coffee disease is only too patent an instance of the injury a uredine can effect.
Eriksson, the most recent authority on the subject, had found himself quite unable to account for sudden outbursts of rust which it did not seem possible to attribute to the result of infection. In 1897 he launched his celebrated theory of the Mycoplasm. He supposed that a cereal subject to rust was permanently diseased and always had been; that the protoplasm of the Uredo-parasite and of the cereal, though discrete, were intermingled and were continuously propagated together; but that while that of the latter was continuously active, that of the former might be latent till called into activity by conditions which favoured it. Ward discussed the theory in his British Association address at Toronto, and was evidently a good deal impressed with it, but nothing short of actual demonstration ever convinced him; and when he proceeded to investigate the actual histological facts on which the theory rested he promptly exploded it.
It is interesting to note that Ward, as I know from correspondence at the time, had himself been embarrassed in investigating the Ceylon coffee disease by the same kind of appearance which had misled Eriksson. It is due to an optical fallacy. When the hypha of a uredine attacks a cell it is unable to perforate it with its whole diameter. It infects it, however, with a reduced and slender filament; this expands again after perforation into a rounded body, the haustorium. In a tangential section the perforating filament cannot be distinguished, and the haustorium looks like an independent body immersed in the cell-protoplasm and with no external connection. It requires a fortunate normal section to reveal what has really taken place. Ward was accordingly able, in a paper in the Phil. Trans. in 1903, to dispose conclusively of the mycoplasm. This cleared the ground of an untenable hypothesis. The complicated nature of the problem which still presented itself for investigation can only be briefly indicated. Sir Joseph Banks, whose scientific instinct was sound but curiously inarticulate, had pointed out that the spores entered the stomata, and warned farmers against using rusted litter. Henslow, one of Ward's predecessors in the Cambridge chair, had been confirmed by Tulasne in showing that the uredo-and puccinia-spores (of the barberry) belonged to the same fungus. De Bary traced the germination of the spores and the mode in which the hyphae invaded the host; the fundamental fact, which he observed but did not explain, was that the germinal filament, after growing for a time superficially, bent down to enter the tissues of its host. Pfeffer in 1883 discovered chemotaxis, the directive action of chemical substances on the movement of mobile organisms. De Bary had previously hinted that the hypha might be attracted by some chemical ingredient of the host plant. Myoshi, a pupil of Pfeffer's, showed finally in 1894 that if a plant were injected by a chemotropic substance a fungus-hypha not ordinarily parasitic might be made to behave as such and attack it.