It must be concluded that the formation of the Vaucheria gall is induced by the mechanical irritation which the Rotifer causes in the protoplasm. These galls are comparable to the hypertrophies in Pilobolus caused by the presence of Pleotrachelus.
Attempts to induce the development of galls artificially by injecting formic, acetic and other vegetable acids, poisons and other substances into the tissues have, however, failed, and even the substances contained in the insect or gall itself only produced negative results. Nothing further was obtained than slight callus formations in some cases. Nor have experimenters succeeded in obtaining more than slight distortions by fixing insects on the growing leaves in such positions that they could scratch the epidermis.
We must therefore conclude that very complex interactions between the plant and insect are here concerned, among which may be the infiltration of some liquid from larva to plant—many of these gall larvae are strongly scented, and Kustenmacher says that fluids excreted by the larva are absorbed by the gall-tissue apparently as nutriment. This would point to the symbiotic character of galls and their guests.
Notes to Chapter XIV
With regard to the action of poisons in small doses see further Johannsen, Das Aether-Verfahren beim Fruhtreiben, Jena, 1900, and, for Botrytis, see Marshall Ward, "A Lily Disease," Annals of Botany, Vol. II., 1889, p. 388.
The subject of enzymes has been exhaustively treated by Green, The Soluble Ferments and Fermentations, Cambridge, 1899, to which the reader is referred for literature. I have taken the statements regarding Fontaria and Dolium from Kassowitz, Allgemeine Biologie, p. 182. The two most important works on chemotactic phenomena are Pfeffer, "Uber Chemotaktische Bewegungen," etc., Unters. aus dem Bot. Inst. zu Tubingen, B. II., p. 582, and Miyoshi, "Die Durchbohrung von Membranen durch Pilzfaden," Pringsh. Jahrb. f. Wiss. Bot., B. XXVIII., 1895, p. 269, and from these the further literature can be traced. As regards the nature of parasitism see Marshall Ward, "On Some Relations between Host and Parasite," etc., being the Croonian Lecture delivered before the Royal Society, Proc. Roy. Soc., Vol. 47, p. 393. On Symbiosis, see Marshall Ward, "Symbiosis," Annals of Botany, 1899, Vol. XIII., p. 549, where the literature is collected. For a general account of galls the reader may consult Kerner, The Natural History of Plants, Eng. ed., 1895, Vol. II., pp. 527-554, and Adler, Alternating Generations, A Biological Study of Oak Galls, etc., 1894.
CHAPTER XV.
SPREADING OF DISEASE AND EPIDEMICS
Dissemination of fungi by the aid of snails, rabbits, bees, and insects—Man—Distribution in soil, on clothes, through the post, etc.—Worms, wind—Puffing of spores—Creeping of mycelia—Lurking parasites—Spread of insects and other animals—Losses due to epidemics.
The dissemination of plant diseases is a subject which has been far too much neglected, but our knowledge of it is slowly increasing. The spores of fungi such as Rusts and Erysipheae are often carried from plant to plant by snails; those of root-destroying and tree-killing Polyporei by rabbits, rats, and other mammals which rub their fur against the hymenophores. Bees have been shown to carry the spores of Sclerotinia and infect the stigmas of Bilberries, etc., with them; and flies convey the conidia of Ergot from grain to grain. Insects, indeed, of all kinds are great disseminators of disease—as witness also the part played by mosquitoes in transferring the malaria parasite to man—and beetles, bees, flies, etc., of all sorts probably play more active parts in this work than has yet been proved, since they not only carry spores attached like pollen to their hairy bodies, but in many cases in their alimentary canal, to be spread later in the dung.
The part played by man in conveying fungi from plant to plant counts for much. Not only do gardeners and farm labourers carry spores on their boots and clothes as they pass from infected to non-infected areas, but carted soil and manure are frequently infested with spores of Smuts, Fusarium, Polyporus, and the sclerotia or rhizomorphs of Sclerotinia, Agaricus melleus, Dematophora, etc. Man also sends diseases through the post, and by rail and ship, by spores or mycelia attached to seedlings, bulbs, fruits, flowers, etc., as shown in several cases of potato, vine, hollyhock, lily, and hyacinth diseases. Every time a carpenter saws a piece of fresh timber with the saw which has been used previously for cutting wood attacked with dry rot, he risks infecting it with the fungus. Similarly in pruning: every cut with a knife which the gardener has used on infected branches may infect the tree.
Cuttings made with a soil-contaminated knife and stuck into ordinary soil in dirty boxes covered with equally dirty glass, present every chance for infection by soil organisms; bacteria and fungi obtain access to the vessels, and derive plenty of food from the juices, and the wonder is not that so many cuttings "damp off," but that any are raised at all under ordinary conditions.
That worms bring buried spores to the surface can hardly be doubted after Pasteur's experiments with Anthrax, and the principle of Darwin's discoveries of the important bearing of the habits of earthworms on this subject, and that the soil attached to the feet of ducks and other birds teems with small seeds, applies to fungi also. Wind is also responsible for distributing fungus-spores over wide areas, as may be easily proved by fixing a glass slide smeared with glycerine in the course of a breeze passing over an infected area.
But although the fungi are, generally speaking, passive in regard to their distribution, such is by no means always the case. Apart from the fact that some forms attract insects by means of honey dew (Ergot), or by sweet odours (Spermogonia, Sclerotinia), the zoospores of Pythium, Phytophthora, etc., are motile, and although they cannot move far in the films of water in which they travel, nevertheless in a wet potato field, with the wind flapping the leaves one against the other, some dissemination of importance must be actively brought about, and similarly with the amoebae of Plasmodiophora in the soil.
The shooting of ascospores into the air by certain species of Peziza, from the discs of which the spores may be seen to puff out in clouds, affords further evidence that fungi cannot be regarded as entirely passive in respect to distribution of their spores. But when we come to certain of the soil fungi—e.g. Agaricus melleus, Dematophora, etc.—the active creeping forward by growth in the soil of their rhizomorphs and mycelial strands afford examples of active spreading of considerable importance in the vineyard and forest, since they pass from root to root and from tree to tree and may infect the entire area in course of time.
Not the least significant mode of dissemination is that by which what I have termed "lurking parasites" are spread: such are fungi which attach themselves to the seeds, fruits, tubers, etc., of other plants and so obtain all the advantages of being carried and sown with the latter—e.g. Ustilagineae and Uredineae which adhere to grain, Verticillium, Nectria, etc., in potatoes and other plants.
The spread of diseases due to animals, especially insects, is of course more active, in consequence of the motility of the distributing agents. This is most marked in the winged species, of which locusts, beetles, moths and butterflies, flies and wasps furnish well-known examples; and is not inconsiderable in the case of wingless and merely creeping species. It is noteworthy that many forms wingless in the parasitic stage are winged at certain periods, e.g. the females of Phylloxera.
That man also spreads insect pests is well known and acted upon, as witness the phylloxera laws—which, however, it is to be feared too often only illustrate once more the adage concerning the shutting of the stable door after the horse has gone.
It would be tedious to attempt anything like a complete account of the estimates of loss in different countries, due to the ravages of insects and fungi, but the following examples should surely serve to convince anyone of the magnitude of these losses and of the economic importance of the whole question, and the reader may be referred to the special literature for further details.
The coffee leaf-disease of Ceylon, due to the fungus Hemileia, is estimated to have cost that Colony considerably over £1,000,000 per annum for several years. One estimate puts the loss in ten years at from £12,000,000 to £15,000,000. The hop-aphis is estimated to have cost Kent £2,700,000 in the year 1882. In 1874 the Agricultural Commissioner of the United States estimated the annual loss, due to the ravages of insects on cotton alone, to amount to £5,000,000; and in 1882 the annual loss to the United States due to insects, calculated for all kinds of agricultural produce, was put at the appalling figure of from £40,000,000 to £60,000,000 sterling. In India, the annual loss due to wheat-rust alone has recently been estimated at 4,000,000 to 20,000,000 rupees, and one insect alone is said to have cost the cotton planters a quarter of the crop—valued at seven crores of rupees—in bad years. Similarly, in Australia the annual loss from wheat-rust has been put at from £2,000,000 to £3,000,000. In 1891 the loss in Prussia alone from grain-rusts was officially estimated at over £20,000,000 sterling. Need more be said? Even allowing for considerable exaggerations in such estimates it is clear that the damage to crops in any country soon amounts to sums which even at low rates of interest would easily yield incomes capable of supporting the best equipped laboratories and staffs for investigations directed to the explanation of the phenomena in detail, the sole basis on which intelligent preventive and therapeutic measures can be based. But it is far from likely that the estimates are exaggerated. The planting and agricultural communities are as a rule opposed to the publication of statistics—or at least have been so in various countries and at different times—and if we knew the damage done to all crops even in our own Empire, the results would probably astonish us far more than the above figures have done.
Notes to Chapter XV
On the dissemination of fungi, the reader will find Fulton, "Dispersal of the Spores of Fungi by the Agency of Insects," Ann. Bot., Vol. III., 1889, p. 207, and Sturgis, "On Some Aspects of Vegetable Pathology and the Conditions which Influence the Dissemination of Plant Diseases," Botanical Gazette, Vol. XXV., 1898, p. 187, both useful papers. Further information will be found in Zopf, Die Pilze, Breslau, 1890, pp. 79-95 and 228, and Wagner, "Ueber die Verbreitung der Pilze durch Schnecken," in Zeitschr. f. Pflanzen Krankh., 1896, p. 144. The estimates as to losses due to epidemics are taken from Watt, Agricultural Ledger, Calcutta, 1895, p. 71; Balfour, The Agricultural Pests of India, London, 1887, pp. 13-15; Eriksson and Henning, Die Getreideroste; the publications of the U.S. Department of Agriculture, The Kew Bulletin, and elsewhere. The reader will find further examples in Massee, Text-Book of Plant Diseases, 1899, pp. 47-51. Both these subjects are well worth further attention, and I know of no complete account of them.
CHAPTER XVI.
THE FACTORS OF AN EPIDEMIC
Illustrations afforded by the potato disease—The larch disease—The phylloxera of the vine.
When we come to enquire into what circumstances bring about those severe and apparently sudden attacks on our crops, orchards, gardens, and forests by hosts of some particular parasite, bringing about all the dreaded features of an epidemic disease, we soon discover the existence of a series of complex problems of intertwined relationships between one organism and another, and between both and the non-living environment, which fully justify the caution already given against concluding that any cause of disease can be a single agent working alone.
The statement of prophecy that a particular insect or fungus need not be feared, because it is found to do so little harm in particular cases or districts examined, will thus be seen to be a dangerous one: any pest may become epidemic if the conditions favour it!
In 1844 and 1845 the potato disease assumed an epidemic character so appalling in its effects that it is no exaggeration to say that it constituted a national disaster in several countries. It was stated at the time that this disease had been known for some time in Belgium, in Canada and the United States, in Ireland, in the Isle of Thanet, and in other parts of the world. Similar, but less devastating epidemics have occurred in various years since. It was generally noticed during such epidemics that the plants themselves were full of foliage, surcharged with moisture, and of a luxuriant green colour promising abundant crops. The now well-known spots, at first pale and then brown and fringed with a whitish mould-like growth—the conidiophores of the Phytophthora—were observed during the dull cloudy and wet weather, cooler than usual, when the atmosphere was saturated for days together, in July and August. The actual amount of rain does not appear to have been excessive, but most observers seem to agree that dull weather with moist air had succeeded a warm forcing period of growth. So rapidly did the disease run its course that in a few days nearly all the plants were a rotting blackened mass in the fields, and the potatoes dug up afterwards were either already rotten or soon became so in the stores. Further experience has confirmed this, and we now know that the epidemic is very apt to appear in any region where potatoes are grown on a large scale, in dull moist weather, especially in fields exposed to mists, heavy dews, etc., about July and August, when the foliage is full and turgid. Similarly on heavy wet soils, unless the season is remarkably open and dry; but also on dry light soils in rainy seasons. So evident was this that many believed that the mists and dew brought the disease—harking back to the superstitions of earlier days. We must remember that prior to 1860 the life-history of Phytophthora was not known. Since De Bary's proof of the germination of the zoospores and of the infection of the leaves, the course of the hyphae in them and in the haulms, the origin of the conidia, etc., and the confirmation by numerous competent observers of the true fungus nature of this disease, we are now in a position to understand the principal factors of the various epidemics of potato disease.
It is not merely that the potato-fields afford plenty of food for the fungus, and that the dull weather causes the tissues to be surcharged with moisture, owing to diminished transpiration, but the mists and dew—to say nothing of actual rain and the flapping of wet leaves—favour the germination and spread of the zoospores throughout the field. Whether the dull light also favours the accumulation of sugars in the tissues, and the partial etiolation of the latter implies less resistance to the entering hyphae, may be passed over here, but in any case it is clear that we have several factors of the non-living environment here favouring the parasite and not improving the chances of the host, even if they do not directly disfavour it.
As another instance I will take the Larch-disease, which is due to the ravages of a Peziza (Dasyscypha Willkommii) the hyphae of which obtain access by wounds to the sieve-tubes and cambium of the stem, and gradually kill them over a larger and larger area and so ring the tree, with the symptoms of canker described below.
Now the Larch fungus is also to be found on trees in their Alpine home, but there it does very little damage and never becomes epidemic except in certain sheltered regions near lakes and in other damp situations. How then are we to explain the extensive ravages of the Larch disease over the whole of Europe during the latter half of this century? The extensive planting, providing large supplies for the fungus, does not suffice to explain it, because there are large areas of pure Larch in the Alps which do not suffer.
In its mountain home the Larch loses its leaves in September and remains quiescent through the intensely cold winter, until May. Then come the short spring and rapid passage to summer, and the Larch buds open with remarkable celerity when they do begin—i.e. when the roots are thoroughly awakened to activity. Hence the tender period of young foliage is reduced to a minimum, and any agencies which can only injure the young leaves and shoots in the tender stage must do their work in a few days, or the opportunity is gone, and the tree passes forthwith into its summer state.
In the plains, on the contrary, the Larch begins to open at varying dates from March to May, and during the tardy spring encounters all kinds of vicissitudes in the way of frosts and cold winds following on warm days which have started the root-action—for we must bear in mind that the roots are more easily awakened after our warmer winters than is safe for the tree.
It amounts to this, therefore, that in the plains the long continued period of foliation allows insects, frost, winds, etc., some six weeks or two months in which to injure the slowly sprouting tender shoots, whereas in the mountain heights they have only a fortnight or so in which to do such damage. That the lower altitude and longer summer are not in themselves inimical to Larch is proved by the splendid growths made by the trees first planted a century ago. Then came the epidemic of Larch-disease: the fungus, which is merely endemic—i.e. obtains a livelihood here and there on odd trees, or groups of trees in warmer or damper nooks—in the Alps, was favoured by the more numerous points of attack afforded to its spores by injuries due to insects—Coleophora, Chermes, etc.—and frost wounds, as well as by the longer periods of moist dull weather, and the longer season of foliation. Moreover, as time went on almost every consignment of young Larch-trees sent abroad was already infected. Here again, then, we find the factors of an epidemic consisting in events which favour the reproduction and spread of a fungus more than they do the well-being of the host.
As a third illustration I will take the case of an insect epidemic. In 1863 a disease was observed on vines in the South of France which frightened the growers as they realised its destructive effects: the roots decayed and the leaves turned yellow and died before the grapes ripened, and such vines threw out fewer and feebler shoots the following year, and often none at all afterwards. In 1865 the disease was evidently becoming epidemic near Bordeaux, and in 1868 it was shown to be due to an insect, Phylloxera, the female of which lays its eggs on the roots, where they hatch. The louse-like offspring sticks its proboscis into the tissues as far as the central cylinder. The irritated pericycle and cortex then grow and form nodules of soft juicy root-tissue at which the insect continues to suck. Rapid reproduction results in the majority of the young rootlets being thus attacked, and since they cannot form their normal periderm and harden off properly they rot, and admit fungi and other evils, in consequence of which the vine suffers also in the parts above ground.
Evidence that the general damage is due to the diminished root-action is found in the peculiarly dry poor wood formed in the "canes" of diseased plants.
By 1877 the epidemic had spread to the northern limits of the French vineyards, and by 1888 half the vines in the country were attacked, and the yield of wine reduced from half a million hectolitres to 50,000 only. Meanwhile the disease had spread to Italy, Germany, Madeira, Portugal, and even to the Cape, though not in epidemic form as in the Bordeaux centre whence it spread.
Now it appears that Phylloxera has long been in the habit of doing damage to vines in America, where, however, it attacks the leaves, on which it makes pocket-like galls, rather than the roots. Moreover, there are species and varieties of American vines which, even when planted in Europe, do not suffer at all from this insect at the roots, either because the rootlets do not push out at the same season as those of the European form, or because they form wood more rapidly and completely, or secrete resinous and other matters distasteful to the insect in greater quantity and are thus capable of healing the wounds, or in some other way they do not respond to the attack or suit the insect. In any case the attack on the leaf rather than the root seems to be the exception in European vineyards and the rule in American species, and we appear to be face to face with a problem of specific predisposition to this particular malady. That the resistant properties of the vines of America—not all, only particular species and varieties are thus "immune"—can be utilised has been proved by European growers; and not only so, for Millardet and others have shown that the European vine grafted on to these resistant stocks suffer less than when on their own roots. It has also been shown that hybrids can be obtained which are resistant.
But the most curious point of all is that Phylloxera was itself a native of America, and came thence to Europe. It had played its part with certain fungi in ruining all the attempts to introduce the European vine into America many years ago. A recent authority on the evolution of American fruits writes as follows:
"All the most amenable types of grapes had long since perished in the struggle for existence, and the types which now persist are necessarily those which are, from their very make-up or constitution, almost immune from injury, or are least liable to attack . . . the Phylloxera finds tough rations on the hard, cord-like roots of any of our eastern species of grapes. But an unnaturalised and unsophisticated foreigner, being unused to the enemy and undefended, falls a ready victim; or if the enemy is transported to a foreign country the same thing occurs."
Further proof that it is in the "constitution" of the European vine that the want of resistance to Phylloxera resides, is furnished by the fact that in California and the Pacific states the European vine was introduced with more success, but is now suffering badly because Phylloxera has spread there also. It must not be overlooked, however, that we are as yet very ignorant of all that is implied in the word "constitution" as used above.
If we enquire further why the Phylloxera epidemic was so much worse in the Southern vineyards than in the more Northern ones of Germany, the opinion seems to prevail that the warmer climates favour the insect. Further, it appears that, in Italy, the vines in loose open soil, provided it is equally rich in mineral food-materials and offers no disadvantages as regards drainage, suffer less than those in closer soils, the reasons alleged being that the young roots can push out more rapidly and widely, and so obtain holdfasts with greater distances between them.