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Volume 40, 1907
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Art. XXVI.—Notes on the Spread of Phytophthora infestans, with special Reference to Hybernating Mycelium

[Read before the Wellington Philosophical Society, 4th September, 1907.]

The behaviour of a well-known plant-disease under changed environment, or in localities where the disease has not previously existed, is a matter of the most vital importance to the student of plant-pathology. Here in New Zealand we have an exceptionally rich field for the study of the biology of many plant-parasites, for in this country the great majority of our most dangerous plant diseases are aliens, having been in the first place imported on one or other of their hosts.

The wide expanse of ocean which separates these islands from other lands forms a barrier which precludes the suppositon that they could have been introduced by spores or other reproductive bodies blown hither by the wind. Once intorduced, the ecological factors for many of these diseases being pre-eminently suitable fot their requirements, they have thriven here as well as, if not better than, in their original home. Others which in other countries are justly looked upon as deangerous plant-parasites have, on acclimatisation here, been quite unable to cause sufficient damage to rank them as markedly injurious. Others again, which in their native country appear to live more or less in equilibrium with their hosts, have, since their intoruduction here, become virulently epidemic, and are the cause of much annual loss both to the farmer and fruit-grower.

In these notes I shall confine myself to some observations on the ecology of the Irish potato-disease)Phytophtyora infestans).

Intoduction into New Zealand

There are no definite records of when and how Phytophthora infestans was first introduced into this country. On the epidemic outbreak of this desease in November, 1904, Professor Thomas made the following satement: “The same disease (Phytophthora infestans) appeared some twelve years ago, but it was not so prevalent as on the present occasion.” Furtehr, he sasys, “Moreover, it is no new thing here, having been in the country, to my certain knowledge, for the past twelve years.” Mr. T. W. Kirk, in the report of the Department of Agriculture for 1905, writes, “Twelve years ago there was a mild outbreak

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in the Auckland Province, but it has not been heard of since till last year.” For my part, I condider that the epidemic outbreak in Auckland in 1904 was in no way connected with the sporadic ones which occurred previously, The cause can be attributed to the fresh importation of Phytophthora mycelium in imported potatoes. There are only two ways that Phytophthora can possibly have been introduced here—either by resting mycelium in diseased tubers, or by asesual spores or oospores; but as these latter have never been definitely discovered, its introduction by means of them is most unlikely. The asexual spores of P. infestans are naturally short-lived, and are in no way provided with any adaptations to withstand the desiccation that they would undergo in passing though the tropics. Therefore it is almost certain that the introduction of thisdisease into New Zealand was by means of dormant mycelium hidden away in the tissues of affected tubers. This view is greatly strengthened by the discovery, on numerous occasions during the past two years, of varieties of imported potatoes in which the presence of Phytophthora mycelium was clearly demonstrated.

All potatoes which are now imported into New Zealand are carefully examined by the Agricultural Department, and those lines found affected with Phytophthora are at once destroyed.

Action of Phytophthora on Potato-Tubers.

There are still many gaps in our knowledge of the full life history of the Irish potato-disease, and the exact pathological processes that obtain in so-called diseased tubers are but imperfectly understood. That the disease is transmitted from season to season by means of mycelium permeating the tissues of the tubers themselves hass been now abundantly proved. Massee has given the name “hybernating mycleium” to that portion of the vegetative body of a fungus which has the power of remaining quiescent during the dormant period of the host's existence, and which can return to normal development as soon as the host commences to devleop. It follows naturally that those fungi which are able to develop hybernating mycelium do no require the same amount of varied spore-formation as those which are not so equipped. The formation of winter spores be decidedly superfluous, and such fungi can be more specialisedin the direction of producing summer and generally short-lived spores, whose object is to rapidly infect large mases of their hosts, provided the environment is suitable.

The finest example of hybernating myucelium is found, as Freeman has shown, in the fungus affecting the various species of Lolium, especially L.temulentum. In over 70 per cent. of

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the seed of this weed that I have examined masses of resting mycelium have been found in the tissue interior to the aleurone layer. This mycelium remains inactive until the Lolium seed begins to germinate, when it develops and keeps pace with the growth of the host, and finally re-forms resting mycelium in the developing seed. No mode of spore-formation, either sexual or asexual, has ever been noted, and the fungus appears to live in perfect harmony with its host. This almost symbiotic union between the fungus and hostis of the utmost biologic importance in the economy of hybernating mycelium, for it is at once apparent that if the fungus can live on its host throughout the dormant season without causing any marked injury, there is all the more chance of the host developing in a normal and more or less healthy manner during the next season, and thus allow the peerpetuation of the fungus; whereas if the resting mycelium caused serious pathological changes in the host, the latter woul probably be killed outright, and the mycelium would die at the same time.

A considerable amount of material has been examined by me during the past three years, showing both the ordinary and the hybernating mycelium of Phytophthora infestans, and a certain amount of interesting information on the perpetuation of this fungus has thus been gathered together.

The exact manner by which the mycelium of P. infestans reaches the tuber has not been as yet satisfactorily ascertained, and for this purpose detailed and careful examination in the field would be necessary. Two methods have been suggested—firstly, that the mycelium spreads from the leaf downwards through the stem until it reaches the tuber; and, secondly, that spores developed on the conidiophores fall to the ground, and are washed by rain or carried by other agencies directly on to the surface of the tubers. I am inclined to think that both these porcesses occur in nature, but hat the mycelium, which descends down the stem and then enters the tubers, alone forms hybernating mycelium, and that the spores which reach the tubers by mechanical and other means do not develop into resting mycelium, but are more or less directly responsible for the rapid rotting that so often occurs with Phytophrhora attack. This view gains great weight from the fact that healthy tubers on the surface of which Phytophthora spores are scattered, but on which resting mycelium has not been observed by me, rapidly develop a rot. Recently Matruchot and Molliard have declared that Phytophthora does not of itself cause a rot in potato tubers, but that after the tubers become affected the rot that sets in is due to microbes that become associated with the P. infestans. This view, notwithstanding the high authorities from

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whence it has originated, I am inclined to combat; although I must admit that in the majority of the cases I have examined secondary infection by bacteria and other fungi, notably Fusarium oxysporum, plays an extremely important part in the rotting that occurs in tubers which have been primarily attacked by P. infestans. It is a well-known fact that tubers bearing the characteristic marks associated with Phytophthora attack often during the whole winter without any trace of rot setting in. When these are examined under a miagnification of about 100 diameters, large amounts of dormant mycleium will be seen in those portions of the tissues of the tubers that abut on the darkened and discoloured areas, which are said to be caused by Phytophtora; and in such cases no other fungi or bacteria will be found associated with the Phytophthora mycelium. This observations gains considerably in significance when it is stated that in those tubers on which an active rot is present resting mycelium can hardly ever be discovered, although there will be an abundance of bacteria and other fungi, both parasitic and saprophytic.

If slices of potatoes showing resting mycelium of Phytophthora infestans are placed in petri dishes, and kept moist in a temperature of about 600 to 700 Fahr., the mycelium will rapidly become active, and in a few days an abundant crop of spores will be developed. This shows that the mycelium is not in a very dormant condition, but is really on the border-line between active and hybernating mycelium. This is a very important point, inasmuch as it shows the liability, under certain conditions of the mycelium to become active even when no growth on the part of the host takes place, and in this particular distinguishes it sharply from the more specialiesed resting stages in the life-history of other fungi, such as many of the Ustilagineœ.

When tubers with the resting mycelium of P. infestans are planted, the fungus develops rapidly through the tissues of the developing plant, and if the weather is humid and warm the mycelium becomes markedly negatively geotropic, and if the conditions remain favourable for its development it quickly makes its way into the leaves, on the under surfaces of which it soon produces an abundant supply of spores, which, blown by the wind, can soon spread infection far and wide. If, however, the weather-conditions remain unfavourable for its development, no spores at all may be produced, and to all outward appearance the potato-plants remain quite healthy. In such a case it is not known wheather the fungus can again form resting mycelium without the intervention of a spore-producing stage, as is done in the case of Lolium temulentum, but I am inclined to consider that such can and often does occur.

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The Tropic Movements of Phytophthora Mycelium

A peculiar feature of the biology of the mycelium of Phytophthora infestans is that, after primary infection has taken place on the leaf or stem, it is markedly positively geotropic. On the other hand, as has been mentioned previously, the mycelium developing from the dormant portion becomes negatively geotropic. For my part, I attribute this not to the action of gravity but to the result of chemotactic stimulus, and that the mycelium in all cases follows the direction in which food materials are stored. This would account for the apparently contradictory influence which gravity has up to the present been considered to produce.

An important point, and one which appears to have been lost sight of, is the fact that in tomatoes attacked by Phytophthora the direction of the mycelium is in general negatively geotropic, or, as I take it, the mycelium is chemotactically attracted by the food material stored in the fruit; whereas in the potato the mycelium is positively geotropic, being attracted downwards by the chemotactic stimulus of the tubers.