Skip to main content

Full text of "Text-book of the diseases of trees"

See other formats

Universityof Oxford 

SiBTHORPiAN Library 


*^ BY 

William Somerville kbedsc 



Digitized by tine Internet Archive 

in 2009 with funding from 

NCSU Libraries 


















The Right of Translation and Reprodtcciion is Reserved 

Richard Clay and Sons, Limited, 
london and bungay. 


[By H. MARSHALL WARD, D.Sc, F.R.S., F.L.S., F.R.H.S.] 

The foundation of a science of Mycology by Berkeley, de 
Bary, and Tulasne, pursued by Brefeld, Zopf, and others, has 
led to a knowledge of the biology of fungi highly creditable 
to the industrious observers who have explored this domain of 
the vegetable kingdom ; while the gradual building up of the 
science of plant-physiology from the days of Knight and Hales, 
De Saussure and Boussingault, to those of Sachs and Pfeffer, 
has placed us in possession of a vast amount of information 
as regards normal life processes in plants. Until much more 
recently, however, it cannot be said that we have had a science 
of the pathology of plants — i.e. the study of abnormal physio- 
logy — of anything like the same importance, in spite of the 
splendid and progressive attempts of Berkeley, Frank, and 
Sorauer to found one. 

In the particular department he has cultivated, Robert Hartig 
has succeeded in founding a plant-pathology really worthy 
of the name, and I would especially emphasize this, that his 
researches are so thoroughly elucidative of pathological pheno- 
mena, in that he studies not only the nature of the structural 
lesions and of the physiological disturbances consequent on 
these, but also the factors of the environment which throw light 
on the question. No better illustration of this could be selected 


than his admirable discussion of the complex phenomenon of 
"Leaf-casting" on pp. 1 10-117 of the present work ; while his 
work on the ZersetzmigserscJieimingen des Holzes is a model of 
thoroughness and scientific accuracy and acumen to which all 
workers in this branch of botany must look up. 

In this country we are awakening rapidly to the necessity 
of placing ourselves abreast of the new ideas involved in this 
compound study of plant-pathology ; but we are perhaps as yet 
by no means so alive to the practical importance of the new 
discipline as it might have been inferred (from our national 
pride in being practical) we should be. 

Animal-pathology is studied with zealous and expensive 
enthusiasm — I suppose because, being animals, we are at once 
alive to its importance ; but plant-pathology, in the real sense 
of the term, scarcely obtains recognition as yet, no doubt owing 
to our interest in the culture of trees and agricultural produce 
having seemed to be less pressing than coming events are likely 
to prove it really to be. From this condition — only apparently 
apathy — we are doubtless awakening, and, as is usual, when we 
English do awake to new necessities, we at once enthusiasticall}- 
set to work to recover lost ground. 

Now, in this department, we have to awaken to some startling 
new facts, the practical bearings of which have deeply impressed 
our Continental cousins for some years past. 

One of these new facts is that we may know a very great 
deal about the systematic position and the morphology of 
parasitic and destructive fungi without knowing much — or, 
indeed, necessarily anytJiing — about the diseases and injuries 
the}^ induce ; another of these new facts is that pathology — 
i.e. the study of disease — cannot be fruitful unless the student 
is experimentally acquainted with plant-physiology, and espe- 
cially (though by no means only) the physiology of nutrition. 

To these statements I would add that we have, as a nation, 
to force ourselves even more than we have yet done out of 
the groove in which plant-physiology is looked upon as a mere 
branch of chemistry and physics. It is no undervaluing of 


the true status of agricultural chemistr}-, or of the study of 
the physics and chemistry of soils, &c., to insist upon it that 
no one can appreciate even the rudiments of plant-physiology 
Avho does not make himself master of the facts of structure 
and the essential phenomena of life by experimental investiga- 
tion ; nor to point out that, as we come to know more about 
the physiology and pathology of plants, we learn that the 
chemistr}' of the soil is one of the least important factors we 
are concerned with. 

These truths have to be faced, and in spite of the at first 
sight depressing inference that the study of plant-pathology 
— and I suppose the same applies to animal-pathology — de- 
mands rigorous and active acquaintance with several other 
branches of science. As I have stated in substance elsewhere, 
we demand that the surgeon or doctor who attends us shall be 
qualified properly to do his work, though we are perhaps not 
always alive to the extravagance of our demands on his ability 
and training ; and just as we cannot expect him to do his work 
in diagnosing the disease or injury, and explaining and com- 
bating or removing its cause, unless he is properly qualified, 
by the requisite instruction in the physics, chemistry, structure, 
and normal working {i.e. physiology) of the healthy body and 
in the pathology of the case concerned, so can we as little 
expect any one to deal with diseased conditions in plants who 
is ignorant of *their structure and physiology, and the patho- 
logical conditions of the case concerned. There is, however, 
the comforting assurance that the processes in plants, complex 
as they are — and we must not err in underrating this — are 
simpler than in animals, and must throw useful lights on all 
general problems in Biology. 

The objection that there are epidemic diseases of plants 
which we have as yet failed to prevent or overcome is obviously 
no more valid than the cry that we cannot as yet stay the 
progress of epidemics of influenza or cholera : the cases are 
exactly parallel, and since we do not abandon or depreciate 
the study of medicine because medical science is not yet in a 


position always to cope as successfully as we could wish with 
human ailments, so we must not undervalue the importance 
of the triumphs of the much younger baby science — plant- 

Probably few people in this country are really aware of the 
enormous strides towards lusty and vigorous youth the new 
science is now making, and what important contributions to 
human progress its study is affording. 

Educated and properly trained agriculturists and foresters 
have long been familiar with the fact that great advances are 
being made in these directions, and perhaps the only fault 
that a very severe critic could find with them is that they have 
remained a little too long deterred by the failures which have 
always to be acknowledged (and met) in a progressive experi- 
mental science. That much more general interest in this pro- 
gress is now evident, however, is best proved by the various 
publications on the treatment of plant-diseases which are 
springing up around us ; for even the very sceptical will admit 
that, on the one hand, the treatment of diseases depends on 
knowledge of them and their origin, and, on the other hand, 
that such eminently practical works would not be published 
unless they were read. 

Foremost among such publications are the reports of the 
various experimental stations on the continents of Europe and 
America, and it is a matter of the highest credit and con- 
gratulation that the Americans are devoting large sums of 
money to the experimental study of methods of treatment, 
based on knowledge of the diseases treated, at several of their 
enthusiastically planned experimental stations. I need only 
point to the reports published by the United States Department 
of Agriculture (Section of Vegetable Pathology), and to the 
ZeitscJirift fur Pflanzen-Krankheiteu emanating under the 
auspices of the " Internationalen Phytopathologischen Kommis- 
sion," as showing how necessary it is becoming to have special 
organs in this branch of science, and to the increasing number 
of text-books on the subject of plant-diseases and their treat- 



mcnt now being published, as evidence for the justice of the 
above statement. 

Among these latter, Kirchner's Die Krankheitcn und BescJidd- 
igiingen jmserer LandivirtJiscJiaftlicJien Pflanzcn stands high 
in the list as a treatise on the diagnosis and treatment of 
agricultural and horticultural plant-diseases. Far more of a 
classic, however, but dealing more especially with the diseases 
of forest trees, is Robert Hartig's beautiful text-book now intro- 
duced to the English public. Another encyclopaedic work, Hess's 
FortscJmtz, deals more particularly with the " dodges," if I 
may use the word, practical foresters are devising to combat 
the principal maladies to which forest trees are subject, and 
of this work English readers are also promised a translation 
at an early date. These more special treatises may be men- 
tioned as supplementing — and in part supplanting — the more 
academical and general works of Frank and Sorauer previously 
referred to. 

The great charm of Hartig's book lies as much in the ex- 
cellent plan and simple method of exposition of the facts and 
principles concerned, as in the astounding richness of infor- 
mation it conveys. This is unquestionably owing to Hartig's 
prominence as the leading investigator and authority in Ger- 
many in this special branch of knowledge — the fungoid diseases 
of forest trees. 

His pre-eminence as a sturdy and patient inquirer, as an 
admirable anatomist and physiologist, and as a bold and 
original thinker, is well known to the few who are acquainted 
with his special scientific publications, particularly his laborious 
memoir on the destruction of timber by fungi. In the present 
work I think he shows himself also a master of the art of 
teaching the principles as well as the facts of his subject. 

Of course there are points of view to be considered in 
criticising such a book. 

The specialist on the Morphology of the Fungi will probably 
complain of the author's classification of these organisms, and 
of his somewhat lax use of certain morphological terms ; but 


it should be remembered that, in the first place, the student's 
attention is not here directed to the fungus itself, as an object 
of morphological study, so much as to the action of certain 
fungi in inducing specific diseases in trees ; and, in the second 
place, it is assumed that the student is already acquainted with 
the main facts in the biology of the fungi — including their 
morphology — before he attempts this particular branch of 

Again, the professed botanist may remark how much is 
assumed concerning the structure and physiology of the host- 
plants — the trees — whose diseases are here treated of. The reply 
is, as before, the student cannot extract all, or nearly all, of 
value from such a work as this, unless he is thoroughly 
acquainted with the principal facts of the normal anatomy 
and physiology of the higher plants. 

A third platform of criticism is that of the " practical 
forester," who may object that the author gives too little 
information as to jhe details of combative or therapeutic 
treatment of the special diseases. To this the obvious reply 
is that it is not necessarily the duty of the scientific pathologist 
to devise the particular mode of attack to be employed in 
special cases — these plans of remedial treatment involve the 
outlay of money, labour, &c., which vary in different countries 
and in different cases, and enough has been done by the in- 
vestigator who indicates the factors involved. Special works 
must be consulted regarding the details of treatment, though it 
seems to me the author, while clearly recognising this, goes even 
out of his way to give practical hints as to treatment, and has 
in many cases put the principal factors concerned in the treat- 
ment so clearly that every thinking practical man can do the 
rest himself. No better illustration of the thoroughly practical 
nature of his writings could be selected than his recommenda- 
tions for the treatment of Dry-rot. 

But it is by no means solely on the ground of the information 
capable of direct application which the book contains that it 
should be judged. I would especially urge the value of this 


study to the student of botany as calculated both to test his 
knowledge in other departments of his science and to open 
out new lines of thought as he considers the interactions 
between one plant and another, and between both and other 
factors — living or not — of the environment. 

Hartig's ingenious explanation of the spread of the well- 
known Larch -disease may or may not be accepted in all its 
details, on the evidence given, and dissent from his explanations 
of such diseases as " Canker," &c., has been expressed, but I 
would maintain — apart from my acceptance of the general truth 
of his arguments — that they teach the student very clearly how 
to investigate and think out these complicated matters for 

We are still decidedl}' wanting in information concerning 
many diseases of trees. Standing elms and other trees are 
occasionally found in this country with a species of Hymeno- 
mycete growing from the trunks six feet or more above the 
ground : are these parasitic or not, and what is their mode of 
action .'' How does Polyponis squainosus attack timber .-• What 
are the exact biological relations of Polyponis fomentarius, 
Fistidina Jiepatica, and a number of other forms found in this 
country, to the trees on which they grow .'' 

These and numerous other questions await solution, by means 
of thorough investigations properly conducted in this country, 
along such lines as Hartig has laid down in Germany, and it 
should be borne in mind that such studies offer stores of facts 
likely to be of the utmost interest to investigators in other 
branches of Botany. To give one instance only : the study 
of the destruction of the walls of the tracheids, and other 
elements of which timber is composed, by the h}'phae of fungi, 
shows that there is considerable variety in the processes of 
piercing, delignifying, corroding, and dissolving them, and it 
seems a safe conjecture that valuable information as to the 
intimate structure of these walls may be derived from the 
examination of the way the fungus unbuilds them, so to 


Among the Ascomycetes and the Uredineae are numerous 
questions already framed for investigation, and I know of no 
department of botanical study more fascinating than the scien- 
tific hunting for the hetercecious forms of ^cidiomycetes : the 
work combines all the excitement of a true hunt with the 
intense intellectual pleasure implied in the demand for the 
severest critical observation, and skilful and delicate manipula- 
tion of the microscopic cultures. 

Again, the whole question of wound-rot, opened up on 
pp. 236-237, is one which demands long and thorough inves- 
tigation ; not only to clear up the many chemical problems 
involved, but also to explain the exact behaviour of sapro- 
phytic fungi and bacteria, and the part they play in the 

My duties as Editor of this work have not seemed to be 
such as to demand that I should express my own opinions on 
the subjects raised, and I have almost confined myself to merely 
noting the occurrence of the principal diseases in this country 
(since the original is written for German readers), and to adding 
a few explanatory sentences wherever it has seemed useful in the 
interests of the lay reader to do so. In some respects Hartig's 
book is a popular one — by which I mean it appeals to a wide 
circle of readers not professionally engaged in the study of 
this branch of science — and it has seemed advisable, therefore, 
occasionally to interpolate a short note in explanation of some 
of the more technical terms employed. Short notes are apt 
to be insufficient in such cases ; but it would so obviously have 
been out of place to overload the author's work with long 
disquisitions on the matters referred to, that I have been 
constrained to risk their being occasionally too brief 

Here and there I have ventured, however, to go a step 
further, and add a reference which may be useful, and this in 
face of my full recognition of the fact that Hartig's book is 
an exposition of his own view of his own work rather than 
that of others. 

In all cases I have been careful to place my remarks, more- 


over, in footnotes between square brackets, so that the run of 
the author's text is uninterrupted. In one case at least I have 
expressed dissent from Hartig's views, but here again the 
reader has the option of neglecting the footnote, and at any 
rate the matter is one of evidence. 


Cooper's Hill, Ifanh 1894. 




Development of the Study of Vegetable Pathology . . 

The Causes of Disease— Sickliness, Natural and Accidental 
Death, Debility of Old Age, Normal and- Abnormal Predis- 
position to Disease, Heredity and Disease, the Investigation 
of Disease i6 


Phanerogams — Lonicera, Triticum, Melampyrum, Rhinanthus, 
Pedicularis, Euphrasia, Lathraea, Orobanche, Monotropa, 
Viscum, Loranthus, Arceuthobium, Cuscuta 22 

Cryptogams — Pseudo-parasites — Thelephora, Lichens ; Schizomy- 
cetes — Bacterium, Bacillus ; Myxomycetes — Plasmodiophora, 
Schinzia 35 

Fungi — Their Structure and Biology, Mycelium, Hypha, Germ- 
tube, Sclerotia, Sporophore, Spore-production, Sexual and 
A-sexual Generations, Vital Conditions, Nutritive Adaptation, 
Parasites, Saprophytes, Mode of Infection and Distribution, 
Epiphytes, Endophytes, Action on the Tissues of the Host- 
plant, Prophylactic and Therapeutic Measures, Cfassification . 40 

Peronosporeae — Phytophthora, Peronospora, Pythium, Cystopus . 57 

Ustilagineae— Tilletia, Ustilago, Urocystis 66 

Ascomycetes— Erysiphete — Erysiphe, Oidium ; Tuberaceee — Ela- 
phomyces. Tuber, Mycorhiza ; Pyrenomycetes — Trichosphteria, 
Herpotrichia, Rosellinia, Dematophora, Cucurbitaria, Sphas- 
rella, Stigmatea, Gnonionia, Nectria, Polystigma, Claviceps, 
Aglaospora, Plowrightia, Physalospora, Coniothyrium, Gloe- 
sporium, Didymosphaeria ; Discomycetes — Rhytisma, Hys- 
terium, Peziza, Rhizina, Sclerotinia, Botrytis ; Gymnoascece — 
Exoascus 69- 

Imperfectly known Ascomycetes— Cercospora, Pestalozzia, 
Phoma, Gloesporium, Septogloeum, Septoria, A new Parasite 
of Seedlings, Valsa 135 

Basidiomycetes — Uredineee — Puccinia, Phragmidium, Gymno- 
sporangium, Melampsora, Coleosporium, Cronartium, Peri- 
dermium, Chrysomyxa, Isolated /Ecidium-forms 153 

Hymenomycetes— Exobasidium, Trametes, Polyporus, Hydnum, 
Thelephora, Stereum, Agaricus, Fungi destructive to structural 
timber, Dry-rot, Merulius 184. 



Healing and Production of New Tissues— Epidermis, Periderm, 
Phellogen, Bark, Cortex, Phelloderm, " Wound Cork," Tyloses 
Intermediary Tissue, Occlusion of Wounds, "Wound Wood,' 
Cinchona Culture, Effects of Pressure, Resin, Wound-rot 
Treatment of Wounds, Prev^entitious and Adventitious Buds 
Spheroblasts, Dwarf Shoots, Dormant Eyes, Root Suckers . . 225 

The various kinds of Wounds — Barking by Game, Mice 
Cattle, &c.. Wounds due to Crushing and Collection of Resin 
Ring Wounds, Pruning, Shortening Branches, Removing 
double Leaders, Coppicing, Injuries to Roots, Propagation by 
Cuttings and Grafts, Defoliation by Insects 241 


Soil in relation to Water and Plant Food— Stag-headed or top- 
dry condition. Premature Ripening of Cereals, Rupturing of 
the Cortex of Trees 270 

Circulation of Air in Soil— Root-rot 275 

Plant Poisons — Sea-water, Contaminated Water, Carbonic Acid 

Gas, Coal Gas 279 


Frost, Bark-scorching, Sun-cracks, Deficiency of Light, Hail, 

Snow, Wind, Fire, Smoke, Lightning 282 

Classified List of Diseases 305 

INDEX 317 




During the present century, and especially during the last 
few decades, the forests of Germany have been threatened with 
dangers of a magnitude formerly unknown. These have been 
occasioned by the gradual relinquishment of natural regener- 
ation, and by the substitution of pure even-aged woods for 
woods consisting of trees of different species and of various 
ages, but most of all by the displacement of broad-leaved trees 
by pure coniferous woods. It is especially noticeable that 
enemies from the animal and vegetable kingdoms find favourable 
conditions for rapid development in our modern forests, so 
that the complaints of increasing devastation of woods appear 
to be by no means unfounded. The foresters of the last cen- 
tury had already made themselves familiar with a large number 
of the enemies and diseases of trees, as is proved by the 
appearance in 1795^ of a work which probably contains the 
first compilation of the observations on plant-diseases scattered 
throughout the older literature. We may assume from this that a 
large number of diseases which have only been properly explained 
during the last few years, e.g. the damping off of seedling 
beeches, the resinous degeneration of pine-tops, the red-rot of 
the spruce, &c., were known to foresters more than a hundred 
years ago, though of course the explanation of the causes was 

1 Schreger, Erfahrimgs7ndssige Awweisung zur richtige7i Ke7intniss der 
Kra7ikhette7t der Wald- iind Garte7ibdit77ie. &c. Leipzig, 1795. 518 pages. 



bound to be defective in accordance with the position of 
botanical science at that time. 

Some fifty years ago a number of able investigators, of whom 
only Saxesen, Th. Hartig, and Ratzeburg need be named here, 
applied themselves to the study of insects. The life-history of 
forest insects, their harmfulness or usefulness, soon became the 
favourite study of many practical foresters, and in a few decades 
the joint efforts of numerous workers were rewarded by the 
elevation of Forest Entomology to the position of a much- 
appreciated subject of scientific instruction, which has become 
the common property of all educated foresters. 

The case was otherwise with those plant-diseases which 
cannot be ascribed to the injuries of animals. Their investi- 
gation was delayed until quite recently ; for it was only after 
botanical science, by the aid of its chief instrument, the 
microscope, had obtained a clear insight into the normal struc- 
ture and vital phenomena of plants, and especially after the 
study of fungi had been prosecuted in the last few decades by a 
series of distinguished investigators, that the examination of the 
phenomena of disease in the life of plants could be undertaken 
with a prospect of success. 

During the period from 1833 to 1841 three text-books of 
plant-diseases did indeed appear — namely, those of Fr. Unger,^ 
Wiegmann,- and Meyen "^ — which bear witness that in attempt- 
ing to explain the phenomena of disease in plants the progress 
already made in the knowledge of the structure and life of 
plants was not left out of account ; but the erroneous views as to 
the nature of fungi, and the absolute ignorance of the history 
of their development which prevailed, impeded progress towards 
a clear understanding of the processes of disease. Independent 
investigation was especially interfered with by the mistaken 
attempt to apply to the study of the diseases of plants the 
scientific results which J. von Liebig in particular had obtained 
in the department of agricultural chemistry. After it had 

^ Fr. Unger, Die Exantheine der Pflanze7i und eiiiige jnit diesen verwa7tdte 
Krankheiten der Gewdchse. Vienna, 1833. 

2 Wiegmann, Die Krankheiten und krankhaften Missbildungen der 
Gewdchse. Brunswick, 1839. 

3 Meyen, PJianzenpathologie. Lehre von dem krankhaften Leben und 
Bilden der PJlanzen. Berlin, 1841. 


been recognized how great is the importance for the welfare of 
plants of the quantity and condition of the mineral matter in 
the soil, and how an irrational treatment of the soil, such as 
scourging tJie ground, in sylviculture, agriculture, and horticulture 
can and must lead to exhaustion of one or other of the nutritive 
ingredients, which betrays itself in the stunted growth of the 
crop, it was supposed to be permissible, though unsupported by 
any exact investigations, to proceed a step further, and to regard 
acute diseases of crops, so long as they could not be ascribed 
to external causes, as the results of the want of one or other of 
the nutritive substances of the soil. The fact that unhealthy 
symptoms make their appearance quite as often on very fertile 
soils as on poor ones led to the assumption that a superfluity 
of nourishment may also be the means of causing diseases in 

The works of De Bary^ and Tulasne- first opened the way 
for the investigation of plant-diseases ; and with the appearance 
of these a new period began, for from that time onwards very 
great attention has been devoted to the life-history and action 
of parasitic fungi. The view hitherto held that all fungoid 
growths appear only as the result of previously existing pro- 
cesses of disease, or as indications of the incipient death of the 
part of the plant which is attacked, was shown to be erroneous. 

Investigation was now directed chiefly to the diseases of 
farm and garden crops. Amongst others Jul. Kiihn^ especially 
enriched science by a series of most valuable investigations. 
Further research gained a surer basis with the appearance of 
de Bary's* Morphology and Physiology of the Fungi. 

So far the attention of investigators had been almost entirely 
directed to agricultural crops, a circumstance which is sufficiently 
explained by the fact that but few scientific botanists had the 
opportunity presented to them of carrying their researches into 

^ De Bar}', Untersiechungen iiber die Brandpilse tend die durch sie veran- 
lassten Krankheiten der Pflansen viit Riicksic/it auf das Getreide und andere 
NdhrpflcDizen. Berlin, 1853. 

- Tulasne, Selectafnngorum carpologia. Paris, 186 1. 

^ Julius Kuhn, Die Kranlcheiten der Ciiltia-gewdchse., Hire Ursac/ie?i tend 
VerJiiitung. Berlin, 1858. 

■* De Bary, Morphologic und Physiologie der Pi/se, (S:c. Leipzig, 1866, 
and Vergleichende MorpJiologie und Biologic der Pilze. Leipzig, 1 884. 

1! 2 


the forest and of giving their attention to the diseases of trees. 
The credit of having first stimulated interest in this direction 
undoubtedly belongs to M. Willkomm.i Hallier's attempt to 
collate the scattered materials in the form of a text-book^ was 
subsequently repeated with happier results by P. Sorauer^ and 
Frank/ whose handbooks are useful compilations, in which the 
matter diffused through numerous periodicals and works is 
collected and systematically arranged. My own investigations 
have been published partly in periodicals and partly as 
independent works.'^ 


In the present state of science it is scarcely possible to draw 
a sharp line of distinction between those conditions of the 
plant known, on the one hand, as healthy, and, on the other, as 
diseased. The development of any plant depends upon a series 
of external factors of nutrition, and these, such as light, heat, 
the kind and proportion of the nutritive materials, and of the 
water and oxygen contained in the soil, of the carbonic acid 
present in the atmosphere, &c., are available for the plant in 
very different quantities. When all these external factors 
influence the development of the plant in the most favourable 
manner, it is vigorously nourished and flourishes well. But 
probably the case is never realised when all these factors of 
life act simultaneously and concurrently in the most favour- 
able manner possible : on the contrary, one or more is sure 
to be deficient or superabundant, and this causes interference 
to a greater or less extent with the development of the 
plant. We cannot as yet say, however, that such plants are 

' M. Willkomm, Die Mikroscopischen Fei?ide des Waldes. Dresden, 1866, 

- E. Hallier, Phytopathologie. Die KrankheiteJi der Cidtu7-geiucichse. 
Leipzig, 1868. 

^ P. Sorauer, Handbuch der Pfa7izcnkrcmkJieitc?i. Berlin, 1S74. 2nd 
Edition, 1886. 

* B. Frank, Die Kra)ikheitcn der Pflanzeji. Breslau, 1880. 

* R. Hartig, Wichtige Krank/ieiten der Waldbdwne. Berlin, 1874. Die 
Zcrsetzungserscheinungen des Holzes der Nadelholzbdwne und der Eiche. 
Berlin, 1878. U?itersuchunge?i aus dcm forstbotanischen Institut zu 
Micnchen. I. Berlin, 1880. III. Berlin, 1883. Die echte Hausschwamtn, 
Mcriilius lacryvtans. Berlin, 1885. 


unhealth)- ; it is only when the Hfe-processes have sunk to very 
small proportions that we speak of a plant as " sickly." 

Such sickly plants recover, as a rule, when the deficiency of 
light, heat, nutriment, or whatever the cause of the sickliness 
may be, is removed. It is the province of physiology to discover 
the conditions under which plants thrive best. I do not regard 
the investigation of the phenomena of mere sickliness as the task 
of pathology. It is only when the sickly condition leads to the 
death of some part of the plant that we may speak of actual 
disease. Suppose, for instance, that the soil of a wood has 
suffered through removal of litter, a diminution of growth will 
result, which, however, is not as yet disease ; but if a moribund 
condition of the tops of the trees sets in, we are confronted with 
the disease known as " top-drying " or " top-drought." This 
example shows how gradually the condition of sickliness merges 
into that of disease, and how it is only the partial death of the 
plant that can be regarded as giving external indication of the 

It is quite as difficult to draw the boundary line between healthy 
and diseased, and between normal and abnormal, in the case of 
those phenomena which we are accustomed to designate as 
monstrosities. In the nature of organisms there is a tendency 
towards variation both morphologically and physiologically, and 
it is upon this, in fact, that progressive evolution in the organic 
world depends. 

Variation is, therefore, a normal phenomenon, and depends on 
causes which are probably almost always operative in the earliest 
stages of the life of the organism before, during, and immediately 
after the fertilization of the oosphere. 

It is impossible to establish a strict line of demarcation between 
normal variation and malformation ; and thus all the phenomena 
connected with the latter, which we are not in a position to 
explain, have been separated and grouped together to form the 
special study of teratology apart from pathology. 

In this text-book therefore we shall confine ourselves essen- 
tially to describing and explaining those phenomena which bring 
about the premature death of the plant, or of any part of it, how- 
ever small. 

This limitation leads us to the answer to the question whether 


plants all die a natural death, or whether they, at least in part, 
succumb to external influences — that is to say, are subject only 
to accidental death. 

Experience teaches that, at any rate among the more highly 
developed plants, each individual dies sooner or later, but that in 
the case of perennial plants, particularly trees and shrubs, the 
cause of death is always to be found in unfavourable external 
influences. In the case of the more lowly organisms, which only 
multiply by division and as yet exhibit no sexual reproduction, 
one can scarcely speak of a natural death, because each part is 
as old as the parent organism by the division, &c., of which it was 
formed. Were a natural limit set to the life of a certain species 
of plant which can only multiply by dividing, the result would 
be that when this limit was reached every part, and therefore 
also the offspring which had originated by division, would perish. 
It is known, however, that this state of things does not exist. 
In the case of those plants which are also reproduced by sexual 
processes many different conditions are met with. In the case 
of annual plants the vegetative part dies each year, and only the 
embryos originating from the fertilized oospheres remain alive. 
When, from these, plants capable of bearing seeds have deve- 
loped, all that is preserved of them, in their turn, is the forma- 
tive product arising from the sexual cells. Thus the vegetative 
part of each plant dies owing to internal causes, though these, 
in part, depend simply upon exhaustion consequent on the 
formation of seed. We see then that natural death of the vege- 
tative organs of the plant occurs from internal causes, whereas 
the sexual cells only die if they have not been fertilized, or if, 
owing to external causes, the product of fertilization has not 
given rise to a new plant. Upon the unlimited duration of the 
life of this part of the plant — that is to say, of those sexual cells 
which do not fall victims to accidental death — depends indeed 
continuity in the organic world, in other words, the development 
and preservation of the vegetable and animal kingdoms. 

In the case of perennial plants it is only certain parts that 
succumb to natural death each year. Amongst herbaceous 
plants, for instance, it is the parts above ground which thus die 
off: in the case of deciduous trees and shrubs, it is the outer 
cortical tissues, the leaves, &c. 


The plant-individual proper, however, only dies in consequence 
of unfavourable external influences. As a matter of fact, every 
tree is rejuvenated each year by the cambium forming new 
tissues at its periphery, and by new shoots and buds. It is a 
matter of experience that the duration of the life of all trees is 
limited, but it is not proved whether this is to be ascribed to in- 
ternal causes, or is the result of the innumerable influences which 
act more or less prejudicially on the plant from without. The re- 
duction and final cessation of the growth in height of a tree, after 
attaining a certain maximum, must be ascribed to interference 
with the factors of nutrition, and, in all probability, especially 
to the fact that the forces which conduct the water and nutritive 
materials to the highest bud of the tree are limited in their 
action, and that sooner or later, depending on the specific and 
individual nature of the plant, these no longer suffice to provide 
for the continuance of growth in height. If we cut a slip from 
an old tree, it will pursue the same cycle of development as the 
parent tree, thereby proving that by vegetative multiplication 
the life of a plant may be indefinitely prolonged. Hitherto no 
phenomenon has been discovered from which one may conclude 
with certainty that internal natural causes of death are peculiar 
to all, or even to any perennial plants. In this connection the 
question is at once suggested whether " the feebleness of old 
age " is a factor which must be regarded at all in considering the 
diseases of plants. In discussing how diseases arise we shall 
show that old age, quite as well as youth, may predispose a 
plant to some disease or other. In itself, however, the feebleness 
of old age is not a natural condition attributable to internal 
causes, but is a .state induced by external influences. The older 
a tree is, so much the more numerous are the dangers through 
which it has had to pass, and so much the greater is the 
number of its injuries and wounds through which parasites 
and saprophytes can find an entrance into its interior. Again, 
the older a tree is, the narrower are its annual rings, and 
with so much the more difficulty and tardiness does it suc- 
ceed in occluding a wound. Finally, the older a tree is, 
the more sluggish are its nutritive processes, because, on the 
one hand, the soil in which the roots are fixed has become 
denser, thereby impeding the entrance of air, and, on the other 


hand, one or other of the nutritive materials may be partiall}' 

With the reduction in the transference of nutrient matters to 
the crown of the tree, the latter becomes stunted and partly dies, 
and this is followed by diseases which finally kill it altogether. 

There are, however, always demonstrable external influences 
at work in the matter, so that the question whether the debility 
of old age is in itself a natural condition manifesting itself, for 
instance, in the enervation of the organization of a cambium 
cell, or in the separation of a bud from a tree, must in the mean- 
time be answered in the negative. Thus, when we speak of the 
natural duration of life of a plant-species, we are to understand 
the period of time during which a plant is able to liv^e without 
succumbing to the unfavourable external agencies in the soil 
and the climate, or to the varied attacks of parasitic and 
saprophytic organisms. 

The above considerations lead us to the natural classification 
of the different kinds of disease which we shall examine in the 
following pages, according to the external influences which 
induce them. 

1. Diseases induced b)^ Phanerogams. 

2. Diseases induced by Cr}'ptogams. 

3. Wounds. 

4. Diseases due to unfavourable conditions of the soil. 

5. Diseases due to unfavourable atmospheric conditions. 

In the case of most diseases it is demonstrable that the 
individuals of a given species of plant, which are subjected to 
certain prejudicial influences, do not all succumb to these 
influences to an equal extent, but that certain individuals or 
varieties prove perfectly or almost perfectly resistant, while 
others soon become diseased or die. These observ^ations show 
that it is not the environment alone which determines the origin 
of a disease, but that, on the contrary, a plant contracts disease 
onl}' when subjected to definite pre-existing conditions ; that a 
predisposition or tendency to disease must exist, and that there- 
fore, to a certain extent, the origin of a disease is determined by 
the co-operation of two factors. The first factor is the external 
cause of the disease, and this is, as a rule, easy of demonstration. 


The second factor, however, has its inception in a peculiar 
condition of the organization of the plant, which is either present 
only at certain times, or is only peculiar to and innate in certain 
individuals, or, finally, has been acquired under the influence 
of definite external conditions. All these peculiarities in the 
organization of the plant may be quite normal in their nature — 
that is to say, the organism as such appears to be perfectly 
healthy — in which case the predisposition is said to be "normal." 
On the other hand, however, the predisposition to disease may 
be " abnormal," as is the case when the plant is only predisposed 
to one disease because it is already suffering from another. 
Abnormal or disease-inducing predisposition may arise, for 
example, in the neighbourhood of a wound through which alone 
some particular parasite could gain entrance to the plant. The 
entire group of infectious wound-diseases may be placed in this 

Under normal predisposition, therefore, zve are to tinderstand 
every condition, even if only temporary, in the anatomical struc- 
ture, in the chemical constitution, or ivt, the vital functions of an 
organism, which, though not in itself prejudicial to the individual, 
induces a disease zvhen a second, and that an external, factor co- 
operates in addition, even though the latter is in itself innocuous 
to the plant. 

In addition to these cases of normal and abnormal disposition 
residing in the organism itself, we may also speak of a pre- 
disposition to disease which is due to the locality. 

There are a great number of fungi which can only attack a 
certain species of tree when plants of another species occur in 
the vicinity on which the particular fungus, at certain seasons 
of the year, may complete its development. Localities in 
which m.any aspens grow impart to the pines a predisposition 
for the disease known as "Pine-twist" (caused by Melanipsora 
TremulcB pinitorquuni). Rhododendrons abounding in a district 
make the spruces liable to " Leaf-blister " (caused by Clny- 
somyxa Rhododendri), while barberr\' bushes are associated 
with the " rust " of wheat. The mere existence of uninter- 
rupted woods, composed of a single species of tree, may give 
rise to dangers leading to extensive epidemics. Pure larch 
woods away from mountainous regions almost always succumb 


to canker, whereas larches mixed with other trees may remain 
unaffected. The cHmatic conditions pecuHar to a given district 
may render-it specially liable to outbreaks of certain diseases. 
Thus in Alpine districts proximity to lakes and narrow valleys 
specially predisposes to certain fungoid diseases, because the 
moist air of such places favours the fructification of fungi in 
a high degree. In the forest one meets with certain localities, 
so-called " frost-beds," which favour the injurious action of 
frost. The character of the soil may predispose to definite 
diseases, in that, for instance, it specially favours the growth of 
underground parasitic fungi, or the conditions may induce the 
appearance of "root-rot." In very many cases one can say 
forthwith of certain localities that they predispose to definite 
diseases, and the latter must occur when some factor or other 
of the environment is present, although in other localities the 
same factor may be harmless to the vegetable world. Of course 
this predisposition which is linked to the locality only forms 
a part of the multifarious circumstances favourable to the 
occurrence and spread of diseases that are to be ascribed to 
the environment of the plant, and it must not be confounded 
with the idea of a predisposition to disease in the narrower 

In the first place, the normal predisposition of plants may 
consist in phases of development which naturally exist for a time 
in every plant. To this class belongs the period of youth of the 
plant, and the young condition of its new shoots, leaves, 
and roots. These are at first covered only by a delicate 
epidermis, which is but slightly if at all cuticularized, and 
which can offer no resistance to the attacks of parasitic 
fungi ; whereas later on in life, when a cuticle has been formed 
on the outer cell-walls, and when periderm and bark have been 
formed on the axial organs, the predisposition for many forms 
of disease disappears. 

On the other hand, later periods of life may also induce 
a predisposition to certain diseases. Young conifers which 
possess resin-canals are almost perfectly protected from 
infection by wood-fungi, at least in so far as these find an 
entrance only through wounds caused by the removal of branches, 
because each fresh wound is at once covered by a protecting 


substance due to the exudation of turpentine. It is not until 
after the development of duramen, which no longer conducts 
water, that a predisposition for wood-diseases sets in, because 
now, when a branch is broken off, the inner wood no longer 
protects itself against attack by pouring out turpentine, for it 
is only in the watery alburnum that turpentine and resin are 
forcibly pressed out of the resin-canals. In the case of trees 
advancing age is also, as a rule, accompanied by a diminution 
in the breadth of the successive annual rings, and the result 
of this is that wounds are not so quickly occluded as when 
they occur on young vigorously growing trees. It is easy to 
perceive that, as a consequence, the prejudicial results of injuries 
are increased as age advances. In this sense alone can we 
speak of the feebleness of old age, and increasing susceptibility 
to external dangers in consequence. 

The condition of vegetation in regard to the season of the year 
has great influence on the power of the plant to resist dangers. 
It is well known what low temperatures a plant can stand during 
the period of winter rest, whereas in spring, after the beginning 
of vegetative activity, and before it ceases in autumn, it is 
killed by a few degrees of frost. 

The capacity of resisting the attacks of parasitic fungi on 
the part of the tissues also differs much according to the 
season of the year. Between the living cell of the host-plant 
and the cell of the fungus-parasite there is a struggle, in which 
(in the case of many parasites infesting the tissues of the cortex 
or cambium) the latter can only kill the former if this is in 
the condition of vegetative rest — that is to say, not actively 
growing, &c. If processes of metabolism are energetically at 
work in the cellular tissue of the host-plant itself, it is then 
enabled to stave off the attacks of the fungus. The action 
of the latter on the cellular tissue of the host — depending 
as it does on the secretion of an enzyme * — is only prejudicial 
when this tissue is defenceless, as it were, owing to the condition 
of inactivity in which it is found. These cortex-fungi grow 
only from autumn till spring, and their further development 

* [Enzymes are a peculiar class of bodies, often called unorganised or 
soluble ferments, capable of producing powerful molecular changes in 
organic substances in presence of water. — Ed.] 


is checked with the beginning of vegetative activity in the 
host-plant. A similar condition of things is met with in those 
fungi which at all seasons luxuriate in the wood of trees, and 
even kill its living cells, but which are incapable of penetrating 
the living tissues of the cortex till these have succumbed to 
drought due to the death of the wood, when they may be 
easily occupied. The tissues of the wood and those of the 
cortex appear to differ in their power of resisting parasites. 

The amount of water in plants, determined by the weather, 
also influences the development of endo-parasites. During 
periods of much rain, when the plant-tissues contain more 
water than during periods of drought, many perennial fungi 
flourish with special vigour inside the plant. This is particularly 
evident in the case of Melanipsora TroniilcB pinitoi-qiiiun and 
Rosellinia qnercina. 

In contradistinction to those phenomena of predisposition 
which have been already discussed, and which to a certain 
extent only appear periodically, there is a second category of 
peculiarities which are innate, as it were, only in certain 
individuals or varieties, which are thereby specially predisposed 
to certain diseases. Variation in the vegetable kingdom ma)' 
find expression in morphological, chemical, and physiological 
peculiarities, and in each of these directions forms may tend to 
occur which are more or less susceptible to one disease or other. 
As regards the morphological aspect, it need only be called to 
mind that there are varieties of potato which possess a very 
delicate skin ; others, a thick periderm ; and it is easy to 
explain why the former are far less secure against the attack 
of the fungus that causes potato disease than those with a 
thick skin. 

Amongst the Douglas firs there is a bluish glaucous variety 
whose leaves, owing to an abundant waxy covering, are much 
better protected against atmospheric drought than the pure 
green form. That the latter possesses a predisposition to perish 
from drought in a continental climate is to be expected from 
the fact that it is natural!}' confined to the west-coast region of 
North America. 

That individual differences with respect to the chemical com- 
position, and especially to the amount of water, occur in plants 


is undoubted, and it may be safeh' assumed that these differences 
also involve differences in behaviour towards prejudicial external 
influences. At present, however, we know very little in this 
connection, and we can in the meantime only conjecture that 
the explanation of the individual differences in the behaviour of 
plants towards frost, drought, and even towards the attacks of 
fungi will partly be found in such chemical differences. 

Even more striking are the cases where differences in the 
physiological behaviour of plants serve as disease-inducing con- 
ditions. It is well known how certain trees of the same wood 
awake from their winter rest and become green at different 
times, although in other respects they are perfectly similar. In 
a young spruce plantation differences of two or even three weeks 
may be easily perceived in the opening of the buds of different 
individuals, and this must be accounted for, in most part, by 
differences in the heat-requirements of the plants. It is evident 
that early unfolding of the leaves implies a disposition for injury 
by late frosts, but it may also become the chief stimulus to the 
development of fungoid disease. If, for instance, the spruce-leaf- 
rust {Chrysomyxd) is, in the spring, at the stage when its spores 
are being shed, all those spruces whose buds have not begun 
to elongate into shoots w411 remain entirely unaffected by the 
fungus, which is only able to force its way into the delicate 
leaves of young shoots. A disposition for this disease, therefore, 
attaches to the individuals which begin to grow earh'. In 
other years it may happen that those individuals which first 
begin to grow are so far advanced in development when 
Chrysomyxa sheds its spores that the leaves are already too old 
to be susceptible to infection. In this case it is perhaps just 
the late varieties that contract the disease. 

The observation that amongst the individuals of a plant 
species there are always some whose requirements as regards 
heat are less or more than those of others, and that these are 
therefore disposed to suffer from cold to a greater or less extent, 
and that, further, demands on the moisture of the air and other 
factors of growth vary with the individual, has probably led to 
importance being attached to the place of origin of the seeds 
which we employ in cultural experiments with exotic species of 
plants. We endeavour to obtain seeds from districts where, in 


the course of time, varieties have spontaneously arisen whose 
power of resisting frost, or atmospheric drought, as the case may 
be, has become enhanced. 

A further group of disease-inducing conditions embraces all 
these peculiarities which have only been acquired in the course 
of the development of the plant, and which may lead to a disease 
if certain external influences be present. If plants are reared in a 
moist atmosphere, e.g. in a greenhouse, the epidermal system 
develops in response to the moist air which surrounds it, so that 
it is only slightly cuticularized. If such plants are placed in a 
dry atmosphere — for instance, in the air of a heated room — they 
become sickly, because the transpiration of the leaves is unduly 

Trees, especially those with smooth periderm, that are reared 
in a very dense wood, and then suddenly isolated in later life, 
suffer from scorching of the cortex. Such trees possess a pre- 
disposition for scorching which is absent in the case of those 
plants of the same species which have been grown, from youth 
upwards, in an open or light wood. The disposition to disease 
in this case consists in the fact that the external covering is 
less strongly developed. Plants grown in the shade also prove 
to be unduly susceptible to the direct action of the sun, in 
that the chlorophyll in the cells of the upper layers of their 
leaves becomes destroyed. Oaks grown in a close beech wood, 
and consequently with small crowns, incur a predisposition for 
top-drought when they are isolated, whereas, under similar 
circumstances, trees with full crowns do not suffer from this 

During the first few years after being transplanted, many 
trees show a predisposition to be easily " frosted," which is again 
lost with the development of a strong root-system. On shallow 
soils evergreens, and especially conifers, are far more susceptible 
to injury from coal-smoke than those on deep soils, for the 
reason that their root-system, being characterised by superficial 
development, is unable to take up water in winter. Desiccation 
of the leaves in consequence of action of the sulphurous acid 
takes place in their case much more easily than in that of trees 
which are able to take up water from greater depths, and this 
even in winter. 


All the disease-inducing conditions which have been dis- 
cussed may be designated as normal, because the peculiarities 
noted are in themselves quite in accordance with the nature of 
the plant-organism, and only become prejudicial when some 
other external circumstance co-operates, and which is termed 
the cause of the disease. 

There still remain to be noticed numerous abnormal disease- 
inducing conditions which depend on an unsound state of the 
plant. To these belong all those wounds in whose train some 
disease or other of the interior of the plant may follow. 

When a tree is pruned it thereby incurs an abnormal predis- 
position for a series of wound-diseases, infectious or otherwise, 
which can be got rid of by the application of timely and appro- 
priate — that is to say, antiseptic — dressings. Injury to a root, 
e.g. the severance of a rootlet, is in itself damage, but when this 
leads to decay spreading from it into the stem, we designate 
such an injury as an abnormal disposition to disease. 

Insects of various kinds live in the cortex of sound trees, 
which they injure, and thus open doors, as it were, to the entrance 
of parasitic fungi into the interior, so that the trees are ultimately 

A hailstone strikes the cortex of a tree and injures it. This 
creates an abnormal condition, which may lead to an infectious 
disease should certain fungi settle on the cortex. 

When trees or shrubs are transplanted in any year, and their 
development is so much retarded by the operation that the new 
shoots have not completed their development when frost appears 
— that is to say, when lignification has not been completed — 
they possess an abnormal disposition to injury from frost. 
Such plants may survive in mild winters, but if intense cold sets 
in they may die off completely. 

From what has been said it will be clear how endless are 
the phenomena which dispose to disease, and also how only one 
group of these, "the inherent tendencies," possess the character 
of inheritability. The phases of natural development, which 
were first discussed, and which are passed through by ever)- 
plant, may be left out of account in connection with the ques- 
tion of inheritability. Neither acquired predisposing causes nor 
those due to an unhealthy state can, however, be transmitted 


from parents to descendants ; at least nothing is known so far 
that indicates such an inheritance. This holds good not only 
for the causes but also for the diseases themselves. 

A transmission, by inheritance, of diseases to descendants is 
unknown in the vegetable kingdom. One may without hesita- 
tion make use of the seeds of plants suffering from any con- 
ceivable disease for the propagation of new plants. In par- 
ticular one may without scruple collect the seed of such trees as 
are dwarfed owing to poverty of the soil. Indeed, as a matter 
of fact, this is done, for instance, in the case of the Scotch 
pine, the cones of which are gathered by preference from those 
trees whose proportions are so diminutive, owing to their grow- 
ing on barren moors, that the collection of the cones may be 
accomplished with ease without climbing the trees. It is only 
when a question of individual properties consisting in dwarfed 
habit of growth, spiral stems, or other undesirable peculiarities 
that are innate in the plant is involved, that the law of in- 
heritance comes into consideration, and then propagators of 
plants have to exercise the greatest care. 


Reference will here shortly be made to the methods of investi- 
gation which we have to follow when we wish to determine the 
causes of diseases in plants. 

In the case of diseases of men or animals the difficulties of 
diagnosis are much increased by the fact that in the great 
majority of cases the disease of a single organ or part of the 
body is followed by secondary phenomena which impede the 
discovery of the proper seat of disease. In the bodies of plants, 
where the nervous system is absent, a disease as a rule remains 
localized, at least at first. The division of labour is not yet so 
far differentiated as in the bodies of the more highly organised 
animals, where disease of any organ, often even a small one, 
involves the whole body sympathetically. A large part of the 
body of a plant may be diseased and even killed without the 
plant being necessarily perceptibly injured in its general health. 
If we have succeeded in observing the disease in its first stage, 


the further investigation offers comparatively Httlc difficulty. It 
is more difficult, as a rule, to determine the true cause of disease 
and death in the case of plants already dead, although the skilled 
plant-pathologist will seldom fail to recognize with certainty the 
true character of a disease. 

If we are dealing with injuries caused by animals or plants, 
we shall discover and recognize them, or at least their traces, 
with most certainty in the preliminary stages of the disease. In 
very many cases it is not sufficient, where we are dealing with 
injuries due to animals, including insects, that we catch the 
creature at work and seek to observe it and its mode of life in 
nature, as has hitherto generally been done ; but, and particularly 
in the case of insect-injuries, w^e must determine whether the 
injured plants did not already possess some predisposition to 
disease before they were attacked by the insects, &c. Especially 
does this hold good for the great family of the bark beetles, 
which often only appear in the train of other prejudicial agencies, 
and especially of injuries caused by parasitic fungi. In the case 
of parasitic plants, again, it is not to be concluded forthwith from 
the presence of a fungus in the dead tissues that death has been 
caused by that fungus. True, where we find the mycelia of fungi 
vegetating in the apparently Jtualtered living tissues of a plant, 
there is practically no room for doubt that we have to deal with 
a parasite ; but even in the latter case the attempt must next 
be made, by means of suitable infection-experiments, to induce 
arbitrarily, and in a somewhat artificial manner, the disease that 
we are seeking to investigate. 

If spores or gonidia of the suspected fungus are to be had, we 
make use of these in carrying out the investigation, after having 
first proved that they are capable of germinating. Should no 
material capable of germinating be at our disposal, we must, if 
possible, undertake artificial cultures in a damp chamber, and 
await the ripening of spores, or even the production of sporo- 
phores.* According to the character of the disease, infection is 
secured by scattering the spores on the leaves, or by placing 
them in a wound artificially made in the host-plant. In the 

* [Spores may be of several kinds, and the term is used as a general one. 
Gonidia are a-sexual spores. Sporophore is a general term to denote any of 
the various kinds of spore-bearing structures met with among Fungi.— -Ed.] 


case of diseases of the cortex it is sufficient to make a fine inci- 
sion with the point of a scalpel, to which a drop of water, with 
spores suspended in it, is attached ; in the case of diseases of 
the wood the latter must be pierced by the wound, which is 
then allowed to absorb the drop of water with its contained 

In dealing with diseases of the cortex or wood, infection by 
means of mycelium* is the much surer course. Having removed a 
small piece of the cortex from that partof a diseased tree where 
the mycelium is still young and vigorous — that is to say, from the 
boundary between the dead and living tissue, we place it on a 
spot in a healthy tree from which a piece of cortex of the same 
size and shape has been removed. We may proceed exactly as in 
budding roses, but it is generally better if the edges of the piece of 
cortex containing the mycelium are brought into intimate contact 
with the edges of the cortex surrounding the spot operated upon, 
and which, moreover, should be prepared immediately beforehand. 

Desiccation should then be prevented by applying grafting-wax 
or other dressing. If it is desired to infect the wood of the stem 
of a tree with mycelium, a small cylindrical block is removed 
(by means of Pressler's growth-borer, an instrument specially 
adapted for such work) from the boundary between the sound 
and diseased wood, because it is usually only in this region that 
the m}'celium contained in the wood is still capable of such 
vigorous growth as to be able to extend beyond the surface of the 
infecting block. With the same borer an exactly similar hole is 
then made in the sound tree selected for infection, the diseased 
cylinder being substituted for that which was withdrawn, and 
the hole closed externally with grafting-wax. 

If, finally, we have to do with parasites which vegetate under- 
ground, it suffices, as a rule, to plant a diseased specimen in 
immediate proximity to a healthy plant of the same species. In 
doing this greater success will probably be secured by bringing 
a root of the diseased individual (known to contain mycelium 
still living and capable of growth) into immediate contact with 
a root of the plant to be infected. 

* [The mycelium is the vegetative part of the fungus, and in many cases is 
more easily obtained in the requisite quantities than spores are : a rough 
equivalent for the word in English is " Spawn " (of mushrooms, &c.) — Ed.] 


It would be a mistake to attempt to answer the question 
whether a fungus is really a parasite or not after the failure of 
one or a few attempts at infection. Let us only consider how 
numerous are the factors regulating success even in the 
sowing or planting of our forest trees, with whose conditions 
of life we are to some extent familiar. Of the fungus to be 
investigated, however, we, as a rule, know almost nothing ; we 
do not know the external conditions of germination, we often 
scarcely know whether the spores are really ripe, whether they 
are in too damp or too dry a medium, whether sufficient oxygen 
is admitted to them, or whether the season of the year is the 
right one for sowing — for spores, like the seeds of forest-trees, 
require to rest for different periods after ripening, before they will 
germinate. What has already been said about the numerous 
conditions predisposing plants to disease will show sufficiently 
how, even with the best material for infection, experiments ma}- 
give only negative results. Since even trained mycologists and 
pathologists often succeed only after innumerable abortive 
experiments in making themselves acquainted with the con- 
ditions under which a plant becomes infected, it will be clear 
that it may be regarded as simply an accident when an amateur 
succeeds at all with an infection-experiment. 

When the infection has succeeded, it is not enough merely to 
follow the course of the disease through its various stages — in 
doing which, moreover, it is of the utmost importance to compare 
cases of disease met with in the forest — but it is also necessary 
to discover the external influences which restrict or further the 
development of the disease. 

This part of the investigation is the most difficult. It 

demands, very specially, the power of accurate observation ; 

apparently unimportant accessory circumstances must be noticed 

and compared ; and, above all things, excursions to the forest 

must be made as often as possible. Investigations of the 

diseases of our forest trees will seldom lead to any definite 

result unless we make careful and extensive observations and 

comparisons in the forest itself. At the same time, still less 

prospect of success will attend observations of diseases in the 

forest if they are not accompanied and supported by exact 

scientific investigation. 

r o 


If investigation shows that neither animal nor vegetable 
organisms are the primary cause of the disease, then the latter 
must be due to some influence in the inorganic environment. If 
it is suspected that unfavourable properties of the soil are 
responsible for the disease, the unhealthy tree should be removed, 
and — if possible, at the spot where it stood — a hole should be dug 
to the depth reached by the lowest roots. During the opera- 
tion attention is to be directed to the consistency of the layers 
of soil, and to the quantity of water which they contain, and 
especially to their greater or less accessibility to atmospheric 
air. In the forest a change in the amount of mineral food, so 
great as to induce disease in a previously healthy tree or planta- 
tion, would occur only under circumstances that would at once 
attract the attention of the skilled observer. For instance, top- 
drought may be a consequence of the removal of litter or the 
laying bare of the soil ; sickness or death may be occasioned 
by the presence of injurious substances derived from factories, 
or owing to flooding with sea-water, &c. Chemical investigation 
will very seldom be necessary. More frequently the cause is 
attributable to atmospheric influences, such as variations of 
temperature, moisture, or precipitations, or to lightning, noxious 
gases, &c. If it can be determined when the disease first 
appeared, the task will often be more quickly mastered by col- 
lecting information, and by ascertaining the external conditions, 
than by investigating the diseased plant, although the latter 
course will, in many cases, lead to the desired end. 

As a rule, diseases produced by animals and plants are 
characterised by their occurring first of all on single plants or 
parts of plants, and then gradually extending from these 
centres ; whereas diseases which are due to the influence of soil 
or atmosphere generally appear regularly and simultaneously 
over large areas, because such influences are seldom bounded 
in the forest by narrow limits or confined to the neighbourhood 
of single plants. 

Mistakes are most likely to occur when a disease is preceded 
by an abnormal predisposition, because this alone, and not the 
disease rendered possible by it, is apt to be kept in view. It 
frequently happens that we meet with different diseases on the 
same tree, each of which is at work independently, and when 


this is the case we should not at once stop the investigation 
when a cause of disease has been discovered. Very often we 
encounter, e.g. in the low lands of North Germany, devastated 
woods of Scotch pines, in which many trees have been killed by 
Trametes radiciperda. More exact investigation, however, often 
results in the discovery that in the same wood, in consequence 
of insufficient circulation of air in the soil, root-rotting is much 
more destructive even than the root-parasite. 

Only the most careful research, supported by thorough know- 
ledge of the forms of disease, which are so numerous and 
varied, can protect us against error. 


This is not the place to discuss the many conditions brought 
about by the struggle for existence — for space, food-materials, 
water, and light — between plants of the same or opposite species. 
Any plant may prove injurious to another if it makes the same 
or similar claims on the constituents of the soil. When two 
plants compete with each other, success does not alone depend 
upon the rapid rate of growth of any particular species on a 
given situation, but is determined largely by the rate of growth 
which characterises iiidividual plants ; and it is this which is 
mainly decisive in pure woods. It has long been known that 
superior individual growth manifests itself in the earliest stages 
of the life of a tree ; in fact, sometimes — for instance, in the 
case of the oak — it is recognizable in the size of the fruit. -^ 
It is therefore of the greatest importance not only to exer- 
cise care in the selection of the seed, but also to remove 
weak plants when transplanting in the nursery. When 
crowded, all plants must struggle with their nearest neigh- 
bours ; but I do not consider that it is the province of vegetable 
pathology to discuss these conditions : rather do I hold that 
I should closely confine myself to the consideration of those 
injuries which consist in the direct attack of one plant on 
the life and health of another. 

1 Th. Hartig established this fact experimentally in the nursery at 
Brunswick thirty years ago. 



No sharp boundary can be drawn between parasites and 
plants that are only indirectly injurious owing to their proximity, 
or to their competition for food-materials, light, and so forth.* 
One gradually passes from the latter to plants which, while 
not subsisting on the substance of others, still directly attack 
them, and induce pathological phenomena. 

Allusion may be made, for instance, to Loniccra Periclyme7iiim,\ 
whose stem, when opportunity offers, winds round young trees 
with the result that a few years later the descent of the plastic 
materials in the bast is forced to take place in a definite spiral 
course. With increasing thickness the tree is soon subjected 
to direct pressure by the twining plant, which prevents the direct 
vertical descent of the solutions of food-substances coming 
from the leaves. It frequently happens that the part of the 
stem immediately below that of the honeysuckle is deprived of 
nourishment, the result being that the cambium in that region 
gradually dies ; whereas the portion above the passively tighten- 
ing stem of the honeysuckle on the one hand exhibits very 
vigorous growth, and, on the other, experiences abnormal altera- 
tion in its younger parts owing to the spiral direction imparted 
to all the organs of the vascular bundles. 

While there is no doubt that the immediate cause of the 
movement of plastic substances in the cortex and bast is the fact 
of their being produced in one place and utilised in another, neces- 
sitating a transference from the place of origin to the place of con- 
sumption, still the assumption that the movement of the plastic 
materials in the bast takes place much more easily and quickly 
in a vertical than in a lateral direction receives support from 
the fact just mentioned, and illustrated in Fig. i, as well as 
from many other phenomena. In fact, lateral movement is 
so difficult as sometimes to induce complete cessation of the 

* [In such cases the plants not vakied are regarded as " weeds " ; but it is 
obvious that any plant may act as a weed towards others, and a little reflec- 
tion shows that it may act as a weed towards its own species if crowded, 
&c.— Ed.] t [Honeysuckle.— Ed.] 



nourishment of the band of cambium situated under the stem 
of the honeysuckle. 

Tritiaim repeiis* may also be mentioned here, the sharp-pointed 
rhizome of which has the power of piercing and growing through 

the fleshy roots of other plants which it 
may encounter in the soil. This has 
been speciall}- observed in beds of oak 
seedlings, though it may be remarked 
that the piercing of the roots has re- 
sulted in no apparent damage to the 

The passage to the true parasites — 
that is to say, to those which subsist 
entirely on the plastic materials of 
other plants — is formed by a group of 
plants in which one cannot at first per- 
ceive a parasitic existence, seeing that 
they are provided with leaves containing 
chloroph}'ll, and take water and inor- 
ganic nourishment from the soil with 
their roots. While they assimilate plastic 
materials for themselves, they also attach 
themselves to the roots of other phane- 
rogamic plants from which the\- abstract 
organic substances by means of an absorb- 
ing apparatus (Jiaiistoriiivi) on certain of 
their roots. To such plants belong the 
RhinantJiecs, a sub- family of the Scro- 
pJmlariacece. The cow-wheat {Melampy- 
nim arvense), the common }-ellow-rattle 
{RJmianthus Crista-galli), and the louse- 
wort {Pedicularis) and eye-bright {Eii- 
phrasia) furnish well-known examples 
of this kind of life. As these plants 
are parasitic onl}- on the plants of pastures and meadows, we 
cannot here spare time to examine them more closely. The genus 
LatJircca, also, which contains the common species Lat/inva 
squaviaria, the tooth-wort, has not yet been proved to be wholly 
* [" Couch grass " or '• twitch ." — Ed.] 


parasitic. Its roots arc parti}' attached to the roots of manj- 
different plants, including several trees such as beech, hornbeam, 
hazel, and alder. 

Notwithstanding traces of chloroph)il in the OrobancJiacece, 
these must undoubtedl)' be classed as true parasites, which derive 
their nourishment exclusively from the host-plants to whose roots 
they are attached. Of the numerous species, some occur so 
plentifully on cultixated plants as to cause appreciable damage 
e.g. Orobanchc raniosa on tobacco and hemp, O. lucoriivi on the 
barberry and bramble, O. Hcdcra; on ivy, O. riibens on lucerne, 
and O. minor on red clover. The parasitism of the yellow bird's- 
nest {Monotropa Hypopitys) is still doubtful, but, as its roots are 
found in contact with those of conifers and the beech, it is 
extremely probable that though most of its nourishment is got 
from the humus a certain amount is also abstracted from these 
plants. Besides Monotropa we have the Orchidacccr that are 
destitute of chlorophyll, which are all of a saprophytic nature. 

Nor can the Loranthacccv be regarded as true parasites, 
because for the most part they abstract onl}' water and inorganic 
food-materials from trees and shrubs, organic substances being 
appropriated to a very limited extent. They possess leaves 
containing chlorophyll, and behave towards their host in exactl)- 
the same way as a scion does to a stock. In fact they yield up 
a portion of the plastic substances which they themselves have 
prepared to the host-plant, which employs them for its own growth. 
Whether, however, this occurs in the case of all, or even most, 
LorantJiacecE is doubtful, but at all events such reciprocal nourish- 
ing takes place with Loraiithus eiiropcuus. The manner in which 
the different species of this family abstract water and nourishment 
by means of their roots from the plants which they inhabit varies 
extremely, especiall}- when the exotic species are taken into 

The best-known species is J^iscnni albiu/i, the common mistletoe, 
which is distributed throughout the whole of Europe, and Asia to 

^ See Solms Laubach in Pringsheim's JaluinicJiern f. tviss. Bot. VI., 
pp. 575 et seq. R. Hartig, Ziir Kennt7riss vo7t Lora7ithus europacus iiud 
Visciim album., with, a table, ZeitscJmft fiir Forst- u. Jagd-Wesen . 1876, 
pp. 321 et seq. Dr. C. v. Tubeuf, Beitrdge ziir Kenntiiiss der Baiunkrank- 
heiteii., pp. 9 — 28. Springer, Berlin, 1888. 


Japan. It is met with on almost all dicotyledons and conifers, 
but exibits a preference for certain species, e.g. silver fir, Scotch 
pine, poplars, and fruit-trees ; whereas on others, e.g. spruce, oak, 
beech, Spanish chestnut, alder, and ash, it has been met with 
either very seldom or not at all.^ With regard to the appearance 
of this familiar plant, it need only be mentioned that narrow and 
broad-leaved varieties occur on different species of trees. The 
mistletoe is distributed by thrushes (especially Tiwdiis viscivorus), 
which feed upon the berries and carry them off. The birds dis- 
engage the sticky seeds from their beaks by rubbing them against 
the branches on which they perch, and to which the seeds there- 
by become attached. The seeds, which germinate in spring, first 
of all develop a kind of sucker, from ^vhose centre a fine root 
appears which pierces the tissues of the cortex. This main root 
penetrates to the wood of the branch or stem, which, however, 
it is too delicate to enter. This then finishes its apical growth 
in length, but, on the other hand, owing to the presence of 
meristematic tissue behind the apex (such tissue being situated 
in the region of the cambium of the host-plant), it is enabled to 
elongate at the same rate as the branch increases in thickness, 
by the formation of a ring of wood and of bast (" intermediary 
growth in length"). Owing to the fact that the wood-ring en- 
velops the apex of the root of the mistletoe, the latter appears 
to bore deeper into the wood each }'ear, but this is really due to 
its being embraced by the stem as it grows in thickness. The 
growth in length of this root, as of all the " sinkers " that after- 
wards originate in the roots that are met with in the cortex, 
very closely resembles the growth in length of a medullary ray 
possessing cambium of its own in the cambium mantle that 
covers the whole stem, and is thus enabled annually to elongate 
both towards the wood and towards the bast. Several lateral roots 
next appear on that part of the radicle which is situated in the 
cortex, and these proceed to grow both upwards and downwards 
in the branch. These " Rhizoids " or " Cortex-roots " push their 
pencil-like apices along the }'oung soft bast, without however 
coming into contact with or altering the cambium zone. The 
organs of the soft bast are dissolved in front of the point, and it 

1 Nobbe, " Ueber die Mistel, ihre Verbreitimg, Standorte., tuid forstUche 
Eedeiiiiaig" Thorander forstliches Jahrbuch., 1884. 


ma)- be taken for granted that the products of sohition are absorbed 
by the cortex-roots and used for their own growth. From investi- 
gations made on the Scotch pine and the silver fir, the annual 
growth in length of the cortex-roots is 75 mm. in the former case 
and 1 75 cm. in the latter. The growth in thickness would appear 
to be somewhat irregular. Once a 
year, very seldom twice, often only 
each alternate year, a " sinker " 
originates on the inner side of the 
cortex-root near the apex. This 
wedge-like outgrowth is of the 
same breadth as the cortex-root, 
but varies much in thickness, and 
breaks through the cambium zone 
until it just reaches the wood of 
the host-plant, where it elongates 
in the same peculiar manner as 
has been already described in the 
case of the radicle. If the cortex- 
root with its sinkers is exposed, as 
is represented in Fig. 2, we may 
trace back from the apex of the 
root c, and accurately determine 
how many years have elapsed since 
the various sinkers have originated, 
because each year these are em- 
braced by a wood-ring. Even in 
the most recent descriptions of 
the mistletoe we still find, as a 
rule, Schacht's illustration repro- 
duced, which erroneously represents 
young sinkers between older ones 
on the same cortex-root. Water 
and inorganic nourishment are 

absorbed by the whole series of sinkers through their lateral 
surfaces, which are in immediate contact with the water-con- 
ducting organs of the wood, and are first of all conveyed to 
the cortex-root, and through it to the leafy part of the mistletoe. 
From the peculiar way in which the sinkers increase in length 

Fig. 2. — Roots of Visciini album 
in Finns sylvestris. The cortex- 
root, which pushes its apex, c, 
along the bast tissues, b, puts 
forth eight sinkers on the inner 
surface, atid buds and shoots on 
the outer. The oldest portion 
has already been pushed out 
nearly to the dead bark. At e 
the sinkers ofa cortex-root which 
has already been enveloped in 
the bark are shown. 



it is evident that the)' elongate not onl}' towards the side next 
the wood but also towards the side next the cortex. With 
the formation of new phloem-tissues the cortex-roots are con- 
stantly being pushed away from the cambium mantle, as may be 
seen from Fig. 3. In the case of such trees as the silver fir, 

whose stem remains smooth for manj' 
decades before the formation of true 
bark begins, the cortex-roots may 
thus be pushed away from the cam- 
bium mantle without their suffering 
any appreciable injury. They may 
attain to an age of forty years, a 
corresponding age being reached by 
the sinkers, whose length increases 
in proportion to their age. On the 
other hand, trees like the Scotch pine, 
which form bark early, show only 
short sinkers, of a length of 3-4 cm., 
and an age of twelve to fifteen years. 
This is to be explained by the fact 
that owing to the usually more active 
formation of new bast the cortex- 
roots are more quickly pushed away 
from the cambium mantle. The outer 
parts of the cortex are converted 
into bark, and as soon as a portion 
of cortex containing a cortex-root 
is converted into bark, it dries up, 
along with that part of the mistletoe 
root which it contains, and the con- 
nection with the sinker is broken. 
This is distinctly brought out in 
Fig. 3. The sinker then ceases to 
grow, and is covered over sooner 
or later by the new wood-rings. Of course the death of a cortex- 
root does not take place simultaneously throughout its whole 
length. On the contrary, the oldest part — that is to say, the part 
situated farthest from the cambium — dies first, whereas those 
younger portions which arc still enveloped in living cortex 

Fig. 3. — Cross section of a stem 
of Abies pectmata containing 
Visctiin album, a, dead bark 
showing dead cortex-roots ; b, 
region of living bast ; c, cam- 
bium region ; d, cross section 
of a cortex-root showing a 
sinker six years old ; e, a 
sinker eighteen years old ; the 
cortex-root has lately been 
enveloped in the bark, while 
the apex of the sinker has 
withered in the duramen ; at 
/" the cortex-root and the por- 
tion of the sinker in the bast 
have been dead for two years ; 
a cortex-root is shown at g 
which has been dead for six 
years. The boundary line 
between the duramen and al- 
burnum lies at hh ; at x two 
sinkers are shown, those por- 
tions situated in the alburnum 
being still alive. 


remain alive. These, however, are in the same position as the 
roots of a tree that has been felled — that is to say, they can no 
longer conduct nutritive substances to the leafy part of the 
mistletoe, which must therefore die when its feeding-roots are all 
confined to the bark. Its place is taken by numerous root- 
shoots which arise from buds formed on the outer side of 
those portions of the cortex-roots which are still alive. The 
mistletoe represented in Fig. 2 is just such a root-shoot. These 
shoots, which are represented in Fig. 4, form a new root-system 

Fig. 4. — Fart of the stem of a silver fir showing a group of mistletoe plants. The 
bark has been removed from one side in order to show the position of the corlex- 
roots and sinkers. 

of their own, and thus it happens that an old stem attacked by 
mistletoe contains numerous young and old cortex-roots, as well 
as old and young sinkers. In this way the tree comes to bear as 
it were a plantation of mistletoes, which is constantly being re- 
generated by the production of new root-shoots, and which is 
always taking possession of a larger part of the tree. On old 
silver firs and Scotch pines it is by no means rare to meet with 
such mistletoe plantations a yard long and half a yard broad. 
It ought to be mentioned that the living sinkers begin to die at 
their apices (Fig. 3) whenever they become enveloped by the 


advancing duramen. Even in the case of the silver fir and spruce 
it is only the outer wood which conducts water, and in the bole 
this region seldom embraces more than forty to fifty annual 
rings, while in the branches it is much narrower. 

The damage done by the mistletoe to forest, fruit, park, 
and avenue trees is by no means inconsiderable. In the 
Reichswald, in the neighbourhood of Nuremberg, I have seen 
woods of middle-aged Scotch pines where scarcely a tree had 
escaped, and where the foliage of the mistletoes competed for 
effect with the natural foliage of the pines. Where practicable, 
as in orchards, &c., the infested branches should be entirely 
removed before the mistletoe has had time to spread to any 
considerable extent. Simply breaking off the plants only 
induces the formation of vigorous root-shoots at the same 

A few words may here be devoted to the genus Arccuthobiiivi, 
of which a species, ArceutJiobimn Oxycedri, occurs in the south 
of Europe, and also in Austria, where it forms dense bushes on 
Jiiniperns Oxyccdnis ; while in North America quite a number 
of species attack forest trees, especially the Abietinea. These 
grow in the same wa}- as the European form, or induce the 
formation of witches' brooms by the rhizoids which live in the 
cortex causing considerable elongation of the infested branch, from 
whose cortex numerous shoots i — 2 cm. long break through at 
irregular intervals, as happens in the case of ArccutJiobiuni 
Douglasii?- In the case of these plants, also, the nutritiv'e sub- 
stances are absorbed by simple sinkers consisting of a single row 
of cells, or in other cases by vascular sinkers. The injury caused 
to forest trees by these plants is very considerable ; still one 
need not anticipate that these parasites will find their way into 
Europe with the introduction of North American conifers. 

More interest attaches to Loranthns curopLVus, a parasite 
specially common in Austria, but also found occasionally in 
Saxony, the formation of whose roots differs entirely from that 
of the LorantJiacccv already described. 

LorantJius enropcEus is, for the most part, found on the 
common oak, on which account it is known as oak mistletoe, 
though it also attacks Casta)iea vcsca ; and in Austria, especially 
^ See C. V. Tubeuf, /.(■. 



in the Wiener Wald, it has proved \-ery destructi\-e in stored 
coppice, where b\- kilHng the tops of the oak standards it 
prejudicially affects their growth in height. An irregular 
swelling of the size of a 
man's head (Fig. 5) often 
occupies the place of the 
leading shoot. The oblong 
seeds (Fig. 6,/) of the plant, 
which is deciduous, are af- 
fixed to branches by thrushes, 
as in the case of Visanii. 
There they germinate, and in 
a few years the base of the 
parasite becomes completely 
enveloped in a large excre- 
scence which forms on the 
tree (Fig. 6, r). 

The root-s}-stem is to be 
distinguished from that of 
the Lo7'anthaccce alread}^ de- 
scribed by the fact that the few rhizoids which arise on the 
radicle always grow downwards — that is to say, in a direction 
opposed to that of the ascending water — and b}' these rhizoids 

Fig. 5. — A swelling on Quercus Cen-is, a, 
bearing an old plant of Loranthiis, />/>. 

Fig. 6. — Loranthiis euj-opcsiis on a branch of Qiiernis Cerris. a, a young planL ; /', 
a five-year-old plant ; c, an outgrowth of the oak ; d, longitudinal section of a 
root of Loranthtts ; x, apex of the root ; e, cross section of a root ; /, a seed. 

taking up water and food-materials directly from the wood 
without forming sinkers. 

The pointed apex of the root (Fig. 7, x) does not grow outside 
the cambium zone, but in the voung wood— that is to say, in 


the part of the branch that is not completely lignified — and 
always exactly parallel to the longitudinal course of the xylem 
elements. The apex of the root, which is flat upon its inner 
side, advances in a definite region of the young wood, at the 
same time pressing out, splitting, and dissolving the still 
unlignified elements by means of its con- 
vex outer surface. This goes on till future 
progress in the original direction is pre- 
vented by the resistance consequent on 
lignification in the outer layers of the new 
wood which the roots of the parasite are 
unable to split off and dissolve. The root, 
whose apex is thus in a cul-de-sac, is forced 
to form a new growing-point some distance 
behind the apex — namely, at the place where 
the convex outer side comes into contact 
with the cambium zone (Fig. 7, y). From 
this new point growth in length begins 
afresh, and is continued in a zone situated 
nearer the periphery of the wood. During 
the development of an annual ring the root 
of LovantJms (which, of course, can only 
grow during the period when the cam- 

d c b 

Fig. 7. — Youngest por- 
tion of a root of L. 
eiiropceiis. a, cortex 

and bast ; /'. cam- bium produces young wood) is generally 

bium ; c A, young wood ; 

d, the portion of the 
wood-ring of the cur- 
rent year in which 
growth has been com- 
pleted ; c, wood-ring 
of the previous year ; 
z, root of Loraii/hus ; 
X, the apex of the root ; 
y, the point whe-e a 
new root-apex is form- 

thrice compelled to shift the direction of 
its growth farther out, the result being 
that the inner side of the root shows a 
corresponding step-like arrangement, which 
accords with the advance of the wood, as 
is shown in Figs. 6 and 7. The distance 
between two steps measures from 5 to 8 
^^^' mm., while the annual growth in length 

of the root amounts to about 1-5 cm. As the roots grow in a 
direction opposed to that pursued by the ascending water, the 
latter flows directly from the conducting elements of the wood 
into the roots of the Loranthns, at the points of depression. 
The root possesses the power of growing vigorously in thick- 
ness, whereby it is enabled for a series of years to keep pace 
with the increase in thickness of the oak-branch, and thus 


to protect itself against being overgrown. The root usually 
continues to grow in thickness for eight }-ears, though it occa- 
sionally ceases to grow in four, when it is enveloped in the adjoin- 
ing wood owing to the formation of callus.* Thus, while it still 
continues to grow at the point, those portions of a greater age 
than eight years lie embedded in the wood, but they remain 
capable of performing their functions perfectly and of taking 
in food-substances, so long as they are not in the region of the 
duramen (heart-wood), where water is no longer in motion. Even 
then, however, nutriment may still be furnished to ih.Q Loraiithus. 
Here and there processes similar to medullary rays run from 
the roots enveloped in the wood to the cortex, and at this point 
root-shoots may be formed from adventitious buds, although 
this occurs but seldom. 

The gnarled swelling which forms on an oak-branch attacked 
by LoraiitJuis is a very striking object. While the upper part 
of the branch ultimately dies, the rugged protuberance increases 
in thickness, and envelops the whole of the lower part of the 
LorantJms along with its branches. The part of the oak-branch 
that bears the swelling also increases in thickness, although it 
possesses no leaves of its own, and there can be no doubt that 
the products of assimilation of the parasite serve both for its 
own nourishment and for that of the host-plant. 

As it is not expedient to shoot the thrushes even for the 
purpose of preventing the spread of the seeds of Loraut/nis, 
we must, in this case also, minimise the evil by cutting off 
branches which are infested by the parasite. 

Although the Dodders, Cuscutece} which are true parasites 
destitute of chlorophyll, are for the most part injurious only 
to herbs, they are still met with on woody plants with 
sufficient frequency to merit a short description in this 
treatise. Their seeds germinate in spring on the ground. 
The young plant perishes unless its long thread-like stem 

* [Callus is the cushion-like mass of growing tissue which arises at the 
edges of a wound and eventually covers over (occludes) the damaged surfaces 
—see Part II. under the discussion of wounds, &c.— Ed.] 

1 See Sorauer's Handbiich, 2nd Edition, Part II., pp. 32— 48. v. Solms- 
Laubach, Ueber den Ban. und die EnHvicklung parasitischer Phanerogamen. 
Pringsheim'sycr//;-/^. vol. iv. 



finds a suitable host-plant, in which case it twines round the 
stem of the latter and sends numerous absorbing roots, or 
" Haustoria," into the cortex. Although the root which 
originally connected the plant with the ground disappears, the 
dodder nourishes itself by extracting nutritive materials from 
the host-plant round which it twines by pushing its sucker-roots 
as far in as the vascular bundles, in which, by breaking up into 
unicellular threads, they often assume a brush-like appearance. 
If the plants are small they may soon be killed, but in the case 
of larger plants it is only their development that is interfered 
with, and, so far, I have not noticed any appreciable damage to 
woody plants. 

The Ciisaitece are distributed by means of numerous seeds, 
which are produced by dense globular inflorescences, situated 
at some distance apart on the stem ; and it has lately been 
discovered that the plants themselves may survive the winter. 
The only protective measure that is practically applicable in the 
case of this parasite consists in using seed uncontaminated by 
dodder. At the same time, the eradication of the dodder-plants 
which arc so common along hedges and fences is also to be 
attended to. These are the places where we most frequently 
meet with dodder, and there too it will oftenest be found on 
various woody plants. Ciisciita eiwopcea* the greater dodder, 
is the species most frequently met with. It is parasitic on 
almost all trees, as, for instance, Corylits, Salix, Popuhts, Prurms 
spiuosa, but especially on Hiiunihis, Urtica, and Galium. As 
the lesser dodder, Cusciita Epit/iymnin, is specially liable to 
attack clover and lucerne, it is the species most to be feared. 
Besides having numerous other host-plants, e.g. Tliyinus, Genista, 
Callitna, &c., it has also been met with on Vitis. Cuscuta 
Epilinnm is commonest on Limini iisitatissinniin. The other 
species are but seldom met with. 

*[C Europcea is far less common in this country than is the Lesser 
Dodder, C. Epithymum, and C. Epilinnm is not often met with. — Ed.] 




Among cryptogams, also, we meet with plants which, although 
not parasites in the narrower sense of the term, may prove 
directly injurious to other plants by the manner of the attack 
which they make upon them. A case in point is furnished by 
ThelepJiora laciniata} whose thallus '' li\'cs on the humus con- 
stituents in the upper 
layers of the soil, but 
whose sporophores grow 
up and embrace young 
plants, as is shown in 
Fig. 8. Commencing at 
the ground, they enve- 
lop leaves and branches 
so completely as to 
smother and kill them. 
I have found the fer- 
ruginous, sessile, more 
or less confluent sporo- 
phore, with its lacerate 
pileus,"]" commonest on 
young spruces, silver firs, 
and Weymouth pines, 
on which it ascends 

to a height of eight inches from the ground. For similar 
reasons, though to a much less extent, trees may be injured 
by an excessive growth of lichens. A luxuriant growth of 
lichens on the stems and branches of trees is a sign that the 
air is permanently humid. It is also connected, however, with 
the quality of the soil and the rate of growth of the trees, and 

' R. Hartig, Untersuchiingen a. d.forstbot. Inst., I. p. 164. Berlin, 1880. 

*[Thallus is the term applied to the cellular vegetative body of many 
lower plants. — Ed.] 

+ [The pileus of a mushroom or similar fungus is the expanded upper part., 
on a portion of whose surface the spores are produced. — Ed.] 

D 2 

Fig, S. — Thelephora laciniat? 


it is well known that the stems of beeches grown on the best 
soils, especially such as are calcareous, bear but few lichens, 
whereas on the poorer soils, especially such as are sandy, lichen- 
covered stems are very abundant. When a beech grows very 
rapidly in thickness, the formation of periderm* must also be 
rapid, and, since the dead cork-cells on the outer side of the cor- 
tex are soon exfoliated and pushed off, luxuriant development 
of lichens is impossible. Where growth in thickness progresses 
very slowly, the dead cork-cells adhere to the cortex for a much 
longer period, so that lichens are enabled to grow longer and 
develop more vigorously. Under such circumstances, too, mois- 
ture is longer retained, and this also favours the growth of 
lichens. The same remarks apply to trees, such as the spruce, 
whose outer layers of periderm are cast off as scales, or whose 
moribund layers of cortex are thrown off in later life as plates 
of bark. The slower the growth of a tree, the more slowly do the 
outer cortical layers die, and so much the more suitable are the 
conditions for the growth of lichens. Consequently, although the 
presence of lichens is primarily the sign of a permanently humid 
atmosphere, or of a slow rate of growth, it cannot be denied 
that they do some small amount of damage to trees. During 
summer the tree takes in oxygen by means of numerous lenticels, 
and this process goes on even in the older parts of the stem. 
The presence of oxygen in the interior of the tree is absolutely 
necessary for maintaining the processes of metabolic, or chemical 
and vital changes. Now, if the passage of oxygen to the lenti- 
cels t of the cortex is impeded by a luxuriant growth of lichens 
or mosses, we may assume that the tree suffers more or less in 
health. This may furnish us with a reason for the death of so 
many branches of spruces and larches whose crowns are over- 
grown with lichens. 

* [Periderm is the corky covering which replaces the dehcate epidermal 
layer as the stem grows older. Cortex is the green, living cellular tissue 
covered by the periderm, &c. For the connection between "periderm" and 
" bark," see later in Part 11. On a two-year-old twig of a tree, about June or 
July, we usually find epidermis on this year's growth, and periderm on the 
browner, older part. — Ed.] 

f [Lenticels are the perforated or pervious corky warts noticed on the 
periderm of twigs — c.g.^ they are very evident on twigs of chestnut, elder, 
&c.— Ed.] 



It was not till a few years ago that bacteria were recognized 
as plant-infesting parasites, and only in extremely isolated cases 
has it been placed beyond doubt that these low organisms are 
the primary cause of disease in plants.* 

Whereas the processes of decay, and most of the infectious 
diseases of man and animals, may be traced to bacteria, the plant- 
organism is protected against them by the peculiarity of its 
structure, and especially by the absence of circulator}^ channels 
for conducting the nutrient fluids which could serve to distribute 
any lowly organisms which might happen to be present in the 
food. It is only by means of the vessels and intercellular spaces 
that they can distribute themselves in any great numbers in the 
body of the plant, for in other cases they have to pass through 
the cellulose or woody cell-walls, which offer great resistance to 
their attack. 

In addition to this, the vegetable juices, most of which show 
an acid reaction, are unfavourable to their growth. As a matter 
of fact, bacteria have hitherto been found only in the tissues 
of plants whose cells are parenchymatous in character and 
possessed of very delicate walls, as, for instance, bulbs and 
tubers. Sorauer ^ applies the collective name bacteriosis 
to diseases due to bacteria. These diseases are characterised 
by the fact that the succulent parts of the infested plant are 
converted into a slimy glutinous pulp, which emits a most 
repulsive stench. Owing to the action of those bacteria 
which have advanced more rapidly along, and spread out from, 
the vessels, the delicate walls of the cells are dissolved, being 
employed, along with their protoplasmic and other contents, in 
nourishing and fostering the bacteria, whereas the starch is often 
left intact. 

The yellow "bacteriosis" of the bulbs of hyacinths {Bacterium 
HyacintJii) is a common disease. Here the yellow slimy masses 

1 Soxdi\\&i\ Ha?idbi/ch. 2nd Edition. Pp. 74— 112. 

* [Russell has recently put together the literature on this subject in a 
dissertation to the John Hopkins University, Baltimore, 1892. — Ed.] 


of bacteria, called B. HyacintJii by Wakker, occur in the vessels, 
and completely decompose the surrounding tissues. 

Under normal conditions the bacteria do not attack perfectly 
healthy well-developed bulbs. Wounds of some kind are 
necessary, which may be easily caused in transplanting the 
bulbs, or the bulbs are previously attacked by filamentous fungi, 
amongst which a species of HypJiojiiyces almost always accom- 
panies the disease. In a damp situation the bacteria enter 
the wound and cause it to putrefy. 

The wet-rot or "bacteriosis " of the potato, which generally 
appears as an accompaniment of the decomposition of tubers 
and stalks due to Phytophthora infestans, is also a disease 
produced by bacteria.* 

The investigations conducted by Vuillemin a few years ago^ 
have shown that Finns Jiakpensis is subject to a disease induced 
by bacteria which may prove fatal to the tree. The first sym- 
ptoms are that the stem and branches show small outgrowths 
which gradually enlarge till they embrace the whole circum- 
ference, when the portion of the tree situated higher up dies and 
withers. When, as is usually the case, these swellings occur on 
most of the branches, the tree succumbs altogether. 

The olive, also, suffers from a disease which is induced by a 
species of bacterium {^Bacillus OlecB tiiberailosis)?- 

Lately a disease of apple- and pear-trees has been described 
by J. Burrill, of Urbana, Illinois, under the name of " blight," the 
cause of which, according to this investigator, is to be ascribed 
to the invasion of a bacterium. The disease appears to bear 
resemblance to the tree-canker produced by Nedria ditissima; 
and as, in the case of this fungus, large numbers of small 
gonidia resembling bacteria arc produced in the cortex, it remains 
to be seen whether this disease has not been erroneously ascribed 
to a bacterium. 

^ C. R. Seances. November 26th, 1888, and December 31st, 1S88. 

" L., Lcs A fa/adies dc r Olivier, Coniptcs Rendus, December 6th 
and 20th, 1886. 

* [It is extremely probable that in these and other similar cases the 
minute bacteria travel into the tissues, down the tubes of the filaments 
(hyphse) of the fungus, feeding on the decomposing protoplasmic contents of 
the latter. — Ed.] 



Amongst the Myxoniycetes, several — though the number is a 
small one — live as parasites, and cause peculiar swellings to form 
on the roots of their host-plants. To these belongs Plasinodio- 
pliora Brassiccu} which causes '•' club-root "* in cabbages, &c. The 
roots and lower parts of the stems of cabbages which are attacked 
by this parasite exhibit excrescences which vary in size but are 
often as large as one's fist. These soon decay, and the enfeebled 
plants frequently fail to give any return. The disease is com- 
bated, on the one hand, by burning the stumps of all infested 
cabbages, so that the parasite is prevented from spreading in 
the soil, and, on the other, by ceasing to cultivate cabbages for 
some years on ground where the disease has appeared. 

Alder-roots, even when very young, are generally beset with 
much-branched tuberous outgrowths, in whose 
cells Woronin has discovered a fungus which 
he has named Schinzia Alni. 

Recently Moller- has referred the plasmo- 
dium-like structures which occur in the cellu- 
lar tissue of the excrescences of the roots of 
alders to a Myxomycete belonging to the yih. 9.— Excrescence 

genus Plasvwdiop/iora, which he calls Plasmo- o" ^ ''oot of the 

,• , ., ■ ;t., , , . . . , . , . , alder, due to 

aiopliora Aim. vVhether this is identical with Schinzia Aim'. 

ScJiinsia Alni, or distinct from though oc- 
curring simultaneously with it, cannot be decided without 
further investigation. 

The tubercles on the roots of Leg^nninosa^ and Ela^agiie<^, 
in whose parenchymatous cells plasmodium-like structures occur, 
also require to be further investigated. 

^ Woronin in Pringsheim's _/<?//;'/'., vol. xi. p. 54S. 

2 H. Moller, "■ Plasmodiophora Alni:;' Ber. Dciitsch. hot. Cos., 1885, Pt. 3, 
p. 102. 

* [Often termed " Fingers and Toes " and "Anbury." — Ed.] 

t [These tubercles are caused by symbiotic organisms of quite different 
nature from Myxomycetes. See Phil. Trans. 1887 and Proc. Roy. Society, 
1889.— Ed.] 




Every fungus consists of a mycelium and a sporophore. 
The former takes in and elaborates the nutrient materials, and 
discharges all vegetative functions, whereas the sporophores 
produce the organs of reproduction, which may have a sexual or 
an a-sexual origin, in the latter case being produced in a vegeta- 
tive manner by division and abscission, a process analogous to the 
formation of buds in the higher plants. The mycelium has its 
first inception in a tubular outgrowth which is produced during 
the germination of a fungal cell, and which by absorbing 
water, and usually food-materials as well, forms what is called 
a fungal filament, germ-tube, or " Hypha." The germ-tube is 
characterised by apical growth, and by the formation of lateral 
branches, whereby a system of fungal tubes (hyphae) is formed 
which is constantly anastomosing, and which has erroneously 
been compared to a stream with its tributaries and springs. 
This comparison is not apt, because fungal hyphae are almost 
uniform in diameter,* there being usually but little growth 
in thickness of the oldest part of a system of filamentous 

In the case of some species, no partitions form in the fungal 
filaments or hyphs, but as a rule transverse septa, \\hich divide 
the internal space into chambers, are formed a short distance 
behind the apex. Such a hypha is said to be " septate." When 
quite young its contents consist of protoplasm, which is usually 
colourless, and only at some distance from the apex does a 
granular appearance manifest itself, which is generally due to 
the formation of fat globules. The cells of the mycelium are 
frequently filled with large drops of fat, and this is especially the 
case when the mycelium assumes a condition of inactivity, in 
which it remains till growth is again resumed. The potato 
tuber, by storing up reserve materials (in this case chiefly 
starch), which are not utilized in the formation of new 

* [Nevertheless, as the author himself points out on the next page, the 
hyphas developed later are often progressively finer and finer than those 
first produced. This is strikingly obvious in the case of some moulds— ^.^^'.j 
Mucor. — Ed. 


tissues till the following year, behaves in a physiologically 
similar manner. The fatty oil is not unfrequently coloured, and 
especially in the case of " rusts," whose oil is of a golden yellow 
colour, a yellow hue is imparted to the tissues of the leaves and 
cortex in which the mycelium grows. Usually drops of cell-sap, 
or so-called vacuoles, also appear at an early stage in the proto- 
plasm, and these, by forcing most of the protoplasm against the 
walls, impart a frothy appearance to the contents. 

It is only when nitrogenous food-materials are present in 
abundance that the contents of the hyphae are retained for a 
long time. This occurs when mycelia vegetate in or amongst 
the tissues of the cortex, bast, or leaves, which for the most 
part consist of parenchymatous cells. On the other hand, 
the contents disappear early when the mycelium vegetates in 
tissues containing little nourishment, as is markedly the case in 
the wood of trees. When the mycelium of a fungus spreads in the 
interior of a tree, it finds abundance of nitrogenous food-materials 
in the contents of the cells of the medullary rays and the wood- 
parenchyma. It is thereby enabled to produce vigorous hx-phae, 
even when traversing the empty lumina of tracheides, wood- 
fibres, or vessels. When the hyphae have to pass through 
regions of tissue containing no proteids, their apices are supplied 
with protoplasm which is sent forward from behind at the 
expense of the older parts of the hyphae. The latter are 
therefore soon emptied, and become filled with air. Although 
the empty mycelial hyphae persist for some time, they ultimately 
disappear under the decomposing influence of the fungus itself. 
The consequence is that one may frequently fail to find anything 
of the fungus itself, although numerous punctures in the walls of 
the cells show clearly that the fungus had formerly been present 
in that part of the tissues. In proportion as the mycelium 
develops in the wood, so does a dearth of proteids for the 
production of new fungus-protoplasm set in, and this is 
strikingly manifested in the diminished thickness of the new 

The walls of the hyphae, which consist of fungus-cellulose, 
are at first very delicate, though in the course of time they 
occasionally attain such a thickness that the lumen almost 
entirely disappears. In this way it sometimes happens that a 


fungus-body which consists of these thick-walled hyphae 
becomes almost as hard as stone.* On the other hand, there are 
instances of the entire walls — or only their outer, less frequently 
their inner parts — being converted into a mass of slime, and 
under certain conditions the walls, when treated with iodine, 
become as blue as starch would under similar circumstances. 
This occurs, for instance, in the mycelium of Hysteriinn^ and the 
apices of the asci of Rosdlinia quercina. 

At first the hyphae are almost always colourless, but in later 
life the walls very often assume a light or dark brown colour. 
In rarer instances other colours are produced, e.g. the blue green 
of Pezisa csnighwsa, which causes the so-called green-rot in 
the dead wood of the oak, beech, and spruce. Sometimes the 
coloration is confined to the outer or to the inner layers of the 
cell-wall. The mycelium, which increases acrogenously and pro- 
duces lateral branches, generally remains in a filamentous condi- 
tion — that is to say, the mycelial filaments remain isolated, or, at 
most, coalesce only at the points where they cross each other. 

A mycelium which vegetates on the outside of leaves, fruits, 
&c., as is the case for instance with the mildews {ErysipJie), 
is said to be epiphytic. When it vegetates in the inside of 
plants it is called endophytic. In this case it either grows 
from cell to cell by piercing the walls (intracellular), or it 
advances between the cells (intercellular). In the latter case 
it behaves, as a rule, like most epiphytes, sending out short 
branches, known as sucker-tubercles or haustoria, into the interior 
of the cells, in order to extract the nutrient contents. 

When the opportunity is preseated for the filamentous 
mycelium to develop vigorously outside the nourishing 
substratum — as happens most frequently in the case of the 
wood-inhabiting Hyinciiomycetes — it forms a skin-like layer, 
Avhich often attains large proportions. In other cases it may fill 
cracks or other cavities in the stems of trees. Such layers, 
crusts, and masses of fungal growth are best known in the case 
of Polyponts sjilphnreus, P. vaporarius, P. borcalis, Hydiunii divcr- 
sidens, Prametes Pini, Mernlins lacrytnans, 8zc. 

* [Such indurated masses of fungus-mycelium are usually termed Sclerotia, 
in reference to their hardness : they commonly serve as storehouses in the 
sense indicated on p. 40. — Ed.] 


Then, again, the mycelium frequently assumes the form of 
branching strands, which enable the fungus to traverse strata 
containing little if any nutriment. In such a case the strands 
are either formed by the loose union of similar hypha^, when 
they are called R/iicoctonics, or they are peculiarly constructed of 
various kinds of organs. The strands of the dry-rot fungus — 
AI. laoyinaus, for instance — possess, first, organs with wide 
lumina and perforated transverse walls which resemble vessels ; 
secondly, thin sclerenchymatous filaments ; and, thirdly, delicate 
hyphas rich in protoplasm, and provided with clamp-cells. In 
addition to such strands we have the so-called Rhizomorphs, 
which, externally, present a close resemblance to the fibrous 
roots of higher plants, while their internal structure displays 
peculiarities which depend entirely upon the species of fungus 
to which they belong. The best known of these are the 
rhizomorphs of Agaricus melkus, which, \\-hen they have room to 
develop, assume a round shape. In the cortical tissues of living 
trees they spread out in a fan-like manner. Their internal 
structure shows characteristic features, by which they ma\' at 
once be distinguished from the rhizomorphs of other fungi, e.g. 
DematopJiora nccatrix. 

Functions similar to those of the tubers and rhizomes 
of higher plants are to be ascribed to the so-called sclerotia. 
These are peculiarly constructed masses of mycelium, in which 
rich stores of nutriment, especially protoplasm and oil, are 
deposited. After remaining quiescent, it may be for a long 
time, they germinate on the recurrence of favourable conditions, 
and produce either a new filamentous mycelium or the 
sporophore of the special fungus. 

The simplest form of such resting-mycelia is represented 
by the cell-groups of Cercospora acerina. Then come the 
sclerotia of Rosellinia giiercina, and the well-known sclerotia of 
Claviceps purptirea."^ 

The sporophores which spring from the mycelium bear the 
organs of reproduction — that is to say, the spores which give rise 
to new individuals. The same species of fungus frequently 
produces different kinds of reproductive organs, which develop 

* [Rhizomorphs are also, in a sense, extended Sclerotia — see also pp. 40 
and 41. — Ed.] 


on or in variously shaped sporophores. The shape of the 
sporophore is much more characteristic of the species than the 
mycelium, and as the sporophore (which often grows to a large 
size) is almost alwaj's outside the nutrient substratum, whereas 
the mycelium is hidden in it, the uninitiated often regard the 
sporophore as the whole fungus, and pa\- little or no attention 
to the mycelium. 

When the sporophore consists only of single filaments 
springing from the mycelium, it is called a simple or filamentous 
sporophore, whereas the compound fungus-body goes by the 
name of a compound sporophore. On account of the great 
variety in shape and structure exhibited b}' the sporophore, it 
would be going beyond our limits were we to consider it more 
closely at this time.* Cells which are called spores are separated 
off in some way or other in or on the sporophore, and these by 
germinating give rise to new individuals. The cells, from which 
spores directly originate, are known as sporogenous cells. The 
spores are produced either internally, as in the case of the 
sporangia of the Phycoviycctes, and the pouches or asci of the 
Ascoviycetes, or by apical abscission, in which case the mother- 
cell is often called a basidium. 

In the case of some groups of fungi, sexual processes have 
been proved to exist. "f* The course of development, as in other 
plants, has been divided into two sections (generations), of which 
the one called the a-sexual generation begins with the germi- 
nation of a sexually fertilized cell, and leads to the production 
of spores (oospores ; zygospores). The germination of these 
spores gives rise to the second generation, which, in form and 
development, is essentially different from the a-sexual plant. It 
ends with the formation of the male and female sexual 
apparatus and sexual cells, and on this account is called the 
se.xual generation. Spores which do not mark the close of the 
a-sexual generation, but which, like buds, brood-cells, and other 

*[The student may be referred to the works of De Bary, Zopf, and Von 
Tavel for details. — Ed.] 

t [Such a book as this is, of course, not concerned with the morphological 
details of this difficult and involved matter, and the student is referred to the 
works of Tulasne, De Bary, Zopf, and Brefeld for particulars of the subject. 
The outcome of recent researches is to show that sexuality has disappeared 
in the case of most fungi. — Ed.] 


organs of vegetative propagation, produce the same plant-form 
as that from which the)' themselves sprung, are called gonidia. 
Following the example of De Bar}-, this term may be taken to 
replace that of conidia, introduced b\' Fries. 

The gonidia serve chiefly for the rapid propagation of 
fungi during the growing season, whereas, in general, the 
sexual spores serve to carry the species over from one year 
to another. 

I may here briefly sketch the mode and conditions of life 
of the fungi. Just as in the case of phanerogams the ger- 
mination of seeds, and the length of time during which they 
will retain their vitality, are much influenced by external 
factors, so in the case of spores and gonidia the power to 
germinate— varj-ing with the different species — appears either 
immediate!}' after ripening, or not till after the lapse of a long 
period of rest. 

On the other hand, in the case, for instance, of the gonidia of 
the rust-fungi, the power to germinate is lost a few days after 
ripening, whereas the oospores of '<Phytophtliora omnivora may 
remain dormant in the ground for at least four years, without 
losing their vitalit)'. 

The demands as to heat are not so great as in the case of the 
higher plants, and thus it is that we see the most luxuriant 
fungus-vegetation in autumn, at a time when the growth of trees 
has ceased. The optimum temperature for fungi, as for other 
plants, varies ver}' much, but in this connection we still await the 
results of reliable investigations. In the case of those fungi 
which concern us here a temperature over 212° F. (100° C.) is 
undoubtedl}' alwa)'S fatal. 

One vital condition, of extreme importance for fungi, is a high 
degree of humidity of the air or of the substratum in which they 
develop. This is due not only to their requiring large quantities 
of water, but also, and much more, to the ease with which the 
mycelia or young sporophorcs die in a dry medium from the 
effects of excessive evaporation. On this account it is very 
seldom possible for the mycelium to develop in the open air. 
It is for this reason also that in all the rusts and smuts, and even in 
the case of a great number of Disconiycctes, the sporophores — 
which usuall)' require to scatter their spores outside the plant — 


are formed under the protection of the epidermis of the host- 
plant, which is ruptured only after the spores have ripened. 

The fact that in summer, in spite of a more favourable 
temperature, far fewer so-called " Toad-stools " spring from the 
ground than in October, when the atmosphere is relatively much 
more humid, shows clearly how dependent on a constant supply 
of moisture in the air is the development of sporophores that 
expand entirely outside the substratum.* The extensive 
distribution which the larch-fungus, Peziza Willkoinmii, has 
experienced in the plains of Germany is almost entirely due to 
the abundant production of fructifications and spores which have 
ripened perfectly in the moist, stagnant air of the dense low- 
lying woods ; whereas in the breezy Alps the fructifications 
almost always wither before they have had time to mature. 

Not only does a moist atmosphere affect the ripening of the 
fructifications and the germination of the spores outside the 
plant, but it also appears to have great influence on the develop- 
ment of the fungus even inside the plant. This assumption is at 
least supported by the fact that Caoina pinitorqnmn, which is a 
perennial in the shoots of the pine, assumes the proportions of a 
plague when the month of June is wet, but causes scarcel}- any 
appreciable damage when the weather is dry. 

As regards their adaptations for nutrition, fungi ma\- be 
arranged in two great divisions. Parasite is the term applied 
to those fungi which draw their nourishment from living 
organisms ; saprophyte, to those which live on dead bodies. It 
is not possible, however, to draw a sharp line of demarcation 
between these two categories. To begin with, it may often 
be disputed whether an organic body is to be called dead 
or living. By far the greater part of the wood of trees is made 
up of dead cells, the walls of which alone remain ; and only 
a relatively small part, consisting of the parenchymatous cells 
of the wood and medullary rays, remains alive and contains 
protoplasm. Seeing that many fungi live only on the old 
stumps of trees and on trees that have long been felled or 
otherwise killed, whereas others destroy growing trees, it would 

*[The whole subject needs thorough investigation, however, and it would 
be particularly valuable to have more information as to the importance of 
sunlight and other factors in this connection. — Ed.] 


appear to be necessary that we should regard the sound wood of 
a growing tree as being aHve, even although only a portion of its 
cells may exhibit the phenomena of life.* In man}- cases it is 
difficult to decide whether wood — e.g. the duramen in many trees 
— was actually living when attacked by the fungus-mycelium, 
or whether its parenchymatous cells were then dead. But apart 
from those doubtful cases, in which it is difficult to decide at 
once whether a fungus is existing as a parasite or as a sapro- 
phyte, there are many fungi which occupy a position somewhere 
between those which are strictly saprophytic and those which are 
strictly parasitic. Numerous fungi are in a position to complete 
the whole course of their development as saprophytes, though, 
under certain circumstances, they may also live in a purely 
parasitic manner. Agariciis vielleiis and the genus Xectria may 
serve as examples. Such fungi are designated facultative 
parasites. Other fungi, which, as a rule, go through the whole 
course of their development as parasites, but which arc capable 
of growing as saprophytes, at least during certain stages of their 
existence, are designated facultative saprophytes. To this group 
belong, for instance, PJiytopJithora ovinivora and Ccrcospora 
acerina. We have thus to distinguish four groups : i. Obligate 
saprophytes. 2. Facultative parasites. 3. Facultative sapro- 
phytes. 4. Pure — that is to sa}', stricth' obligate — parasites^ 
which can only grow parasitically, e.g. the group Uredinecc. 

The spread of an infectious disease may take place in two 
distinct ways, either by infection caused b\- the m\'celium, or 
by infection caused by the spores, including the gonidia. 

Infection by the mycelium is met with in nature most frequentl}- 
in the case of those parasites which grow below ground. This is 
to be explained by the fact that the varying amount of moisture 
in the air admits of the development of the mycelium above- 
ground only in exceptional cases, as, for example, in HerpotricJiia 
and TricJiospIuvria. 

In the case of infection by the mycelium, it is, to a certain ex- 
tent, the same individual fungus that spreads from root to root and 

* [In many such cases the destructive fungus may be looked upon as a 
Saprophyte, when viewed with regard to the immediate seat of its action — 
e.g., wood— but as 2l parasite with regard to the tree as a whole, whose life is 
destroyed in consequence of the secondary results of the damage. — Ed.] 


from branch to branch. Thus, when a disease spreads in a wood 
in this way, it does so with relative slowness, but, in dense woods 
at least, it is, as a rule, characterised by the death of all or most 
of the trees inside the local area of distribution. The result is 
that blanks, varying in size, gradually occur in the wood. 

In the case of Trmnetcs radicipcyda,\\\& most dangerous enemy 
of spruce and pine plantations, contact of the diseased root 
containing the fungus with the sound root of a neighbouring tree 
is necessary in order that the latter may be penetrated by the 
mycelium which protrudes from between the bark-scales. In the 
case of Agaricus inelleus mycelial strands, in the form of rhizo- 
morphs, spring from the diseased roots, and proceed to spread 
underneath the surface of the ground. The roots of sound 
conifers that are encountered are embraced, and an entrance is 
effected between the bark-scales : these are forced off b}- means 
of the conical apices of the rhizomorphs, which then bore into 
the living tissues. 

In the case of Rosellinia qiiercina, which destroys the roots of 
the oak, the delicate filiform mycelium — which here and there 
forms rhizoctones — spreads during moist warm weather from the 
diseased plant into the upper layers of the soil, where it attacks 
and destroys the roots of neighbouring plants in a manner which 
will be described more fully later on. On account of the 
mycelium being capable of forming small round sclerotia on oak- 
roots, and of assuming a resting condition, the parasite is after- 
wards enabled to resume the growth which has been interrupted 
by such unfavourable conditions as cold or a temporary lack of 
moisture in the soil. 

Deinatophora necatrix spreads in a similar manner in vine- 

The distribution of a parasite by spores and gonidia is not, as 
in the case of infection by the mycelium, confined to plants in the 
immediate neighbourhood, although these are certainly most 
exposed to the danger of infection. It may, in fact, happen 
that trees at a great distance are infected, while those in the 
immediate neighbourhood remain sound. When treating of 
special cases we shall have occasion to bring into prominence 
how various are the conditions that influence this question, and, 
in particular, how animals and men, by spreading the spores, may 


induce the outbreak of an epidemic. Here a few examples may 
be cited in illustration of this point. 

PhytophtJiora oiiuiivora produces spores (in this case called 
oospores) in the interior of the seedling, as the result of sexual 
fertilization. When the plants decay, these spores get into the 
ground, where they may rest for a series of years and produce 
the disease afresh, should the right kind of seedlings be present. 
But, in addition to these oospores, the parasite produces numerous 
gonidia on the surface of its leaves. These are capable of 
germinating at once, and are blown by the wind, or conveyed by 
animals or men, to plants in the neighbourhood, the result being 
the formation of new hotbeds of infection. 

In the case of Tranietes radicip:rda, which, on the spruce at 
least, almost always produces its sporophores in holes in the 
ground, new centres of infection are usually established by spores 
that have been distributed by mice. 

The smut of wheat is generally induced by employing seed to 
the outside of which spores of the smut-fungus have adhered^ 
but it may also be caused by manuring with fold dung if infected 
straw has been used as litter. 

The conditions become most interesting in the case of heter- 
cecious rust-fungi — that is to say, parasitic fungi which complete 
the various phases of their development not on the same plant 
but on two different species. Here mention need only be made 
of the connection between the fungus of the barberry and rust of 
wheat, or between /Ecidiiun abietimun and Chrysouiyxa RJiodo- 
dendri and Chrysoniyxa Ledi, or, finally, between ^Ecidiuvi 
colinnnare and MelaJiipsora Goeppertiana. In the case of these 
parasites the occurrence of the disease depends on the presence 
of both host-plants : still De Bary has demonstrated that in cases 
of necessity Chrysoniyxa Rhododendri may exist without spruces, 
and it appears to me to be beyond doubt that Melampsora 
Goeppertiana is able to develop without the silver fir. We know 
only one or other of the stages of development of a series of 
rust-fungi, and it remains to be determined what the other fungus- 
forms are with which they stand in relationship. 

The method of attack of parasites, also, reveals the most 
marked differences. Whereas the epiphytes — whose mycelium 
vegetates externally on the epidermis of leaves, fruits, and stems 
— send only delicate absorbing organs into the interior of the 



epidermis, the endophytes must send either their germ-tubes, 
arising from spores germinating externally, or else their mycelia 
into the interior of the plant. 

According to the mode of attack, we may divide parasites 
into two main groups. The first comprises those which have the 
power to attack uninjured plants ; the second, those which can 
effect an entrance only through a wound. It is those belonging 
to the latter group which are accountable for infectious w^ound- 
diseases. The former are partly confined to the very early stages 
of development of the plant, or of the shoots, leaves, or roots ; in 
rarer instances they also force their germ-tube into the stomata 
and lenticels of more mature leaves and shoots. It is only when 
the mycelial growths are very vigorous, like those of Agariais 
vielleiis and Tramctes radiciperda, that they are able, b>' entering 
between the bark-scales of the root and forcing them apart, to 
bore even into cortical tissues covered with corky layers. 

The mode of attack of Rosellinia quercina affords one of the 
most interesting examples of this kind. The main root of the 
young oak is protected against external attack by a corky mantle 
of considerable firmness. The mycelium of Rosellinia is con- 
sequently able to get at the interior only by first killing the fine 
lateral roots, and as these traverse the corky la}^er the hyphae 
form breaches in the protective covering. At the points where 
the lateral roots pierce the cork)- mantle the mycelium develops 
flesh}- tubercles, which then send one or more processes through 
the breach into the interior of the root. It is only some time 
afterwards that the destructive filiform mycelium is formed at 
the apex of these processes. 

Wounds admitting of the entrance of parasites into the interior 
of trees arise in many wa}-s. Reference need only be made here 
to such agencies as animals, man, hail, wind, snow, &c. 

The effects produced b}' parasites on the tissues of the host- 
plant can be explained only by assuming that in each species of 
fungus a peculiar enzyme (ferment) is produced in its protoplasm, 
which by being excreted through the h\'phai is communicated 
to the adjoining cells.* 

Ver)- often the m}-celium \egetates in living parenchj-matous 

* [That such enzymes are really formed and excreted by the protoplasm has 
been proved. For instance, Botrytis excretes an enzyme capable of dis- 
solving cellulose— see Aitnals of Botany, 1888, "A Lily-disease." — Ed.] 


tissues without producing any appreciable effect on them. 
Especially is this the case when the cells have already attained 
the condition of permanent tissue before the mycelium has 
appeared in or between them. 

The mycelium of Calyptospora has no apparent action on the 
permanent tissues of Vaccinuim Vitis-IdcBa ; whereas, in v^xy 
)'oung shoots, it causes enlargement of the parenchymatous cells 
of the cortex, with the result that very remarkable swellings are 
produced on the stem. 

One of the most frequent results of the action of fungi is that 
a stimulus is given to cell-division. Mention may be made of 
the swellings on the stems of silver firs whose cortical tissues are 
infested by ^cidhim elatimim, of the swellings on the stems of 
junipers owing to Gymnosporanghun, &c. Still more frequently 
the infested parts are stimulated to display altogether abnormal 
growth. Flowers, fruits, and portions of stem of various species 
of plants are transformed in a most peculiar manner by fungi 
belonging to the genus Exoascus. It does not necessarily follow, 
however, that their vitality is thereby prejudicially interfered 
with {e.g. witches' brooms of the hornbeam, &c.) 

Changes in the cell-contents are often noticed which are 
indirectly induced by fungi. This is the case, for instance 
when the mycelium of Hysterium inacvosponini kills the elements 
of the bast at the base of spruce-leaves, thereby destroying 
their capacity of conducting plastic materials, while the other 
parts of the leaves still live and assimilate. The result is that on 
account of the newly formed carbo-hydrates not being able to 
get away from the leaf, all the cells become packed full of starch. 

The tannin which is dissolved in the cell-sap offers excellent 
food for the mycelium of Polyponis igniariiis, being absorbed first 
of all by the hyphae which penetrate the sound oak-wood, after 
which it undergoes metabolic changes in the youngest parts of 
the mycelium. The occurrence of mycelia in oak-timber is there- 
fore followed by the disappearance of tannin, the smell of which 
has long been regarded by practical men as a proof of the sound 
condition of the wood. The conversion of a portion of the cell- 
contents or of the cell-walls into turpentine under the action of 
the hyphae of Peridermiinn Pint is also interesting. Although 
it often happens that the starch-grains disappear very soon from 

E 2 


amongst the cell-contents, as, for instance, in the case of attack 
by Phytophthora omnivora, it also not unfrequently occurs that 
starch resists the destructive influence of wood-parasites longer 
than the thick lignified walls of the cells in which it is contained. 
In fact the manner of decomposition of the starch-grains varies 
exceedingly according to the species of fungus that attacks them. 
Similarly as regards the cell-walls. The solvent action of living 
hyphae is manifested in two distinct ways. Where a hypha 
touches a cell-wall it dissolves the particles of calcium oxalate 
contained therein, exactly as a root-hair, by means of the 
acid solution which it exudes, dissolves the particles of cal- 
cium carbonate with which it comes into immediate contact. 
This action is confined to the surface of the cell-wall which is 
actually in contact with the fungus-filament. But every para- 
sitic fungus that lives in the wood of growing trees destroys 
the wood in a manner peculiar to itself When one and 
the same species of fungus, e.g. Polyponis siilpJmreiis, vegetates 
in trees of such different species as oak, willow, and larch, it 
changes the wood so peculiarly in a short time that at first 
sight it is difficult to distinguish these timbers from each other, 
although, in a sound state, they are so strikingly different. This 
can be satisfactorily accounted for only by assuming that each 
species of fungus exudes an extremely powerful and charac- 
teristic ferment, which permeates the walls for long distances, and, 
to begin with, frequently dissolves only the incrusting substances, 
more especially the lignin. 

In the accompanying figure (lo) the upper part of the wall is still 
lignified, whereas the lower part consists of pure cellulose. After 
the removal of the lignin the middle lamella, which is most ligni- 
fied, is the first to disappear, the result being that the various 
organs become completely isolated, as happens when sound wood 
is treated with potassium chlorate and nitric acid. The hyphee 
which pierce the walls with their apices disappear later on, when 
they themselves are dissolved by the ferment. In Fig. ii is 
shown how the elements of the wood of the oak have been com- 
pletely isolated and dissolved by the action of a ferment. 

In the case of other wood-parasites the decomposition takes 
place in the following manner. By the extraction of the incrust- 
ing substances a zone bordering the lumen is first converted into 


cellulose, after which decomposition spreads generally through- 
out the walls. Thus the walls constantl)' become thinner, till 
finally onl\- the corners remain where three tracheids join (Fig. 
12). Several wood-parasites, e.^: Polypoms 
Schiveinitzii -BiXxA P.sulphureus, induce a form 
of decomposition owing to which the walls, 
with the exception of the middle lamellae, 
shrink so much as to give rise to numerous 
cracks which ascend from right to left. By 
certain adjustments of the microscope we of 
course see simultaneously the correspond- 
ing cracks in that half of the wall 
which belongs to the neighbouring fibre, 
and this makes it appear as though the 
cracks crossed each other. The walls, 
which are very rich in carbon, assume a 
brown colour (Fig. 13). We shall direct 
attention in the special division dealing 
with this subject to other forms of decom- 
position, all of which are characteristic for 
some species of fungus. Here it need only 
be mentioned that the question whether all 
the organic parts of the lignified cell-walls 
require to be absorbed by the mycelium 
of the fungus before being decomposed 
into carbonic acid and water, or whether 
to some extent they are directly oxidized 
and converted into these substances, can- 
not at present be finally decided. As a 
large quantity of oxygen must be made 
use of during decomposition, its rapidity 
depends to a great extent on the facilities 
that are afforded for the entrance of air 
to the interior of the tree. A certain 
amount of air is present in every wood}' 

fibre. In dicotyledonous trees the air is conducted to distant parts 
by means of the vessels and intercellular spaces, and in resinous 
conifers by the resin-ducts ; but in the case of the silver fir 
and other conifers destitute of resin-ducts the mode by which 

Fig. 10. — Tracheid of 
Finns sylvestris, de- 
composed by Tranietes 
Pini. The primary 
cell-wall has been com- 
pletely dissolved as far 
as a a. In the lower 
part the secondary and 
tertiary layers consist 
only of cellulose, in 
which lime - granules 
are distinctly visible, 
h : filamentous my- 
celia, c, penetrate the 
walls and make holes 
as at d and e. 



the entrance of air to the interior of the tree is ensured has yet 
to be explained. The carbonic acid . which is formed can 
escape by the same way as the oxygen entered. To what 
extent carbonic acid and oxygen when dissolved in water may 
traverse the wood remains to be determined. 

Fig. II. — Decomposition of oak by Tlie'ephora Ferdix. a, tracheids containing a 
few filamentous mycelia,and showing the perforations in the walls which these have 
occasioned ; b, wood-parenchyma wiih starch-granules, the latter being in process 
of solution, and having to a certain extent disappeared from the neighbourhood 
of the cell-walls; <:, vessels containing hyphre ; d, sclerenchymatous fibres show- 
ing filamentous mycelia and perforations ; e and /, tracheids which are com- 
pletely isolated owing to the dissolution of the primary cell-walls ; the thickened 
rings of the bordered pits are also found isolated between the tracheids. On 
account of the organs being dismembered the openings into the bordered pits no 
longer cross each other ; g, wood-parenchyma, completely dismembered and 
almost entirely dissolved ; h, tracheid just before final disappearance ; /, scleren- 
chymatous fibre much decomposed ; k, a tracheid whose walls have become 
fissured before being dissolved. 

In concluding these general considerations, I have still to 
discuss the question whether any — and, if so, what — means are 
at our disposal for combating the ravages of fungi. I am 
convinced that every forester who has received a scientific 
education will take a deep interest in obtaining a knowledge of 
what tree-diseases are, and how they originate, even though it 



may not be possible to apply any practical remedies. It is by 
no means the first duty of science to call attention to the 
practical value of a new discovery ; nor should research be 
primarily directed to those fields which promise to yield results 

Fig. 12. — Decomjjosition of spruce-timber by Polyporus horealis. a, a tracheid con- 
taining a strong mycelial growth and a brownish yellow fluid which has originated 
in a medullary ray ; at b and c the mycelium is still brownish in colour and 
very vigorous. At d and c the walls have already become much attenuated 
and perforated ; here the mycelium has been less abundantly supplied with nutri- 
ment and the filaments are very delicate ; at /' the pits are almost completely 
destroyed ; at g and h only fragments of the walls remain. The various stages in 
the destruction of the bordered pits are to be followed from / to r ; at i the 
bordered pit is still intact ; at k the walls of the lenticular space have been largely 
dissolved, their inner boundary being marked by a circle ; at / one side of the 
bordered pit has been entirely dissolved ; at in and n one sees a series of pits 
which have retained a much-attenuated wall on one side only — namely, on that 
which is provided with the closing membrane. In making the section a crack 
has been formed in this wall. Between o and r both walls of the pits are found 
to be wholly or partially dissolved, only at / and q has the thickened portion of 
the closing membrane been preserved ; at J the spiral structure of both cell-walls 
is distinctly recognizable. These walls when united form the common wall of 
the tracheid ; at / hyphce are seen traversing the tracheids horizontally. 

capable of immediate conversion into hard cash. The duty 
of science is nobler and higher than that. But if, in our 
search, we succeed in fathoming the mysteries of nature, and, 
at the same time, obtain results of practical value to humanity. 



then it is our duty to direct attention to these. This I have 
never neglected to do, and, although I do not under-estimate the 
many difficulties which foresters will long have to encounter 
in endeavouring to put into practice the 
p results of scientific investigation, still I hold 

that, as the guardians of the forest, it is their 
duty to make themselves acquainted with the 
results of scientific investigation, and care- 
fully to watch over the health of what is 
committed to their charge. Not only must 
they do everything that may prevent disease^ 
but they must also instantly adopt energetic 
measures to nip an existing disease in the 
bud, and so prevent its further spread. 

As every disease must necessarily be treated 
differently, this is not the place to enter on 
the consideration of specific measures. But 
just as human health is better maintained by 
the observance of certain general laws, so 
there are also general rules for the treatment 
of woods, by following which we may pre- 
serve the health of the trees. 

The best prophylactic measure against the 
occurrence and spread of an epidemic is 
the formation of mixed woods. Infection, 
walls consisting ^^j-j^ y^^^^^^ ^^^ ^^^^^.^ gj-Qund, is least likely 

chiefly of lignin. ° ' -' 

Fig. 13. — Portion 
of a tracheid of 
J'iiiiis decomposed 
1) y P I y p rii s 
Schu'cinitzii. Most 
of the cellulose has 
been extracted, the 

to occur when every tree is isolated by being 
surrounded by others of a different species- 
On ground which is infested by root-parasites, 
or which contains resting-spores whose vitality 
is preserved for many years, it may be advis- 
able, under certain circumstances, to abandon 
the cultivation of some particular species of 
tree. One should also try to prevent the 
distribution of spores either by men or 

animals, and especially so in the sale and purchase of young 


In the case of root-parasites, the therapeutic measures to 

be adopted when a disease has broken out consist partly in 

In drying, cracks 
have been formed, 
which however do 
not extend to the 
primary wall, a b. 
These cracks are 
seen to cross each 
other at the bor- 
dered pit (•, and at 
the perforations d 
and t' ; a simple fis- 
sure is shown at f. 


promptly pulling up or otherwise eradicating the diseased plants, 
and partly in isolating the infected area by means of narrow 
trenches. As a general and most important measure, it is 
advisable at once to remove from the wood all plants attacked 
by fungi, so that their spores may not spread infection. Tidiness 
is the first hygienic law in sylviculture. 

Having noted above the more important points that should be 
kept in view in studying the parasitic fungi, I shall now, in 
accordance with the plan of this work, pass on to a systematic 
examination of the parasites that occur in woody plants. 
As regards fungi that are parasitic on farm or garden crops, I 
shall shortly refer only to such as are of general practical 
importance. For plant-parasites not included in this work I 
must refer to the handbooks of Frank or Sorauer. 

Following the most recent classification of the fungi, which 
distinguishes three groups — namely, PJiycoviycetes (Algal fungi), 
Ascoinycetes, and Basidiomycetes — I shall begin with the first 
group. This embraces five orders — namely. Zygomycetes, 
EiitomophthorecE, Saprolegiacece^ PeronosporecB ChytridiacecE, and 

Of these orders there are only two that need be considered 


The PeronosporecE are true vegetable parasites, whose mycelium 
ramifies in the tissues of higher plants, the hyphae being for the 
most part intercellular, though occasionally also intracellular. 
Special absorbing organs (haustoria) are employed for abstracting 
the nutriment from the living cells, which consequently die after 
a shorter or longer period. The sporophores which spring from 
the mycelium either grow through the stomata or burst through 
the epidermis. These in various ways form sporangia, which 
produce gonidia, often motile. 

Having moved about for some time in a drop of water as 
swarm-spores, the gonidia develop a germ-tube, though the 
sporangia may also germinate directly without having first 
produced swarm-cells in their interior. 

* [For details as to the classification of fungi the reader may be referred 
to the text-books of De Bary, Zopf, and Von Tavel. — Ed.] 


In the tissues of the host-plant, though occasionally outside 
of it, female sexual organs (oogonia) originate on the mycelium, 
and during fertilization the male sexual organs, called pol- 
linodia or antheridial branches, are brought into contact with 
these oogonia. The antheridia send a minute process (the fer- 
tilizing-tube) into the interior of the oogonium, which is fer- 
tilized by its protoplasm receiving a small portion of the con- 
tents of the antheridium. This gives rise to the formation of 
the egg-spore (oospore), which is provided with a thick cell-wall. 

While the gonidia, being easily detached and carried by 
wind or animals, provide for the rapid distribution of the 
parasite during summer, the oospores reach the ground in 
the dead and decaying parts of plants. There they pass the 
winter — indeed, in such a position they may remain alive for a 
number of years — after which they either germinate directly, or 
first of all produce sporangia with zoogonidia.* 


The disease caused by this parasite was noticed in forestal 
publications over a hundred years ago as " the disease of seedling 
beeches," and cannot be unknown to any forester employed in 
beech woods. When seedlings are abundant after a rich seed- 
year the disease is to be met with over the whole of Germany, 
and the more plentifully the wetter the months of May and June. 
The fungus also attacks other broad-leafed trees, e.g. Acer, 
Fraxinus, Robinia, as also herbaceous plants such as Fagopymm, 
Clarkia, Scnipej'vivuin, &c. The parasite is equally widely dis- 

' 1 described this parasite in 1875 in the Zeitschrift fiir Forst- uini Jagd- 
wesen, pp. 117 — 123, under the name of P. fagi. A detailed account of the 
history of its development, and of the disease to which it gives rise, along 
with a plate, was contributed by me to the Unteisuchimgen cms dein forst- 
hotanischcn InstHut, 1880, pp. 3 — 57. In 1875 — that isto say, simultaneously 
with me — Schenk described this fungus under the name of P. sempervivi. 
In order to settle the question of priority De Bary selected the name 
Phytcphthora omnivora {Beit rage zur Morpli. und Phys. der Pilze, 1881, 
p. 22). 

* [For an account of the Phycomycetes peculiar to Britain the reader 
may be referred to Massee's British Fungi (Reeve & Co., 1891). — Ed.] 



tributed in the seed-beds of conifers, where it may be met with 
on the seedlings of every species. 

The disease may attack seedling beeches before they have 
reached the surface of the ground, in which case a dark 
discoloration spreads from the primary rootlet, and the plants 
die off. Or, not till the 
cotyledons have unfolded 
does the stem above and be- 
low them, or at their base, 
become dark green and change 
colour (Fig. 14, a,b) ; or simi- 
lar spots may be recognized on 
the cotyledons (Fig. 14, c), or 
on the primary leaves (Fig. 
14, d). Should the weather 
remain long wet, decomposi- 
tion quickly spreads over the 
w^hole plant, while during dry 
weather the plants wither and 
assume a reddish brown and 
scorched appearance. Young 
sycamores, ashes, and robinias 
show similar pathological 
symptoms, and, in particular, 
very black streaks will fre- 
quently be found running up 
or down the stem from the 
base of the cotyledons. Fre- 
quently it is only the apex of 
the stem and the leaves that 
become black, in which case 
the plant recovers ; but if, on 
the other hand, the lower part 

of the stem is attacked, recovery is impossible. Where the seeds 
of conifers are sown in rows, it is not unusual for a large number 
of the plants to perish before they have appeared above the 
surface of the ground. The roots and stem.s usually decay, 
and the young plants die or wither without any mechanical 
injuries being observable. It is worthy of note that, owing 

Fig. 14. — Diseased seedling beech. Stem 
below the cotyledons dark green at a ; 
cotyledons diseased at b and c ; first 
foliar leaves showing blotches as at d. 



to the death and disappearance of the whole of the seedhngs 
at certain places, blanks four inches and more in length may 
be formed in the seed-drills. 

The infectious character of the disease may be gathered 
from the peculiar way in which it is distributed. A diseased 

plant soon becomes surrounded 
by diseased neighbours, and thus 
the epidemic spreads centrifu- 
gally in beds that have been sown 
broadcast, and in two directions 
where the seed has been drilled. 
Should a frequented footpath lead 
through a beech wood that is being 
regenerated b}' seed, all the plants 
growing on the path and along 
the sides contract the disease and 
die in a short time. It has also 
been observed that if the disease 
has once appeared in seed-beds it 
usually recurs in succeeding years 
in a much-accentuated form. The 
disease is known to be greatl}' 
favoured by rainy weather — espe- 
cially if accompanied by heat — and 
by any kind of shading, whether 
produced by standard trees or b)" 
artificial covering. The first ap- 
pearance of the disease in any year 
can only be due to the oospores 
of the parasite, which lie dormant 
in the soil during winter, and 
infect the germinating seedlings in 
spring. The mycelium spreads in 
the tissues of the seedling, and, 
in the case of the beech, both in the stem and in the cotyledons, 
the latter being probably attacked as they are being pushed up 
through the ground. In the tissues of the cotyledons the mycelium 
is almost entirely intercellular (Fig. 15, b), withdrawing the 
nourishment from the interior of the cells by means of small 

Fig. 15. — Cellular tissue from the 
cotyledon of a diseased beech. 
The starch-grains have been ab- 
stracted from the protoplasm 
which has withdrawn from the 
cell-walls, a ; the mycelial fila- 
ments, which are of varying 
thickness, h b, giow intercellu- 
larly, and are provided with 
minute haustoria ; each fertilized 
oogonium contains an oospore, <(■. 



roundish haustoria. The consequence is that the starch- 
grains soon disappear, and the protoplasm dies and contracts 
from the cell-walls (Fig. 15, a). While the fungus continues to 
spread in the plant, numerous hyph« break through the epidermis 
to become sporangiophores (Fig. 16). The swelling of the ex- 
tremity (Fig. 16,/) gives 
rise to a lemon-shaped 
sporangium, which is pro- 
vided with a papilla at the 
apex, and a short pedicel 
at the base (Fig. 16, g). 
After its abscission from 
the stalk the latter elon- 
gates afresh to produce a 
second sporangium (Fig. 
16, g,h), and meanwhile 
the first generally drops 
off (Fig. 16, i). Should 
the sporangia come into 
contact with water — as, for 
instance, a drop of rain 
or dew which has lodged 
about the cotyledons — 
they germinate directly, 
putting forth one or more 
germ-tubes, which gener- 
ally proceed to bore into 
the epidermis of the host- 
plant. In other cases the 
protoplasmic contents of 
the sporangium form a 
large numberof extremely 

minute and very active gonidia (swarm-spores, or zoospores — 
Fig. 17, c), which are capable of free movement after the apex 
of the sporangium has been dissolved. For some hours these 
swim about in a drop of rain with all the activity of infusoria, 
until they settle on the epidermis of the host-plant, when they 
germ.inate with one or even four tubes (Fig. 17, a, b). Some- 
times the zoospores germinate in the interior of the sporangium 

Fig. 16. — Epidermis of a diseased cotyledon 
of the beech, a, the external wall of an 
epidermal cell ; b, the cuticle ; c, a hypha 
which has intruded itself between the wall 
and the cuticle ; at of it pushes up the latter ; 
at e it appears on the surface, and at /' it 
forms a sporangiophore. After producing 
the first sporangium it branches at g, to 
form a second //, while the first drops off 
at i ; a stoma is shown at k from which 
sporangiophores project. 



when their germ-tubes may either break through the lateral 
walls or push out through the open apex of the sporangium 
(Fig. 17, c). In either case the germ-tubes creep about for 
some time on the epidermis of the host-plant, after which they 
force their way into the interior, and especially at those places 

where the lateral walls of 
the epidermal cells arc 
situated (Fig. 17, ^, d). 
Less frequently the germ- 
tubes reach the interior by 
first traversing an epi- 
dermal cell (Fig. 17, e). 
Under favourable circum- 
stances the development 
of the parasite may have 
progressed so rapidly in 
the plant that three or four 
days after infection new 
sporangiophores may 
make their appearance. 

The sporangia, and the 
swarm-cells that form in 
them, serve to spread the 
disease during the months 
of May, June, and the 
earlier part of July. They 
either fall directly on to 
neighbouring plants, or 
are carried by the wind. 
Their distribution is great- 
ly assisted by animals — 
as, for instance, by mice 
(in the seed-beds) — and game, but most of all b}' man. The death 
of all plants along a path is the result of the sporangia and 
swarm-cells adhering to the trousers or coats of passers-by, and 
afterwards dropping off farther along the path. 

From what has been said, the favouring influence of rain, shade, 
&c., is sufficiently evident. In dense seed-beds the hyphae grow 
directly from one plant to another, and this offers a simple 

I'lG. 17. — The surface of the stem of a seedling 
beech. At a d zoospores are seen which 
germinate and send their germ-tubes into 
the interior at a point where the common 
wall of two epidermal cells abuts on the sur- 
face ; c, a sporangium whose zoospores have 
germinated, d/, in its interior ; at t' a germ- 
tube has grown directly into an epidermal 
cell ; at ^t; a germ-tube has reappeared on 
the surface. 


explanation for the total destruction of all the plants at certain 
parts of the bed. 

As the result of the sexual act the oospores originate in the 
tissues of the host-plant in the following manner. In the inter- 
cellular spaces of the leaf-parench)'ma of the beech, globular 
swellings appear at the apex of numerous short hyphal branches, 
and become oogonia ; while smaller so-called antheridia originate 
in like manner either at the apex of special hyphae, or on the basal 
portion of the stalk of the oogonia. In each case a transverse 
septum delimits the organ from its stalk (Fig. 15, c c). The 
antheridium having very early laid itself against the outer wall 
of the oogonium, and the most of the protoplasm of the latter 
having become aggregated to form an oosphere, the antheridium 
next develops a short process, the fertilizing-tube, which it pushes 
into the interior of the female organ as far as the oosphere. A 
part of the contents of the antheridium is then transferred to fer- 
tilize the oosphere, which is thereby converted into an oospore. 

In the roots of conifer seedlings oospores are formed not only 
in the cortical parenchyma but also in the interior of the tracheides, 
when, in consequence of the restricted space, they frequently 
assume an elongated form. 

The oospores reach the ground in the decomposing parts of 
plants, and there they may remain capable of germinating for at 
least four years. Some soil taken from a diseased beech seed- 
bed in 1875, having been distributed in water and poured 
upon a bed of seedling beeches, caused disease and death of the 
germinating plants not only in 1876 but also in 1878, and even 
in 1879. 

The practical measures at our command for combating the 
disease follow from what has been said. In order to guard 
against the outbreak of an epidemic, we must avoid sowing seeds 
on ground where the disease has once proved destructive, though 
we may cultivate transplants on it. The oospores that remain 
in the ground will prove destructive only to seedlings. Should 
the disease appear in a seed-bed, all contrivances for producing 
artificial shading must be removed, as they prevent the rapid 
evaporation of water from the surface of the cotyledons. All 
dead and visibly diseased plants should be removed. If a 
number stand close together, the distribution of the sporangia and 


gonidla may be most quickly prevented b}' heaping on earth. If 
the diseased plants occur singly they should be carefully pulled 
out and buried in a firmly trodden trench, in order to guard 
against the dissemination of the sporangia. In traversing the 
bed the spread of this disease ought to be obviated as much as 
possible b}- the workman not allowing his boots subsequently to 
come into contact with healthy plants. The seed-bed should 
also be inspected daily. 


Although the fungus which produces the potato disease had 
been introduced from North America before 1845, it is only 
since that year that it has assumed the dimensions of a plague in 
Europe, where it always causes great loss in wet years. In the 
mode of its distribution and in its dependence on wet weather it 
very closely resembles P. ovinivofa. A characteristic feature is 
the occurrence on the leaves of black blotches, w^hich, con- 
stantly increasing in circumference, and finally embracing the 
stems, may bring about the premature death of the parts of the 
plant above ground. Although the tubers of diseased plants are 
generally more or less affected, still this is sometimes the case 
only to a small extent, being recognizable merely b)- a few brown 
specks on cutting into the tuber. During wet years the tubers 
often rot for the most part in the field, those that are less 
attacked decaying in the cellar or pit during winter. These 
changes (wet- rot) are to a large extent due to the action of 

The mycelium of P. infcstans passes the winter in the tubers, 
and when these are planted out it grows into the sprouting shoots 
invading the tissues of both stem and leaf On examining the 
neighbourhood of the black blotches, one recognizes, even with 
the naked eye, a zone which is distinguished by its mouldy 
appearance. Here are to be found the numerous sporangiophores, 
for the most part projecting from the stomata. They agree 
in shape with those of P. omnivora, and bear similar but more 
numerous sporangia, which convey the disease to sound plants, 
and are even carried by the wind to adjoining fields. There is 
* [See footnote on p. 38. — Ed.] 


no doubt that they are also brushed off and distributed by ani- 
mals, as, for instance, by hares. The germination of the sporangia, 
or their production of swarm-spores as the case may be, agrees 
generally with that of the allied species. The sporangia, how- 
ever, reach the ground in large numbers, and are carried down to 
the tubers by the rain-water. Should the ground remain wet, in- 
fection follows upon the development of the germ-tube. The fact 
that varieties of potatoes with thin skins are more easily pene- 
trated by the germ-tube of the fungus than those having thick 
skins may explain how it is that the latter suffer less from 

The formation of oospores, which I have demonstrated in 
the case of P. oninivora, has not )'et been discovered in the 
potato fungus, and possibly it may not exist. Seeing that the 
mycelium passes the winter in the tubers, the existence of the 
fungus does not in this case depend on the formation of oospores. 
The occurrence and spread of the disease is most of all in- 
fluenced by the humidity of the air and soil, because, in a 
moist environment, sporangia are abundantly produced on the 
leaves, and the germination of the sporangia and gonidia, both 
above and below ground, is greatly favoured. 

If dampness prevail during storage in winter, numerous 
sporangiophores are produced on the tubers, especially in the 
region of the eyes, or where a wound ma}' happen to occur ; 
and by means of the sporangia that are formed the disease may 
be conveyed to previously sound tubers even at that season of the 


A decade or two ago this parasite of the vine was introduced 
from America, and in the interval it has rapidly spread through 
the vineyards of Europe. 

Its American designation of mildew, or grape-vine mildew, 
has been changed in France to mildiou. In Germany it is called 
the false mildew of the vine {falscher MeJilthmt der Reben). 

The disease is characterized by the occurrence of large grey 
patches on the under side of the leaves, while on the upper side 
the infested spots become yellow or reddish. The diseased 
spots dry up, and the leaves are shed prematurely. During 



rainy weather the disease spreads rapidly, but dry weather at 
once retards further progress. The fungus passes the winter 
in the form of oospores, which arc produced in the diseased 
leaves. During summer the distribution is effected by sporangia 
and zoospores, as in the case of PJiytopJithora. Infection takes 
place chiefly on the young shoots and leaves, the epidermis of 
which is but slightly cuticularized. The disease proves the more 
destructive to vines and grapes the earlier in the season it 
appears, and especially so when favoured by wet weather. 

It is not improbable that other species belonging to the 
genera Peronospora and PytJihuii do injury to young trees. It 
is especially desirable that investigations be instituted to prove 
whether PytJiiuvi de Baryanuni — which in crowded beds causes 
the death of many agricultural plants — is also injurious in the 
seed-beds of dicotyledonous and coniferous trees.* The genus 
Cystopus also belongs to the Peronosporcz, the best-known species 
of which is Cystopus candidiis, which produces the white-rust of 


Although this order contains only fungi which are parasitic 
on herbaceous plants, especially grasses, still the diseases which 
they produce are of sufficient importance to require a short 
description here. 

In the cvery-day language of the farmer, Smut is a term 
applied to the most varied phenomena of disease in plants. 
In the narrower sense of the word, however, the term is restricted 
to those diseases which produce a dark brown mass of spores 
in certain parts of plants, especially flowers and fruits, less fre- 
quently on leaves, stems, and even roots. In the particular part 
of the plant that is occupied by the copious mycelia of the 
smut-fungus this spore-powder is formed by the abscission or 
abjunction of abundantly developed fungus-filaments, the tissues 
of the plant itself being almost completely destroyed. 

The mass of spores is either formed on the surface of the 

* [An allied form was exceedingly destructive in seed-beds of Cinchona in 
Ceylon in 1880-1881.— Ed.] 

t [For the British forms of Ustilagines see Massee, op. at., and 
Plowright, British UredinecE mid Ustilagineoi (Kegan Paul & Co., 1889). — Ed.] 


plant or it remains enclosed by the epidermis, in which case 
it appears as a black semi-transparent swelling. 

The spores of smut may retain their capacity for germination 
for several years. On the recurrence of favourable conditions 
they usually produce a stout germ-tube called the promycclium 
[I'orkeivi), which, after attaining a length equal to two or three 
times the diameter of the spore, forms a number of smaller 
spores, known as sporidia, at its apex or on its sides. 

Frequently the promycelium breaks up directly into a number 
of sporidia. In the case of those species which produce \v'horls 
of sporidia at the apex of the promycelium, a process of fusion 
takes place between adjoining sporidia, and these afterwards 
drop off in pairs. 

When a germinating smut-spore or sporidium comes into 
contact with a suitable young host-plant, it sends its germ-tube 
through the epidermis, and thus gets into the tissues of the 
stem, where the mycelium grows upwards, chiefly intercellular 
without producing any apparent damage. It is only in the parts 
of the plant where spores are formed that the tissues are 

Those smut-spores which fall to the ground before or during 
harvest usually germinate at once, and perish in the absence 
of suitable young host-plants.* This being the case, the disease 
persists from year to year, for the most part, owing to the employ- 
ment of seed to which smut-spores adhere externally. When the 
corn is being threshed the detachment of the spores from 
smutted plants offers ample opportunity for the contamination 
of the seed-grain. Frequently, however, the spores are conveyed 
to the field in manure which has been made with smutted 

On account of the germination of the smut-sporcs being 
dependent in great measure on moisture in the air and soil, 
the occurrence of the disease is favoured in a soil whose 
physical condition — either naturally or owing to the liberal 
application of farm-yard manure — enables it to retain large 
quantities of water. 

It follows from what has been said that attention should first 

*[Brefeld has shown that the "sporidia" may reproduce by budding 
saprophytically during long periods in the manured soil. — Ed.] 

F 2 


be directed to preventing the transference of smut-spores to the 
field. To secure this, seed which is as clean as possible should 
be used. If this cannot be had, the adhering spores should be 
killed by steeping the seed-grain for twelve to sixteen hours in 
.a one-half per cent, solution of cupric sulphate. Further, the 
use of smutted straw for manure should be avoided. 

The most important kinds of smut (brand) are : — 

The Coal-brand, Stickv-brand, Stink-brand, or Bunt 
of wheat ( Tilletia Caries mid T. Icevis) — which besides attacking 
wheat is also found on quickens, wall barley, and meadow grass 
{^Poa pratensis) — is characterized by the fact that the spore- 
powder (which emits a disagreeable smell when fresh) remains 
enclosed in the grains till the time of harvest. The bunted 
grains being bruised in threshing liberate the spores, which 
adhere to the sound grains, and, both being sown, the young 
plants become infected. 

The Dust-brand [Ustilago) is the most destructive genus, 
and also contains the greatest number of species. Ustilago 
Carbo attacks not only oats, wheat, and barley, but also a large 
number of meadow grasses. It completely destroys the ovary, 
and usually the paleae as well, so that brown spore-powder 
escapes on to the stalk. 

Ustilago destruens, the Millet-brand, destroys the panicles of 
the millet while they are still enclosed by the highest leaf- 

Ustilago Ulaydis, the Maize-brand, produces large swellings, 
completely filled with dark brown spore-powder, on the stem, 
leaves, and cobs of the maize. Numerous other species occur 
on grasses, herbs, and bulbous-rooted plants. 

The Stem-brand {Urocystis) is frequently met with, and 
especially the brand of rye-stems, Urocystis occulta. It is very 
conspicuous, on account of the highest internode of the rye-stem 
rupturing longitudinally, and allowing the black spore-powder 
to escape. 

Other forms often met with are Urocystis J^iolcr, U. Ancnionis, 
and U. Cepul<£. 



This second group of fungi has obtained its name through the 
spores being produced in the interior of sacs (asci). In some 
cases the sporocarp results from a sexual process.* The fungi 
belonging to this group are very numerous, and are arranged 
in four orders — the Erysiphecs, Tiiberacecs, Pyrenomycetes, and 


All the mildew fungi are true parasites. Their mycelium 
vegetates on the surface of plants — that is to say, on the 
epidermis of leaves, fruits, and stems, and obtains its nourishment 
by means of haustoria from the interior of the epidermal cells, 
which consequently turn brown and die. The ascocarps which 
are developed on the mycelium are usually globular and com- 
pletely closed — that is to say, unprovided with an apical or other 
opening,-]- and may be recognized with the naked eye as small 
dark specks. These hibernate and carry the fungus over to the 
following year, while in the course of the summer gonidia are 
formed by abscission on numerous simple erect hypha:?. These 
are at once capable of germinating, and spread the disease during 
the period of growth. On account of the interwoven mycelia and 
gonidiophores, when luxuriantly developed, forming a fine grey 
meal-like covering on the upper surface of the leaf, the term 
" Mildew" has been applied to the disease. 

As a preventive measure, the burning in autumn of leaves 
infested by the cleistocarps of the fungus has been recommended, 
while sprinkling sulphur on the diseased parts after the mildew 
has appeared in summer is said to be efficacious. Unfortunately 
no scientific investigations regarding the action of the powdered 
sulphur on the mycelium of the fungus have as yet been 

*[The Sporocarp (in this case termed an Ascocarp) is often a very com- 
plex body. The question as to its origin and morphological nature cannot 
be discussed here, and the reader is referred to the special works of De Bary, 
Brefeld, &c., already quoted. — Ed.] 

t [These closed Ascocarps are termed Cleisiocarps, in distinction from 
the perforated Perithecia of the Pyrenomycetes and the open Apoihccia of 
the Discomycetes. — Ed.] 


The numerous species of mildews have recenth' been arranged 
in several genera according to the number of asci in the 
cleistocarp, or according to the number of spores in the ascus, 
or, finally, according to the structure of the so-called appendiculae, 
which are peculiar filiform radiating processes of certain cells 
of the wall of the perithccium. Here we need only allude to 
a few species. 

ErysipJie {Phyllactiiiia) guttata forms the mildew of Fagus, 
Carpimis, Cory his, Qiierais, Betula, Almis, Fraxinns, Loniceray 
Pynis covimimis, and Cratcrgiis. The cleistocarps are fur- 
nished with appendiculae which are straight, unbranched, and 
thickened in a bulbous manner at the base, and internally 
produce several asci, each containing two spores. In beech woods 
this parasite sometimes causes premature withering of the leaves. 

ErysipJie bicornis {Uncinnla Accris) very often injures the 
leaves and young shoots of Acer. I have encountered this 
species most frequently on Acer platanoides and A. campestre. 
It covers the whole of the leaf, or forms large greyish white 
blotches on one or both sides (the black patches on these leaves 
are due to Rhytisma acerimuii). The cleistocarps possess several 
asci holding eight spores, and the appendicular are simply forked 
at the apex. The gonidia are elliptical in shape. Even so early 
as August the leaves of the maple are often completely covered 
with these white patches. 

ErysipJie Ttdasnei is closely related to the former species, 
but occurs only on the upper side of the leaves of the Nor- 
way maple. The gonidia are globular. ErysipJie {Uncimild) 
adunca produces the mildew of the leaves of willows and 

ErysipJie {SpJuvrotJieca) pannosa forms the well - known 
mildew on the shoots and leaves of the rose. In w^et years 
especially it is necessary prompth' to pluck and burn the 
diseased leaves. 

Oidium Tiickeri, the fungus which causes the disease of the 
grape, was observed in England for the first time in 1845, but 
has since spread throughout all the vine-growing countries of 
Europe. The mycelium grows on the leaves, shoots, and fruit. 
When the last is attacked the epidermis dies and loses the 
power of expansion, so that as the berry grows the epidermis is 


ruptured, and the grapes in consequence begin to decaj'. So far 
only the gonidia have been discovered, and it remains to be 
determined how the fungus survives the winter. 


The Truffles are distinguished by having round subterranean 
closed fructifications (cleistocarps), in which the asci are produced 
on hymenia which clothe the surfaces of contorted passages. 
Gonidia and sexual organs are unknown. 

Through the investigations of Rees^ it was first established 
that the stag truffle, ElapJiomyces granulatiis, develops its 
mycelium parasitically on the roots of pines. It is further 
known that the edible species of truffle of the genus Tuber are 
parasitic on the roots of the oak and beech. Frank has re- 
cently devoted much study to the occurrence of fungal growths 
on the roots of phanerogamic plants, especially ConifercB and 
Cnpulifcnr, and has proved that mycelial growths are ^^^idel)' 
distributed on the tender apices of the roots of trees. The outer 
surface of young roots may be so closely covered by the 
mycelium, which penetrates into and between the cells, as to 
form a dense fungal mantle. Owing to luxuriant branching 
and growth of the tissues the infested roots to some extent 
display abnormal forms, while a sort of symbiotic condition 
arises similar to what we find associated with many other plant- 
parasites. When the cortical tissue of the roots has been 
infested by the fungus for some time, it dies, and should the 
fungal filaments penetrate into the internal tissues the roots 
themselves die off entirely. Frank has designated these 
phenomena by the name MycorJiiza, or fungus-root. It has not 
yet been determined how many species of fungi take part in 
these phenomena, and, especially, whether fungi belonging to 
other groups besides the Tuber acecB form MycorJiisa. Frank 
holds the view that these root-fungi, by assisting in nutrition 
and by conveying organic plant-food from the soil, play an 
important part in the life of trees.* 

^ Dr. M. Rees and Dr. K. Fisch, UntcrsiicJiiingeii iiber Ban tmd Lcbot 
der Hirschtrii^el^ ""^ ElapJiomyces^^ 1888. 

*[The best account of Frank's views for the student is in \i\% Lehrbiich dcr 
Botanik, B.I. 1893.— Ed.] 


Whether this view will receive confirmation in the future 
remains to be seen, but in the meantime its correctness is open 
to grave doubts. In the first place, it has not yet been proved 
that trees can take in organic food-substances by their roots ; and, 
in the second, it has been established that trees are very well 
nourished without the aid of MycorJiiza, and that, besides the 
infested roots, there is always a very large proportion of roots 
■entirely free from fungoid growth. 


In the case of Pyrenoinycetes the hymenium bearing the asci 
usually lines the inner surface of roundish or flask-shaped 
receptacles, called perithecia, which are distinguished from 
the cleistocarps of the preceding by having an aperture at the 
apex through which the spores escape. The numerous genera 
may be divided into two groups, according as the perithecia 
stand singly (simplices), or grouped in large numbers on a 
common cushion, or sunk in a stroma (compositi). 

The following species, being noteworthy parasites, deserve 
closer attention. 


This parasite is chiefly met with on the silver fir, though, 
according to v. Tubeuf, it also occurs on the common spruce 
and hemlock spruce. It is to be found wherever the silver fir 
is indigenous. Its colourless perennial mycelium grows on the 
under side of the branches, from which it spreads to the under 
side of the leaves, knitting them firmly to the branches. On 
this account the leaves, instead of falling off on dying, remain 
attached to the branches (Fig. i8). 

On account of the mycelium being confined to the lower side 
of the branch, most of the leaves that are met with on the upper 
side survive during the first year at least (Fig. i8, a). The my- 
celium encroaches on the new shoots as they are formed, and, 

1 R.H3.xt\g, Ein 7ieuer Parasit der IVezssfanuc, " Tridiospharia parasitica:'' 
AUgem. Forst- tend Jagd-Zeitg.^ January 1884. 



the young immature leaves at the base of the shoot being killed, 
subsequently shrivel up. The leaves on the middle and apex 
of the shoot, being 
reached somewhat later 
by the slowly advanc- 
ing m)'celium, retain 
their shape. 

The cushions which 
are formed by the my- 
celium on the under side 
of the leaves are at first 
white, but afterwards turn 
brownish (Fig. 19, b b). 
They only partially con- 
ceal the bluish lines 
which are met with on 
the under side of the 
leaves of the silver fir. 
In the course of time 
very minute perithecia 
are formed on these 
cushions (Fig. 20). 

The cushion originates 
in the following way. 
From the hyphae that 
cover the leaf (Fig. 21, a) 
numerous branches, (^, are 
sent out towards the 
epidermis, and these 
form a fleshy cushion, <r, 
consisting of parallel 
hyphae closely united 
to each other. At the 
point, d, where it reaches 
the epidermis of the leaf, 
each hypha sends a fine 

rod-like haustorium into the outer wall, e, of the epidermal 
cells, and owing to the secretion of a ferment these cells and 
the stomata, f, are killed and become brown. The cells of 

Fig. 18. — Branch of the silver fir attacked by 
Triihosplueria parasitica, a, healthy leaves ; 
h, dead and brown leaves whose bases are at- 
tached to the branch by the fungus-filaments. 
On account of their not being fully formed when 
attacked by the fungus the dead leaves towards 
the base of the shoot have shrivelled up. 



Fig. 19. — Lower side of a 
leaf of the silver fir at- 
tacked by T. parasitica. 
At a the colourless my- 
celium spreads from the 
branch on to the lower 
side of the leaf, on which 
it forms white cushions, 

Fig. 20. — Part of the leaf 
of a silver fir, on the left 
side of which the cushion 
bears a number of small 

Fig. 21. — Mycelial cushion of T. parasitica on the under side of the leaf of a silver 
fir. The filamentous mycelium a sends down numerous branches at d to form 
a cushion c, consisting of parallel hyphre ; where the hyphre reach the surface of 
the leaf each sends a rod-like haustorium, c/, into the outer wall of the epidermal 
cells, e e ; at dlhe cushion has been somewhat raised from the leaf, so that several 
of the haustoria have been pulled out of the epidermis; the epidermal cells yy 
have become brown. Although the filamentous mycelium // has penetrated the 
chlorophyllous cells of the leaf-parenchyma^^, these do not become brown till 
somewhat later ; the mycelial cushion grows into the depressions at the entrance 
to the stomata i, where however it is unable to form haustoria ; at these places 
it becomes coated with the adhering waxy granules. 



the interior, ^- which contain chloroph)'ll, do not succumb for 
some time to the action of the mycelium, //, which here and 
there effects an entrance. The depression at the entrance 

to the stomata, being Hned with 
wax)' granules, prevents the en- 
trance of any haustorium, /. The 
dark brown perithecia (Fig. 22) 
which ultimately arise on the 
cushion are scarcely recognizable 
with the naked e}-e. The}^ are 

Fig. 22. — Perithecium of T. para- 
sitica. The dark brown sphere 
shows a round aperture at its 
apex, and bristle-like hairs 
which jiroject from its upper 
half. A portion of the wall 
has been removed from the 
lower left-hand side in order to 
show the pale contents, which 
consists of asci and paraphyses. 
These are shown more highly 
magnified in the lower part of 
the figure ; a representing rod- 
like bodies which are often pre- 
sent ; b, asci with spores, and 
r, isolated spores. 

Fig. 23. — Herpotrichia nigra on the 
spruce ; half natural size. 

characterized by having bristle-like hairs distributed over the 
upper half In the interior of the perithecia are often to be found 
small rod-like organs, a, besides the asci, b, which hold eight 
grey spores, usually consisting of four chambers. Should these 
spores succeed in obtaining a suitable footing on the branch 
of a silver fir, they speedily germinate and produce the disease. 
The m}-celium spreads parasiticalh- from the point of infection 


in all directions, so that large branches may be eventually entirely 
defoliated. In dense young woods it even spreads from branch 
to branch, while fresh centres of disease are produced by the 
distribution of the spores. 

Seeing that young woods which have been naturally regener- 
ated may suffer severely — and especially so where they have been 
formed under shelter trees — it is desirable that diseased branches 
should be pruned off This treatment has produced good results 
where practised on a large scale. 


..^X^\/^^^{ C^^ This parasite is met with in the higher mountain ranges, where 
it chiefly attacks the spruce, mountain pine, and juniper. In the 

jjN^j^j^ijNk^woods of mountain pine, large blanks are met with, which, on a 
'^ cursory glance, give one the impression of having been completely 

^v.gJL'., charred. In nurseries at high elevations, where the young spruces 
are buried under snow during winter and spring, it often happens, 

'^^'^^ directly after the snow has melted, that the plants are overgrown 

—' and killed by the dark brown mycelium. This is especially notice- 

, ^ able when the young trees have been laid prostrate on the ground. 

In the spruce woods of the Bavarian Forest one often finds 
that the fungus has killed the young seedlings over large areas 
either entirely or to the height of twelve or fifteen inches. The 
dark brown mycelium envelops the whole branch or plant, knitting 
the leaves completely together (Fig. 23). 

Instead of forming a definite cushion, the mycelium embraces 
the leaves irregularly (Fig. 24, /;), and on these the perithecia 
are also produced {a). Dark brown tuber-like bodies are formed 
over the stOmata (Fig. 25), while the mycelium also spreads 
over the surface of the leaf and sends haustoria into the outer 
walls of the epidermal cells, which consequently die and become 
brown. The deeper-lying parenchymatous cells are also killed 
by the fungus, even before any mycelial threads have gained an 
entrance through stomata on other parts of the leaf, and pene- 
trated into the interior. 

The surface of the dark brown comparatively large perithecia, 

^ R. Hartig, Hcrpotrichia m\n-a n. sp. All. Forst- und Jagd-Zeitg., 
January 18S8. 


Fig. 24, is beset with numerous branching h)-pha:, which are 
specially abundant on the lower part, near the point of contact 
with the mycelium. These black spheroid bodies are frequentl\^ 
nearly hidden by the mycelium. In the asci the spores arc 
arranged in two rows. At first, and apparently also when 
mature, thc}^ consist of two chambers, but at last four chambers 
are formed. These spores germinate with great readiness. 

It is an interesting biological point that the fungus grows, es- 
pecially when the temperature is low, 
under the snow or during the time it 
is melting, because, under such cir- 
cumstances, the air is completcl}^ 
saturated with moisture. The fre- 
quenc}' of the disease at high eleva- 

FlG. 24. — a and /', two spruce 
leaves attacked by H. nigra, 
twice natural size. The brown 
mycelium forms black tuber- 
like bodies in the stomata, 
which however are much 
smaller than the black peri- 
thecia, one of which, magni- 
fied fifty times, is shown in 
the lower part of the figure. 

Fig. 25. — The growth of the mycelium 
of H. nigra. I'he filamentous my- 
celium a develops a granular mycelium 
on the surface of the leaf, and this 
covers the stomata with tuber-like 
bodies, rod-like haustoria being sent 
into the outer walls of the epidermal 

tions has led to the general adoption of the practice of forming 
spruce nurseries at low altitudes. It has also been found a 
good plan to look over the nurseries immediately after the 
melting of the snow, and to raise up all prostrated plants in 
order that they may be exposed to the wind. It would also 
be a step in the right direction, in planting out trees, to set 
them on hillocks and similar elevations, and to avoid placing 
them in hollows and other depressions. 



The oak-root fungus, Rosellinia qiierciiia, is one of the most 
interesthig of parasites, and especially so because its mycelium 
displays the same diversity of form as that of Agaricus inelleiis. 
The mycelium is one of those parasitic mycelial forms which 
were formerly referred to a special genus, RJiisoctonia. 

The disease produced by Rosellinia qiiercina appears only to 
attack the roots of oaks from one to three years old, but it is 
very prevalent, especially in north-west Germany. In oak seed- 
beds it gives indications of its presence by the young plants 
becoming pale and withered, especially during rainy seasons. 
The leaves near the apex of the shoot are the first to wither, but 
later on the lower ones go too. If a plant showing the first 
.symptoms of the disease be lifted out of the ground, we per- 
ceive a few black spheroid bodies of the size of pin-heads situated 
on the tap-root, especially at the points where the delicate 
lateral rootlets are met with (Fig. 26). It is also observed that, 
at certain points, the roots are closely embraced in a web-like 
fashion by delicate ramifying strands which resemble so many 
threads. These are the Rhizoctonice, which penetrate also into 
the adjacent soil, and, as we shall see, spread the disease 
underground from root to root. In the neighbourhood of 
these black tubers, and wherever the RhisoctonicB have been 
closely in contact with the surface of the roots, the cortex 
turns brown. The apex of the tap-root is often quite 
rotten, but even plants whose roots remain alive to the tip 
display the pathological symptoms already described. 

Upon older plants that are already dead the RJiizoctonicE are 
no longer white, but brown, and there the black spheroid bodies 
are often to be recognized in large numbers. Sometimes the 
latter are also to be found on the lower part of the stem — that is 
to say, on the epicotyledonary axis : they may be most easily 
discovered after the plant has been very carefully washed, be- 
cause then the lustre of the black tubercles readily betrays their 
presence. During damp warm weather all the plants on patches 
a yard or so in diameter may become withered and die. 

^ R. Hartig, Untersuchiingen aus deni Forstbot. Institute I. pp. 1—32. 



Where the seed has 
been drilled, the disease 
spreads from the point 
of attack in two direc- 
tions ; where sown broad- 
cast, it spreads centri- 
fugally in all directions. 
Should dry weather in- 
tervene, or on the ap- 
proach of autumn, the 
disease ceases to spread, 
but on examining the 
roots of plants appa- 
rently sound situated in 
close proximity to those 
already dead one will be 
able to recognize nume- 
rous examples of the 
pathological symptoms 
which have been already 
indicated. If such dis- 
eased plants are trans- 
planted in the following 
year, it will be a question 
of weather whether they 
die, and possibly trans- 
mit the disease to neigh- 
bouring trees, or form 
a new tap-root, if the 
apex was destroyed by 
the disease, and, after 
remaining stunted for 
some years, slowly re- 

If a dead plant be 
placed in a damp warm 
chamber, or be planted 
in July in the middle of 
a bed of healthy young 



seedlings a few months old, a mycelium is very soon developed 
from the black tubers — which w^e may call resting-mycelia 
(Sclerotia) — which breaks through the bark at different places 
and forms a dense whitish-grey mildew-like tissue, and also 
spreads rapidly over the surface of the ground (Fig. 27). This 
mycelium consists of septate hyphas, which are at first 
colourless but afterwards turn brown. These, after a time, 

arrange themselves side by 
side, grow together laterally 
at places, and form the 
fine strands called RJiisoctonice, 
which consist of numerous in- 
dividual hyphse very loosely 
united to each other. Should 
such a mycelium — whether in 
the form of isolated hyphae or 
of RhizoctonicB — come into con- 
tact with the sound roots of a 
neighbouring plant, it embraces 
Y/r >)> them in its meshes, and bores 

mi-' A''- directly into such of the corti- 

cal cells as are still alive. 
These are found in the delicate 
lateral rootlets and near the 
apex of the tap-root. The my- 
celium penetrates as far as the 
medulla, should such be pre- 
sent, and in a short time kills 
the root. In the living cortical 
parenchyma of the tap-root — 
which is only to be found on 
the lowest and youngest parts— the cells become plugged up 
with a luxuriant growth of pseudo-parenchymatous tissue, 
which, owing to the occurrence of numerous oil-globules, 
may be recognized as a resting-mycelium. These bodies, which 
germinate under favourable conditions, may be designated cham- 
bered sclerotia. On account of the formation of a periderm layer 
in its cortex, the older parts of the tap-root are protected against 
the direct attack of the parasite. The outer cortical cells being 

Fig. 27. — Oak root enveloped liy the 
mycelium of /v. qiierciiia, a, on which 
perithecia have developed at /'. 



parti}- shrivelled up and partly cast off, there remains but one 
path of entrance into the interior of the root. After the fine 
lateral roots that pierce the corky covering have been killed by 
the parasite, openings or breaches are left at certain points, 
through which the parasite 
effects an entrance, and this 
it accomplishes in a pecu- 
liar manner (Fig. 28). At 
such places — frequently both 
above and below the base 
of the dead root — fine white 
mycelial outgrowths are first 
formed. These develop into 
fleshy tubers, ultimately pos- 
sessing a dark brown cover- 
ing, which send several fleshy 
processes into the tissues of 
the oak root (Fig. 28, c, d). 

The adjoining cortical tis- 
sues are killed and become 
brown (Fig. 28, e). Should 
dry or cold weather inter- 
vene, the host-plant gains 
time to form a new layer of 
cork in the neighbourhood 
of the infecting tubers along 
the line that marks the limit 
of living tissue. In this way 
the plant may be, for the 
time, saved. Should the con- 
ditions of growth remain 
favourable for the fungus, 
however, the fleshy pro- 
tuberance pushes out a fine 

Fig. 28. — Point of infection by R. qtiercina, 
magnified twenty times. The delicate 
lateral rootlet a, which has been killed by 
the filamentous mycelium, displays fleshy 
infection-tubercles, b c, at the place 
where it has ruptured the periderm 
of the tap-root. These tubercles send 
processes, d, into the internal tissues. 
The adjoining cellular tissue is brown, e, 
but free from mycelium. A Rhizoctonia- 
strand, f, has been produced by the 
upper tubercle, which has consequently 
parted with a portion of its nutritive 

filamentous mycelium, which 

spreads through all the tissues of the root and kills it. 

In the sclerotia the parasite possesses a means of existing 
from one year to another, and of resisting the periods of drought 
during summer which kill all filamentous mycelia as well as 



the sporophores that may be in process of development upon 

In summer the mycelium that vegetates on the surface of the 
ground produces gonidia on the whorled branches of gonidio- 
phores, and these, by being carried on the skins of mice, &c., 
may originate new centres of infection. But besides these, black 
spheroid perithecia, about the size of a pin-head, are produced 
either on the surface of the diseased oaks themselves or on 
the surface of the ground in their neighbourhood (Fig. 27, b). 

It is probable that the spores which are formed in the pere- 
thecia do not as a rule germinate and reproduce the disease 
till the following year. 

Generally speaking, it is only in wet years that the parasite 
does much damage. It may be combated by digging trenches 
round the diseased spots in the seed-bed so as to isolate them. 
One should avoid transferring diseased plants from the seed- 
bed to the plant-bed. 

RJiizoctonia violacea, which kills saffron and lucerne, has not 
yet been scientifically examined in its different stages of develop- 
ment, and it remains to be determined whether this parasitic 
mycelium belongs to a form related to one of the foregoing fungi 
or not. Fuckel's statement that this mycelial form belongs to the 
fungus ByssotJieciuui circinnans appears so utterly improbable 
that it is not worth while to take further notice of it. On the 
other hand, I feel called upon to describe here the following 
important parasite of the vine. 


Amongst the numerous enemies of the vine, the root-fungus, 
D. necatrix, occupies a prominent position. The disease which it 
induces is known as Wurzelpilz, Weinstockfaule, Pourridie de la 
Vigne, Pourriture, Blanc des Racines, Blanquet, Champignon 
blanc, Aubernage, Mai nero, and Morbo bianco, and is dis- 

•^ R. Hartig, Dematophora necatrix Ji. sp., Untersiichiingen aus deni 
Forstbot. Institut zu Miinchen, III., 1883. 

*[Viala has published a very thorough investigation of this disease and 
the devastations it causes in the south of Fiance, &c. : " Monographie dn 
Pourridie des Vigne s et des Arbres fniitiers," 1891.— Ed.] 


tributed throughout France, Italy, Switzerland, Austria, and 
south-west Germany, 

Amongst the root-diseases of the vine, that which is caused 
by PJiylloxcra vastatrix is generally known. Ver)' similar 
pathological symptoms occur on the stems of plants that are 
attacked by the vine-root fungus, and confusion often enough 

Whether Agariais vielleus also is injurious to the vine — -as has 
been maintained — I am not in a position to say, because, so far, 
no specimens have been forwarded to me in which the fungus 
could actually be identified. On the other hand, it appears as 
though, in very wet years and on heavy ground, " root-rot " may 
arise as a result of asphyxia — that is to say, owing to deficiency 
of air in the soil. On such suffocated vines a fungus, Roesleria 
hypog(£a, often occurs, which, it appears to me, is most probably 
saprophytic in character. 

The parasite that we are here discussing spreads in the vine- 
yards from plant to plant by means of its underground mycelium, 
so that w^e often hear of great damage being done. Other plants 
that are cultivated in the vineyards, such as fruit-trees, potatoes, 
beans, beet, and the like, also fall a victim to the fungus. During 
my investigations I found that the mycelium could at once kill 
young maples, oaks, beeches, pines, spruces, &c. 

On plants where the mycelium is vigorously developed, as in the 
case of the vine, Fig. 29, and the young maple. Fig. 30, it forms 
a luxuriant snow-white mass of a woolly or strand-like texture, 
which adheres to the outside of the plants, though it may also 
spread in the ground to long distances. Where this mycelium en- 
counters the fine fibrous roots of other plants, it kills them, and, 
at their base, bores into the interior of the larger roots, Fig. 3 1 a, 
spreading afterwards in their interior in the form of peculiar 
rhizomorphs. Fig. 32, and killing all the adjoining tissues. In 
the soft cortical tissues of the vine-root they retain their strand- 
like appearance, and by ramifying laterally and outwards they 
envelop the root in a network of strands, Fig. ^^'':). 

In structure these rhizomorphs are entirely different from those 
oi Agariais melleiis. In Fig. 34 I have represented somewhat 
diagrammatically the apex of one of these rhizomorphs, and refer 
for details to the description appended to the illustration. 

G 2 




Fig. 29. — A vine that has been killed by 
D. 7iccatrix, and afterwards kept for a 
long time in a moist chamber. The fila- 
mentous mycelium, a, assumes the char- 
acter of white Rhizodonia strands, /', 
which anastomose, c c. At rt^and e rhizo- 
morphs grow out from the interior. 

Fig. 30. — A sycamore infected by 
D. necatrix. The portion above 
ground is represented some four- 
teen days anterior to the rest. 
The plant is enveloped in the 
white woolly mycelium, a ; on 
the subterranean portion Rhizoc- 
touiiv consisting of dark my- 
celium, b b, are to be seen. Nu- 
merous sclerotia, c, project from 
the cortex. 



The branches of the rhizomorph directed outwards break 
through the cortex from within and form a new filamentous 
myceHum, which penetrates into the soil ; or, in other cases, they 
swell up under the cortex to form tuberous sclerotia. Fig. 33 (^, 

Fig. 31. — Longitudinal 
section of the root of 
a vine whose upper 
part has been killed 
by the rhizomorphs 
of D. necatrix as far 
as b, and whose lower 
portion shows an in- 
fection-spot at a. 

Fig. 32. — Magnified five 
times. Boundar}', a, 
of the healthy and dis- 
eased parts of the root. 
The rhizomorphs send 
out lateral branches, 
which may occasion- 
ally reach the epider- 
mis, as at /'. 

Fig. t,t,. — A large vine- 
root infected by D. 
necatrix. A portion of 
the cortex has been 
carefully removed so as 
to show the rhizo- 
morphs which begin to 
appear at a ; zX b the 
mycelial tubers, which 
resemble sclerotia, are 
formed, and on these 
the gonidiophores ulti- 
mately develop. 

which sometimes break through the cortex and appear in rows 
upon the surface, Fig. 35. 

On these tubers the gonidiophores are developed in the form 
of bristles, at whose apex the gonidia are abscinded, Fig. 36. 

It also happens very frequently, however, that these sporophores 



are developed on the filamentous mycelium which clothes the 
diseased plant, or foreign substances, in the form of RhisoctonicE 
or otherwise 

The perithecia of this species have been discovered by Viala. 

O C5 

t; .S c 

i: .- T^ ■z! (u ^ .. Q 

O rt ' 


St? f-' ^- o 

r^ >~, C t. rt ^ ^ 
5. t. oj -^ 

^ •£ ^ ^ C -« -tr; O ° ?J 

■ ° I ^ ^ >^ § ^ -a 

c 'jr s rp 

^^^ t. °A <r: ^ ii r. '^ ^ -9/^. V. 
£ S^ Sr^ ^ '■^ " ■ 

o -^ ^ 

.ti o 

^ .. S =J s "" "^ 

o *-■ 

u _ o 

J o p jr; '5 "P "" 

rv t/)>i^3r-.iiiii-ij-.t-.ti *^T? 


■5 ^'^'^^ S'2'O ^ lU 

,S! '3 ST! ^ ■? B rt ., ^ 'm 
ih r< o o m '^^ ■av. '" 'r. Y. "^ ^-^ 

■go's -S '"^ 

o rt fe rt C^- 

rt-:'SSP.S-S.yrt .- c^ 

ii o c ^ 


^ o 

-G rt ^ -r; ^ 

They form only on vines and fruit-trees that have been dead 
and decayed for a long period. It is only in soil that has dried 
slowly that they are to be found. Associated with bristling ap- 
pendages bearing gonidia the perithecia appear partly on sclerotia 



and parti}' on the mycelium. Between the surface of the ground 
and the depth of some two inches they form clusters of small 
spheroid bodies round the vine or tree. The sporocarps, which 
are very hard, deep brown in colour, approximately spheroid in 
shape, and 2 mm. long, are provided with short stalks. They 
are completel}- closed, 
and enable us to refer 
Deviatophora to the 
Tiiberacei. The elon- 
gated filiform asci, 
however, which swell 
out on one side like a 
lop-sided turnip to en- 
close the eight spores, 
distinguish this para- 
site from all known 
Tiiberacei. Deviato- 
pJiora necatrix is thus 
the sole representative 
of a new genus of 

The method which I 
at first recommended 
of starving the para- 
site b}- means of iso- 
lating trenches, &c., 
having proved too 

tedious a process, it remains to be seen whether the impreg- 
nation of the poles with creosote can do anything to combat 
the disease in the vineyards. 

Fig. 35. — Root of a vine 
showing numerous scle- 
rotia-like tubers, on 
which a few bristle- 
shaped gonidiophores 
have developed. 

Fig. 36. — A portion 
of Fig. 35 after 
the gonidiophores 
have been formed ; 
magnified five 



This parasite frequently gains an entrance through wounds on 
Cytisus Laburnum, destroying the cortex and branches for con- 
siderable distances, and even killing the whole plant. Besides 
the dark brown spheroid perithecia, which are arranged in groups, 

1 Cucurbitai-ia Labiirni, atif Cytisus Laburnum. Freiherr v. Tubeuf. 
Cassel, Fischer, 18S6. 



we meet with the most varied forms of gonidia, which are pro- 
duced either free on the stroma, or in the interior of cavities in 
the stroma, or in pycnida. Owing to the ease with which all 
these organs of reproduction germinate, the parasite is frequently 
very abundant. 

In a similar manner, C. Sorbi appears to attack the bark of 
Sorbits Auaiparia. 

Here allusion may be made in a few words to the " Disease- 
blotches " on the leaves of numerous trees, shrubs, and herbs. 
These often occur in great abundance in autumn, the leaves being 
covered by numerous sharply defined blotches, which are usually 
circular in outline and brown in colour, and frequently surrounded 
by a red margin. They are generally due to fungi belonging to 
the family Sphczrelloidea, and especially to the genera SpJuErella 
and Stigma tea. 

The gonidia are formed on the living leaves, but the perithecia 
only on the dead parts of plants, and usually not until the spring 
after the leaves have fallen. 

SpJicBvella FragaricB produces the diseased blotches on the 

S . piinctiforniis and .S". viaailiforviis produce brown blotches 
on the leaves of the oak, lime, and hazel. 

5. Fagi produces blotches on the leaves of the beech, &c. 

Stigniatea Mespili induces brownness in the leaves of the pear. 

Stigmatea Alni is the cause of blotches on the leaves of the 

Gnomonia belongs to an allied family, Gnonionia erythrostoma, 
producing a brown colour in the leaves of the cherr}'. The in- 
fected leaves die prematurely, but do not fall off. On these are 
developed the perithecia with their unicellular tubular spores. It 
is advisable to remove during winter all leaves that may be 
hanging on the trees. 

A parasite, Valsa Prunastri, frequently proves injurious to the 
apricot, cherry, and sloe. The fungus infests the cortex, and 
causes the death of the branches. The form producing spermatia 
is the first to appear, and ejects its tendril-like masses of sper- 
matia ; while later — namely, in the following spring — the peri- 
thecia develop in the dead cortex. 



The genus Nectria contains a number of parasitic fungi which 
produce their perithecia — which are usually red, and grouped in 
considerable numbers — on the surface of a pseudo-parenchymatous 
wart-like stroma. Before the perithecia make their appearance, 
this same stroma serves for the production of numerous gonidia. 
The gonidia-bearing stroma was formerly referred to a special 
genus called Tnbcrcularia. 

The following three species belonging to this genus are faculta- 
tive parasites, which, like so many other parasites, can also live 
as saprophytes. 


Like all the Nectrias, ^V. Cucurbit2ila is one of those parasites 
which, as a rule, can gain access to the interior of a host-plant 
only through a pre-existing wound. Here the host-plant is 
usually the spruce, in rarer instances the silver fir, Scotch pine, 
&c. The means of entrance which the parasite utilizes in the 
forest are chiefly the injured spots due to GrapJwlitha pactolana, 
Fig. 37, though, less frequently, it also enters through the abra- 
sions caused by hail, or the crack at the base of a branch whose 
bark in the upper angle has been slightly torn by the depression 
due to an accumulation of snow\ 

The germinating ascospores or gonidia push their germ- 
tubes into the tissues of the cortex, and the ramifying mycelium 
ultimately develops most luxuriantly in the sieve-tubes of the 
soft bast, Fig. 38 b, or in the intercellular spaces between them, 
Fig. 1% c. The mycelium is met with in bast tissues that are 
apparently perfectly sound and fresh. The brown colour does 
not appear in the tissues for some time afterwards. The 
fungus would appear to make progress, for the most part, only 
when growth is at a stand-still in the cortical tissues. It 
generally ceases to advance when the plant and its cambium 
awake to renewed activity. From this we must assume that 
the power of resistance of the living tissues of the host-plant is 

' R. Hartig, Untersuchiingen^ I. p. 88. 



greater during the season of growth than at other times. As 
ma}- be seen from Fig. 'i^'j, the parasite may advance longitu- 
dinally more than 2\ inches 
during a growing season. Later- 
ally the seat of the disease 
seldom advances more than 
I — \\ inch. The tissues that 
have been killed b}' the fungus 
become separated from the 

Fig. 37.— a spruce attacked by N. 
Ciiiiirbitida. At a a wound due to 
a hailstone has healed over without 
becoming infected ; l>, the gallery 
of a larva of GraphcHtha pactolana, 
over which a callus has been formed, 
but where infection has occurred 
two years later : the mycelium has 
spread from c to c in the cambium, 
and from d to d in the cortex ; 
numerous groups of perithecia have 
appeared on the dead cortex. 

Fig. 38. — Cross section of the 
cortex and wood of a spruce 
infected a short time previous- 
ly ; a, the wood. /' b, the sieve- 
tubes containing one or more 
mycelial filaments ; c, my- 
celium in the intercellular 
spaces ; magnified 420 times. 

living parts by the formation of a laj-er of cork which prevents 
the further progress of the parasite in the following year. 

If the cortex that has been killed be exposed to wind and sun, 


it dries up even so earl}^ as the beginning of summer. When 
the part of the tree that is attached is thin, the wood also dries 
up, and the top of the tree dies and becomes yellow and 
withered. Ver)' frequently one meets with such withered tops 
in young spruce woods without being able to find a trace of the 
perithecia, which only attain maturity when the cortex in which 
the mycelium is hidden is constantly kept moist. When this 
occurs — as often happens on the lower parts of the stem, where 
the cortex is kept moist by the shade and protection of the 
branches — a large number of white and yellow cushion-like 
stromata develop on the dead tissues. These, which are about the 
size of pin-heads, break through the outer cortical and periderm 
layers, or remain hidden amongst the loose bark scales. These 
cushion-like stromata first of all produce large numbers of gonidia, 
but later on numerous red melon-shaped perithecia are formed, 
whose ascospores are usually disseminated in winter or spring, 
when they find their way to the injuries caused by G. pactolana, 
or to other wounds. 

With the disappearance of the moth — as, for instance, in con- 
sequence of the severe winter 1879 — 80, in which the caterpillars 
were, for the most part, frozen — the injury due to this Ncctvia is 
of course also diminished, because it has fewer opportunities for 
infection. Spruces which are attacked only by the moth and not 
by the fungus hardly ever perish, but after being crippled for a 
few years recover completely. Spruces which are attacked by 
Nectria only on one side may also recover, because in the course 
of time a callus forms over the injured part. The damage, how- 
ever, which is done to young spruce woods by the trees dying at 
the top is so great that it seems advisable to limit the spread of 
the parasite by cutting off and burning all such tree-tops as 
are attacked by the fungus. 


It is the dicotyledonous trees that are chiefly attacked by this 
fungus, many of the varied forms of disease which are usuall)' 

^ R. Hartig, Ufttersicchtmgen, I. p. 209, Plate VI. 

* [This fungus is very common in this country, and I have frequently 
observed and examined its undoubted connection with the canker of apple 
and other trees. — Ed.] 


embraced under the term '" Canker " being due to iV. ditissiiiia. 
This canker-fungus appears most frequently upon the beech, oak, 
hazel, ash, hornbeam, alder, maple, lime,apple, dogwood, and bird- 
cherry. Although as a rule this parasite only gains an entrance 
to the cortical tissues of trees through wounds, I have also been 
able to infect young leaves by means of gonidia and ascospores. 
Abrasions caused by hailstones are probably the commonest kind 
of wounds, Fig. 39. Should no infection supervene on such a 
wound, a callus forms, and occludes the injured part in a short time, 
^^%- 39 '^^- If. however, it is infected by the gonidia or ascopores 
oiNectria, death and brownness spread in all directions, but most 
rapidly in the direction of the long axis of the stem. Although 
in rare instances the mycelium may advance 3 cm. in a year, it 
is comparatively seldom that the annual rate of progress in any 
direction exceeds one third of that amount. The apparent 
deepening of the diseased spot in the course of time is to be ex- 
plained from the fact that not only does the contiguous healthy 
tissue continue to increase in thickness, but it even displays an 
augmented rate of growth. This is satisfactorily enough ex- 
plained when we remember that during their movements in the 
bast the plastic materials assimilated by the leaves are neces- 
sarily confined to the sound side of the stem. As the canker- 
spot dries up, their passage is chiefly confined to its margin, which 
is consequently very richly nourished, and projects as a well- 
marked prominence. Thus in the course of years very striking 
malformations are formed. 

It also frequently happens that the base of a lateral branch, 
whose cortex has been injured in the upper angle, proves the in- 
fection-spot from, which death of the tissues annually proceeds. 
Fig. 40. In the case of the hazel especially it often happens that 
in pulling down the branches to get at the nuts a split is formed 
at the point of bifurcation. This then develops into a canker- 
spot, which constantly increases in size, as is represented in 
Fig. 41. 

I believe that I am justified in assuming that under certain 
circumstances, with which I am not yet familiar, the mycelium 
spreads from the cortex to the wood, in which it progresses up- 
wards, and at certain places attacks the tissues of the cortex and 
cambium from within. In this way canker-spots may be pro- 


duced without the part of the tree on which they occur having 
been previously injured, Fig. 42. The famihar state of things 
where certain trees are covered with canker-spots, while adjoining 

trees of the same spe- 
cies are tolerably clear 
of them, can hardly be 
explained in any other 
way than by assuming 
that the fungus travels 

Fig. 39. — Branch of 
a beech showing 
two hailstone 
wounds, of which 
the upper one, b, 
has been infected 
by N'cctria, while 
the lower one, a, 
has escaped infec- 
tion and has been 
occluded by a 

Fig. 40. — Hornbeam 
infected by N. ditis- 
sima, which has en- 
tered at the angle 
formed by the branch 
and the stem ; na- 
tural size. 

Fig. 41. — Hazel showing the 
canker due to N. ditissima, 
the spores of which have 
germinated in the bifurca- 
tion of two branches which 
have been somewhat pulled 
asunder ; a, b, h, the bound- 
ary of the canker-spot, 
where red perithecia are 
abundant ; c c, the healthy 
side of the branch ; half 
natural size. 

in the wood. This subject, however, requires further inves- 

As the mycelium spreads in the cortical tissues of trees, it 
produces innumerable extremely minute gonidia resembling 
bacteria, which apparently assist in no small degree in the 



almost complete decomposition of the tissues of the cortex, with 
the exception of the outer periderm layers. Only on those 
parts of the cortex which have been killed during the past year 
— that is to say, on the periphery of the 
canker- spot — do white gonidia-bearing stro- 
mata appear. These had already been ob- 
served by Willkomm in his investigations 
into the canker of the beech, by whom they 
were designated Fiisidmm candidiim. On these 
the minute deep-red perithecia originate, but 
they can only be discovered after careful 
search. They are found partly in groups and 
partly singly on the dead cortex, and especially 
in the fine fissures. Fig. 40. One sometimes 
searches for them for a long time in vain upon 
the older canker-spots, for the reason that 
these have ceased to increase in size at all parts 
of their circumference. In Fig. 43 the canker- 
spot is increasing in size only in the upper 
left-hand corner, and it is only there that the 
red perithecia are to be found in abundance. 

In the case of the canker of the beech, I 
have frequently observed that sooner or later 
the mycelium ceases to advance at certain 
places, in consequence of which the shape of 
the canker-spots becomes exceedingly irregu- 
lar. Here and there the canker spreads for a 
series of years, but finally the diseased spots 
may be entirely covered over by a kind of 
callus. See Figs. 43 and 44. 

It may also be remarked that the parasite 
is distributed throughout the whole of Ger- 
many, and that canker of the beech especially 
is met with from the Island of Riigen to the 
south of Bavaria, being very prevalent, for 
instance, in the neighbourhood of Munich. 
Young trees from five to ten years old, as well as trees 140 years 
of age, may be attacked by the disease, which, however, in the 
latter case, is confined to the twigs and branches of the crown. 


42. — Branch of 
a beech showing 
numerous canker- 
spots, which do 
not appear to have 
been preceded by 
any cortex wounds. 


Climatic conditions, especially frost, are essentially without 
influence on the disease, and the same is true as regards the soil. 
Although the damage inflicted by this parasite is by no means 

small, still I doubt if, in practice, 
anything can successfully be done 
to combat it. The injured trees 
remain alive as a rule, and at least 
yield firewood. Their removal in 
the thinnings is certainly advisable, 
so long as the ground is not there- 
by injuriously exposed. In oak 
woods also, whenever thinning and 

Fig. 43. — A stem of the beech, 
half natural size, showing numer- 
ous canker-spots ; these, how- 
ever, are increasing in size only 
at certain places, and it is only 
where increase is taking place 
that the red perithecia are to be 

Fig. 44. — Cross section taken from the 
lower end of the beech stem repre- 
sented in Fig. 43 ; natural size. 

opening out are undertaken with a view to underplanting, the 
cankered trees should be the first to be removed. I cannot advise, 
however, that one should go so far as to fell all cankered trees, if 
this should mean the formation of large blanks in the wood. 


Very frequently N. ditissiina is found associated with ApJiida} 
LacJinus exsiccator produces large galls in the cambium of the 
beech, and when these afterwards burst open they present the 
opportunity of infection to the fungus. The mycelium spreads 
in the cellular tissues with extraordinary rapidity. The beech 
Aphis, too, Chermes fagi, which clothes the stem with a \\h.\\.e 
woolly covering, is often associated with the fungus, which, under 
these circumstances, quickly kills the cortex, without producing 


This Nectria is certainly one of the most widely distributed 
of fungi, and finds its way on to almost all dicotyledonous trees 
and shrubs when they have been killed by frost. Besides living 
saprophytically, it also occurs as a parasite, and that most fre- 
quently on the maple, lime, and horse-chestnut. Infection usually 
occurs at branch-wounds, though also very frequently at root- 
wounds, which cannot be avoided when transplanting either large 
or small trees. The mycelium of this fungus, which grows rapidly 
upwards in the vessels, penetrates into all the elements of the 
wood, decomposing the starch, and leaving a green substance be- 
hind. Fig. 45. The consequence is that the wood turns black, 
while the cambium and cortical tissues remain sound. The wood 
becomes unable to conduct sap, the leaves wither prematurely in 
summer or drop off, and the cortex of the youngest shoots 
dries up after the wood is completely dead. In autumn or the 
following spring the cinnabar-coloured stromata that bear the 
gonidia appear grouped together in large numbers on the dead 
cortex. On account of their size and colour they are conspicu- 
ous even from a distance. The large rough perithecia which are 
formed later are much darker red in colour. 

It is interesting to note that this fungus cannot injure the 

1 Untersuchiingen aiis dem Forstb. Inst, zu Miinchen., I. pp. 151 — 163. 

- H. Mayer, Ueber den Parasitismus von Nectria cinnabarina. Un- 
lersiich. a. d. Forstb. Institut zu Micnchen, III. 

* [This fungus is extremely common on black currant and other trees in 
England. It is the species which, in its gonidial form, is so often observed 
on pea and bean sticks, dotting them over with scarlet points. ^Ed.] 



living cambium and cortex. It is, in fact, only able to invade 
them when they have been killed either by frost or by the want 
of w^ater consequent on the 
wood drying up centrifugally 
under the influence of the 
mycelium of the parasite. 

The simplest way to limit 
the increase of the parasite 
is to cut off and burn twigs 
and branches that are bear- 
'^^S gonidia and perithecia. 
The immediate application of 
tar or grafting- wax to wounds 
of all kinds is the best safe- 
guard ap-ainst infection. 


The various species of 
Polystignia induce the forma- 
tion of red fleshy blotches 
on the leaves of trees belong- 
ing to the genus Prnnus. 
Polystigma rubrujii^ occurs 
on the leaves of the plum 
and sloe. On the under side 
of the leaves, which in sum- 
mer display the large deep- 
red fleshy blotches, numerous 
small punctures will be found. 
These are the orifices of the 
spermagonia, which are buried 
in the leaf- parenchyma, and 
from which hooked colourless 

spermatia afterwards appear. The perithecia only occur upon 
the leaves between their fall and the following spring. By 
sowing the ascospores on young plum-leaves new spermogonia 
are obtained in six weeks. The best preventive measure is to 

^ Tulasne, Select a Fiaigorum Carpologia^ II. p. 76. 


Fig. 45. — Wood of maple containing the 
mycelium of N. cintmbariiia ; magni- 
fied by 1200. The vigorous mycelium, 
a a, traverses the elements and dis- 
solves the starch-grains, b c, first 
attacking the granulose. As the cel- 
lulose and the mycelial filaments, ci, 
are decomposed, a green solution ap- 
pears in the interior of the elements. 
The walls are much perforated, as at e e. 
(After H. Mayr.) 



get rid of the infected leaves by raking them together and 
burning or burying them. 

Poly stigma fiilvum attacks Primus Padiis and Amygdahis. 
This parasite is specially destructive to almond-trees, the yellow 
patches frequently embracing more than half of the whole leaf- 
surface. As the perithecia are formed on the fallen leaves in 
the following spring, it is advisable to burn them. 

Polystigma ochraceitm is parasitic on the leaves of the wild 


Mention may also be made here in a few words of that 
disease of cereals which, from the appearance of peculiar 
sclerotia or mycelial tubers, is designated ergot. 

The well-known black bodies that accompany ergot, and 
which occur on many species of Graminece, fall to the ground 
when the crop is reaped. There they pass the winter, and 
after germinating in the moist soil in the following spring 
each sclerotium generally produces a number of long-stalked 
spherical sporophores. Sunk over the whole surface of these 
reddish spherical bodies are to be seen numerous flask-shaped 
perithecia, whose orifices project somewhat above the general 
surface. Each ascus holds eight filamentous spores, which reach 
the open air by being pushed out through the orifice. Should a 
filamentous spore chance to reach and germinate on the flower 
of a cereal, the germ-tube forces its way into the ovary, where the 
mycelium develops in the tissues and almost completely con- 
sumes them. The ovary, which is entirely enveloped in mycelia, 
displays on its surface brain-like corrugations, which are the 
gonidial stromata. The gonidia, which are very small, oval, 
unicellular, and colourless, are imbedded in a sweet mucila- 
c-inous fluid which is secreted by the gonidial stromata, and 
appears in drops between the parts of the flower, being known 
as honey-dew. This gonidial form of the parasite was formerly 
designated Sphacelia segetum. Only after the formation of 
gonidia is finished does the ergot proper appear, and this it does 

Tulasne, Ajdi. des set. nat.^ 3rd sen, vol. xx. p. 56. 


at the base of the ovary, though completely independent of it. 
Morphologically it is essentially distinct from Sphacelia segetum 
in having a peculiar pseudo-parenchymatous structure. The 
original tissues of Sphaalia segetuui, with possibly some 
remnants of the ovary, die completely, and adhere for a short 
time to the apex of the ergot. 

From what has been said, it will be seen that the disease is 
spread partly by the sclerotia that hibernate from year to year, 
and partly by the innumerable gonidia which, suspended in the 
solution of honey-dew, are carried by various species of insects 
to the healthy graminaceous flowers where they germinate and 
which they infect. 

On account of the sclerotia that reach the fields in the seed 
being still capable of germinating in spring, the farmer en- 
deavours to prevent the disease by using clean seed-corn. 
The ergot should also be gathered before harvest, and this 
is accomplished at little cost, because it fetches a good 
price. * 


Amongst the many forms of canker met with on the oak, 
some of which still await investigation, the one that is caused 
by Aglaospora Taleola is characterised by a number of striking 
peculiarities. It would appear to be confined to woods under 
the age of forty years, and it is only so long as oaks are without 
true bark that they are liable to attack. The disease manifests 
itself in the following way. Both on the dominant and the 
smaller trees large patches of the smooth cortex die and become 
brown, but as this usually occurs only on one side of the stem 
the whole of the tree does not succumb. As the cortex often 
dies over long distances and on various sides of the tree, patches 
of sound cortex are to be met with surrounded on all sides by 
dead tissue (Fig. 46, \, a). A year afterwards numerous round 
or oval cushion-like stromata appear in the dead cortex. Later 
on these break through the periderm in one, two, or three places, 
so as to open up external communication with the gonidiophores 

' Fflrstlich-natiirwissensch. Zcitschrift, January, 1893. 
* [Owing to its use in medicine. — Ed.] 

H 2 

Fig. 46. — Portions of oaks, thirty-five years old, attacked by A. Taleola ; two thirds 
natural size. No. i has been attacked for one to two years, and still shows some 
sound patches, as at a a. Numerous stromata are visible on the dead cortex. No. 2 
has been suffering from the disease for four years. In the upper portion a canker- 
spot will be seen which is still unhealed, while below another is shown which is 
nearly closed. No. 3 represents a cross section showing three canker-spots, five, 
eight, and ten years old respectively. 



Fig. 47. — Cortex with stromata, 
which at a are covered by the 
periderm ; at b the periderm has 
been removed ; c shows the cross 
section of a stroma ; magnified 
five times. 

and the openings of the perithecia (Fig. 47, a). When the peri- 
derm is removed, the stroma appears in the brown cortex as a 
dark brown mass of tissue (Fig. 47, /;). If a section be made 
of the latter (Fig. 47, r), a black 
line will be perceived which sepa- 
rates the tissues of the cortex 
from those of the fungus. For 
each of the openings that appear 
at the surface one will generally 
recognize three perithecial cham- 
bers in the stroma. 

If a section is made at right 
angles to the stroma so as to 
expose the longitudinal view of 
the aperture of a perithecium 
(Fig. 48), it is seen that the 
bounding line consists of dark 
brown mycelium {a), which, be- 
ginning beneath the surface of the dead periderm, separates 
the whole stroma from the cortex, and even traverses the 
outermost sclerenchymatous bundle [b). The tissues thus 

enclosed consist of de- 
composed cortex and 
a large quantity of 
mycelium. The flask- 
shaped perithecia unite 
at ^to form a common 
neck, the aperture of 
which breaks through 
the periderm. In the 
outer layers of the cor- 
tex {c) close beneath 
the periderm are the 
gonidiophores, which 
disperse numerous go- 
nidia by abstriction. 
The gonidia are sickle-shaped (Fig. 49, a), while the ascospores 
that are produced in the perithecia are bicellular and provided 
with peculiar filamentous appendages (Fig. 49, b). Three such 

Fig. 48. — Cross section of a stroma, a represents 
the bounding zone consisting of mycelium; b, the 
sclerenchymatous strands of the cortex ; c, the 
gonidiophores ; d, the point where several peri- 
thecia unite. 


appendages spring from the middle of the spore, and one from 
each end. It is the gonidia and spores that infect the cortex. 
It would appear that before infection can occur the periderm 
must be slightly injured, and this happens with great frequency 
in thick oak woods owing to the rubbing 
of the branches of adjoining trees against 
each other. Soon after the cortex has been 
killed the alburnum becomes brown, and it 
frequently happens that absolute decom- 
position sets in, in the course of the year. 
The sound portions of the tree begin to 

Fig. 49.— «, gonidia, form a callus along the edge of the dead 
and b, asci oi A. ^ ^ 

Taleola. part, SO that sooner or later the latter is 

again covered over. The dead fibrous cortex 
maintains its position for some years, but in the end it is entirely 
cast off, so that the dead wood becomes visible (Fig. 46, 2). Of 
course the same tree may in the course of years be repeatedly 
infected at various points, as the accompanying cross section 
(Fig. 46, 3) shows. 

This disease is accountable for the death of a great number 
of oaks, and renders frequent felling and strong thinning 
necessary. The dominant fast-growing trees recover from their 
injuries more rapidly than those of slower growth, the con- 
sequence being that one is left with a thin wood, consisting for 
the most part of the largest trees. 

When the disease appears in a young oak wood, it is 
advisable at once to fell the infected trees, except where 
they belong to the dominant classes. The latter are thus 
stimulated to increased growth and enabled to recover from 
their wounds, and this result will follow with the greater 
certainty if the wood is underplanted with beeches or some 
other soil-improving species. The chances of infection are 
also thereby reduced, partly owing to the removal of the 
diseased trees, and partly because there are afterwards 
fewer opportunities for the occurrence of wounds induced 
by friction. 

Plowrightia morbosa^ (Cucurbitaria morbosa). The 

^ W. H. Farlow, "The Black-knot." Bull, of the Bussey Institution. 
Bot. Articles., 1876, p. 440. 


Black-knot of Stone-fruit Trees. Although this disease has 
hitherto been met with abundantly only in North America, still 
it may find a place here, because experience has taught that 
the diseases of cultivated plants may be very easily transported 
to us from other parts of the world. It makes its presence 
known by the occurrence on the twigs, of plums and cherries, of 
hemispherical swellings, which project to about -| of an inch and 
are usually congregated in groups. 

The surface of the swellings is covered by the gonidia of 
the parasite. The ascophores, which ripen in January, are in 
the form of round prominent black capsules. The twigs that 
are beset with knots should be removed as completely as possible 
and burned. 

Physalospora Bidwellii is a parasite of the vine which has 
been constantly spreading in France since 1885. The disease, 
which is known as " Black-rot," is usually confined to the 
berries, the young tendrils and stalks of the bunches being 
attacked only in exceptional cases. It makes its appearance 
a short time before the grapes ripen, when it may be recognized 
by the appearance of a small round sooty blotch, which on 
enlarging assumes a reddish colour, getting more intense towards 
the centre. In a day or two the berry is entirely destroyed, and 
three or four days later it assumes a dark colour and becomes 
perfectly withered. The skin and succulent tissues become 
wrinkled and shrunk and adhere to the seeds, without, 
however, showing any wounds. Thus it is not a case of 
decomposition but of withering. Gradually but slowly the 
disease spreads from bunch to bunch and grape to grape. Its 
occurrence would appear to depend on a high temperature and 
a humid atmosphere. 

The perithecia, pycnidia, and spermogonia of this parasite are 
known. The fungus survives from one year to another by 
means of stylospores which are contained in the pycnidia, and 
also by means of sclerotia. 

Coniothyrium diplodiella, a fungus which attacks the vine, is 
only known in the form of pycnidia. The disease which it induces 
has occurred as an epidemic in Italy, France, and Switzerland. 
The fungus for the most part attacks the branches of the raceme 
and the stalks of the berries, and these parts frequently become 


perfectly rotten before the berries themselves are attacked. The 
diseased berries soon assume a pale colour, and the yellowish 
white appearance is so characteristic as to have suggested for 
the disease its popular American name of " White-rot," to 
distinguish it from the dark discoloration induced by Black- 
rot. So far no effective method of treatment has been 

Gloeosporium ampelophagum {SpJiacdoina ampelinnni) 
produces the " Anthracosis " of the vine. On all parts of the 
plant are observed brown blotches which rapidly become black. 
These soon turn to depressions surrounded by a ridge. Later 
on, when the blotches and surrounding tissues dry up, projecting 
portions of the mycelium bearing gonidiophores make their 
appearance on the surface as small white spots. Pycnidia are to 
be found in the ridge. Anthracosis finds the conditions best 
suited to its development in warm situations after prolonged 
damp weather. 

Didymosphaeria populina attacks the Lombardy poplar, and 
produces a disease which is met with in man)' parts of France 
and Germany.^ 

In spring a brown blotch on one side of the young twigs 
situated on the lower branches indicates the point to which the 
mycelium of the fungus has extended. A little later all that 
portion which is situated above the original blotch becomes 
black and bends inwards. The buds situated below the diseased 
spot produce fresh shoots, which become infected in the following 
spring. Branches whose shoots are largely infested wither up 
entirely. The lower part of the tree, which suffers most from 
the disease, becomes very bushy owing to the abundant 
production of suckers, consequent on the stimulus imparted 
to the buds by the destruction of the shoots. These sucker- 
shoots ultimately succumb to the same fate as their predecessors. 
As a consequence of this state of things, all the nourishment 
is used up in that part of the tree, with the result that the top of 
the infected tree withers before the parasite has ascended so far. 
In the month of May the pycnidia begin to break through the 
epidermis to admit of the escape of elliptical hyaline st}'lospores, 

1 Vuillemin, Compt.Rend.2i\h March, 1889, and Prillieux,zW(/. 27th May, 


from which one or two much-septated germ-tubes are deve- 
loped laterally or terminally. The perithecia, which afterwards 
predominate, appear at the same time. As such they remain 
through the winter and the following spring. They are 
spherical in shape, and measure 0'2 mm. in diameter, and 
contain paraphyses and asci, the latter having eight bicellular 

The disease may be combated by lopping off the lower 
branches of infected trees. 

The gonidium form of this fungus is Fusidadiuni TremiilcUy 
which develops on leaves that have been infected in the previous 
spring by the ascospores. 


The essential difference between the Discoinycetcs and 
Pyrenoviycetes is that the asci are not formed on the internal 
wall of a closed spherical or flask-shaped organ (perithecium), 
but on the surface of an open saucer-shaped fructification 
(apothecium). Before the ripening of the spores the asci are, 
at most, protected by a covering which does not belong to 
the fructification itself, but is partly formed out of the epidermal 
layers of the host-plant. 

The Discomycetes are arranged into several sub-families, 
amongst which the P/iacidecs are to be distinguished by the 
hymenial layer, originating not on the surface of the fungus-body 
but in its interior, where it remains covered, temporarily or 
permanently, by the fungal tissues. 

Special mention may be made of the genera RJiytisma and 
Hysterium, which belong to this group. 


This fungus causes one of the best known of the blotch- 
diseases of the Maples. Acer platanoides suffers especially from 
this parasite, A. psendoplatanus and A. campestre in a less degree. 
On the leaves in July we first of all observe round yellow 

^ Cornu, Compt. rend., Ixxxvii. (1878), p. 178. 

* [The reader may be referred to Phillips's British Discomycetes (Kegan 
Paul & Co., 1887) for an account of our native forms.— ED.] 
■]■ [Very common in this country. — Ed.] 



marks from one to two centimetres in diameter. In August 
these begin to turn black (Fig. 50), and the leaves usually fall 
somewhat prematurely, so that, by the end of September, the 
trees are, for the most part, leafless. 

Not till some time during winter or the following spring do 
the numerous, somewhat prominent, vermiform apothecia appear 
on the black blotches of the rotting leaves. During damp warm 
weather these open by a longitudinal fissure. The disease is very 
easily produced artificially by laying such black portions of the 
leaves of the previous year on young maple-leaves during wet 

weather, or in a moist chamber, in 
the month of May. The filamentous 
spores which escape germinate, 
and produce fresh blotches. As 
this fungus agrees very closely with 
the next genus, Hysterium, both as 
regards origin of the perithecia 
and the development of the black 
stroma, I will not pursue this part 
of the subject further here. 

The injury, which consists in the 
reduced power of assimilation of the 
leaves, is not so great as to warrant 
the expense of instituting preven- 
tive measures. These would con- 
sist in raking togetherand removing 
the leaves in autumn. In gardens 
and parks, where this is done from 
other motives — for instance, in the 
English Garden in Munich — one never meets with an example of 
Rhytisina, whereas along the country roads and forest paths in 
the immediate neighbourhood of the city, where the leaves are 
left to lie in ditches and hollows, the disease occurs with great 


This species of R/iytisma, which closely resembles the pre- 
ceding one, is also to be met with on the leaves of Maples. 
They are to be distinguished, however, by the appearance of the 

Fig. 50. — Part of a leaf of the 
Norway maple, showing A'. 
acerimun. The black blotches 
are surrounded by a pale brown 
zone of dead tissue. 


region of the leaf that is occupied by the mycehum : this in 
the former case is a black blotch, and in the latter consists of 
black spots on a green ground. In autumn, when the leaves arc 
quite yellow, the green colour of the chlorophyll persists in the 
blotches for some considerable time. 


Black blotches similar to those produced by R. acerinmn often 
occur on Salix purpurea, nig7'icans, Caprea, aurita, &c. These 
are caused by RJiytisma salicinum, but are of relatively minor 


The genus Hysterhim possesses black elliptical to linear 
fructifications, which project from the leaves as black, lustrous, 
wart-like bodies. 

The spores are linear, their walls being externally 
mucilaginous and swollen. In the case of the three following 
species the germ-tube probably always enters by a stoma. The 
mycelium spreads between the cells in the parenchyma of the 
leaves of conifers, which consequently become brown and die. 
Should the disease attack a leaf near the base at a time when 
the upper parts are still healthy and capable of assimilating 
under the influence of light, and should the transportation of the 
products of assimilation from the leaf be prevented by the death 
of the elements of the bast, the plastic substances collect in the 
form of starch-granules in such large quantities as to completely 
fill up the leaf. 

The tissues of the leaves, which are at first pale green, after- 
wards become brown, and frequently the fructifications do not 
develop in them for more than a year. The ascogenous 
fructifications are often preceded by spermogonia, which in the 
case of the silver fir (Fig. 53) are arranged on the upper side 
of the leaf in two sinuous longitudinal ridges, whereas the 
apothecia which produce the ascospores are united on the under 
side of the leaf to form a single similar ridge. Both originate 
by the mycelium penetrating into and rupturing the epidermal 
cells. Ii then develops luxuriantly and forms a lenticular 
fungus-body, which afterwards becomes deep brown in colour. 



The stroma, which first produces paraphyses and later asci, 
originates underneath this mycehal body, which is firmly 
attached to the epidermal cells. 

The moister the weather, the more rapidly do the spores 
ripen. They are disseminated only when a long spell of wet 
weather has saturated the dead leaves with water and enabled 
the paraphyses and spore-walls to swell by contact with the 
water. This swelling leads to the rupturing of the leaf and 
the formation of a longitudinal fissure, which immediately 

Fig. 51. — Hysteritini niaci-osporum, showing a transverse section through a ripe 

ruptured stroma. 

recloses on the occurrence of dry weather or when the spores 
have escaped (Fig. 51). 


The distribution of this disease is coterminous with that of 
the silver fir, though I have found it really injurious only in the 
Erzgebirge, where large woods of silver firs, even of advanced 
age, had lost the majority of their leaves. Brownness of the 
leaves is always observed for the first time from May to July, 
and occurs on the two-year-old leaves which are entering their 
third year. A few months after the leaves have turned brown 
the spermogonia develop on their upper side, where two sinuously 
corrugated black longitudinal ridges make their appearance 
(right part of Fig. 53). 

^ R. Hartig, Wichtige Krankheiten, pp. 1 14 ei seq. 


1 09 

Later on the apothecia appear on the under side as a longi- 
tudinal ridge on the mid-rib, and ripen in the following April, 
when the shoot is three years old. A large proportion of the 
leaves, however, fall earlier, the perithecia being produced on 

Fig. 52. — The under side of a branch of 
silver fir, showing the perithecia united 
into a longitudinal ridge. 

Fig. 53. — Hysteriuin ner- 
viscqiiitim on the leaf 
of the silver fir ; the 
leaf on the left shows 
the apothecium on the 
under side, that on the 
right the spermogonium 
on the upper side. 


the comparatively few leaves that remain /;/ situ. It may also 
be remarked that still older leaves may contract the disease. 


This disease of the spruce produces the " spruce-leaf redness," 
which in many years occurs with great severity in woods from 
ten to forty years old. 

Its presence may be detected by the leaves of the previous 
year's shoots turning brown in May, or possibly not till autumn, 
and by the invariable occurrence of abundant mycelia in the 
leaves even before they become brown. Leaves which change 
colour in spring reveal the commencement of the formation of 
perithecia in July of the same year, and these ripen next spring 
in April and May. At that time they are present on the two- 
\ear-old shoots. I observed this rapid process of development 
in the humid climate of the Erzgebirge. At Eberswalde, on the 
other hand, the leaves on the two-year-old shoots do not become 
^ R. Hartig, M'ichtige Krankheiten^^. loi. 


brown till October, and the formation of pcrithecia begins on 
the three-year-old leaves in June of the following year, the spores 
ripening in the succeeding March and April. The apothecia 
appear as long, straight, lustrous-black ridges, for the most part 
only on the two under sides of the leaf (Fig. 55). The spores 
are about twice as long as those of H. nerviseqiiiuni. It is 
desirable that further investigation should be directed to this 
and to the immediately preceding disease, because I have not 
yet been able to clear up thoroughly many details in the develop- 
ment of these parasites. Especially has no explanation so far 

Fig. 54. — A spruce-branch, showing 
brown leaves on the upper two-year- 
old portion, and apothecia on the part 
that is three years old. 

Fig. 55. — Apothecia 
on a spruce- leaf. 

been offered regarding the phenornenon of the leaves of the 
youngest shoots of many spruces first becoming brown and 
then dropping off in autumn, so that these shoots become 
almost completely defoliated. Instead of long apothecia- 
ridges developing on such leaves, small isolated apothecia- 
tubercles similar to those of Hysterhiin Pinastri make their 


This is a species of fungus which is everywhere present in 
pine woods, and has been identified by Goppert^ as the cause of 

' Goppert, Verhandl. d. schlesischen Forstvcreins^ 1S52, p. 67. 
* [Common in England. — Ed.] 


the pine-leaf cast. Under the name " Pine-blight" (leaf-cast or 
shedding) the most various diseases have been included. These 
attack young and old pines, and are characterised by the leaves 
becoming brown, and usually also by their being prematurely 
shed. The causes of these diseased conditions are very various. 
In the first place, frost may actually cause the death of the young- 
leaves of pines. On July 23rd, 1878, large pines, especially such 
as were growing along the margin of the wood, were so severely 
affected by frost in the Turoscheln district that those parts of 
the new leaves which had emerged from the sheath died. 

As however the leaves of the Scotch pine do not protrude from 
the sheath before the beginning of June, late frost can do injury 
only in very few cases and in very exceptional localities. In 
many seasons one observes — frequently only on one side, espe- 
cially the east side, of trees— that all the leaves of the youngest 
shoots on trees that are much exposed to the wind become 
uniformly brown, except the lowest part, which is enveloped by 
the sheath. Whether, in such cases, the injury is always due to 
actual frost, or even to severe cooling, I am not in a position to 

In many cases the browning, death, and shedding of the leaves 
are the result of drought.^ In cases where the pine seed-beds 
have been covered with snow in winter, but which has disappeared 
after a few warm sunny days without the ground thawing, it will 
be found that the leaves soon become brown, and that the pines 
contract " the blight." If one examines the discoloured leaves 
after the appearance of the first symptoms of the disease, he fre- 
quently fails to find any trace of mycelia. It is also character- 
istic that the brownness is equally distributed over the whole 
leaf, or spreads back from the apex uniformly to a greater or less 
distance. In such a case we have to do with a drying up of the 
leaves, which do not receive a sufficient quantity of water from 
the frozen ground to compensate for the loss by evaporation 
that takes place in the clear dry weather of winter. Although 
erroneously ascribed to the action of frost, the cause is the 
same, too, in cases where the foliage of Pimts Strobus, the spruce, 
and other conifers, as also dicotyledonous evergreens, becomes 

^ Ebermayer, Die physikalischcti Eiiin>irkimge7i des Waldes auf Liift und 
Boden, 1873. 


withered on the side of the tree which is exposed to wind or sun. 
One should certainly not ascribe the withering of spruce-leaves 
during winter on the sunny side of the tree to the action of frost, 
and there is quite as little reason for relegating to the same 
cause the browning of young pines in frozen ground owing to 
direct insolation and strong air-currents. 

In the height of summer, about the month of July, exactly the 
same phenomenon may be observed during dry weather when 
pines in a drilled seed-bed on sandy soil are left standing for a 
second year. Only those pines remain perfectly healthy which are 
situated at the sides of the paths — that is to say, at the edges of 
the beds. In spring the one-year-old pines remain quite healthy, 
as long as the soil retains a sufficient supply of moisture and 
growth has not begun. Afterwards growth proceeds both above 
and below ground, though most vigorously in the marginal 
plants, whose roots can obtain water and nutriment from the 
paths as well as from the bed. Should transpiration of water by 
the plants in July be greatly increased partly in consequence of 
the air being dry and warm, and partly owing to the formation 
of new shoots and leaves ; and, on the other hand, should the 
soil have lost its winter moisture, then the pines wither in exactly 
the same way as happens in winter when the ground is frozen 
and the sky is clear. Only those plants remain green which 
stand nearest to the paths, or at least to the edge of the bed. 

In the nursery at Eberswalde, after a severe early frost in 
October, that portion of the ground of the pine seed-beds which 
the sun could not reach was still frost-bound at midday. On 
the other hand, the ground which the rays of the sun could affect 
was completely thawed and warmed in the course of the fore- 
noon. The seed-beds all over were beautifully green, and very 

A few days later all the pines in the seed-beds which had been 
shaded were brown, whereas those which had been exposed to 
the sun remained perfectly healthy. I am able to explain this 
phenomenon only by the fact that the frozen ground prevented 
the absorption of water by the roots, whereas the clear sky and 
relatively warm air furthered transpiration by the leaves. In 
this case shading had acted prejudicially. 

In by far the greater number of cases the pine leaf-cast is 


parasitic and epidemic in character, and is to be attributed to 
Hystcriniii Pinastri. Where " the cast " has become a calamity 
which year after year overtakes the seed-beds and young woods, 
it may at once be assumed that the disease is present in this 
most destructive form. 

It may frequently be recognized on young pine-seedlings 
even in the first autumn by the primary leaves acquiring 
brown blotches, while the other parts often assume a purple- 
red colour. 

Even at this early stage one always finds the characteristic 
mycelium of the parasite in the brown blotches. Frequently it 
also happens in the first autumn that a large number of very 
small black spermogonia appear on the diseased leaves (Fig. 56, 
d, e), the spermatia of which do not seem to be capable of germi- 
nating. After wet summers I have found perfectly ripe apo- 
thecia on the leaves of young pine-seedlings even in autumn. 
As a rule the black apothecia (Fig. 57, -r), which are much larger 
than the spermogonia, do not develop till the following year. 
Everything depends very much on the weather. On account of 
the dry leaves being unable to offer any nourishment to the 
fungus, its development, and. that of its sporophore, can proceed 
only during wet weather. Dry summers and cold winters do much 
to hinder the development and distribution of the fungus, whereas 
wet summers and mild muggy winters are specially favour- 
able for its growth. During mild winters the blight frequently 
spreads rapidly in nurseries and in regenerated forest areas. I 
have never observed the apothecia make their appearance during 
the first year on the leaves of pines two years old and upwards. 
They usually appear only in the third year, and generally after 
the leaves have fallen, though they not unfrequently also ripen on 
leaves that have remained /// si'tiL As regards the manner of 
distribution of the blight-fungus, it may be mentioned that the 
ripe apothecia rupture only after long-continued rain. Then the 
tissues of the leaf have been thoroughly softened, and a plentiful 
Supply of water has been able to reach the apothecia from within. 
This causes the asci and spores to swell, a state of things 
which is followed by the forcible rupturing of the apothecium- 
cover. Long-continued rains, however, do not usually occur 
except with west winds. They are less frequent with north or 




south winds. This is to be remembered in instituting preventive 
measures against the Wight. As a rule the diseased leaves of 
one-year-old seedling-pines die off completely in spring, without 
however falling off. On the other hand, one finds that all the 

diseased leaves of the bifo- 
\ \ . i /I .. . liar spurs of two-year-old 

pines suddenly become 
brown after the advent of 
warmer weather in March 
or April. This is followed 

Fig. -56. — A one-year-old pine in spring whicli 
has been attacked by H. /'inastri. a, 
healthy green leaves ; It, leaves with a 
brown apex and green base ; c, green 
leaves showing numerous brown blotches ; 
d, leaves whose upper portion has become 
brown during the previous winter, and 
which now bear the spermogonia of H. 
Pitias()-i ; the basal portion of these leaves 
has become brown more recently; e, leaves 
that are completely dead and covered by 

Fig. 57. — a, a one-year- 
old pine-leaf in April 
showing brown spots 
where infected, but still 
remaining green to- 
wards the base ; /', a 
dead two-year- old pine- 
leaf in April with 
ripe perithecia, x, and 
empty spermogonia, jj'. 

by a cast — that is, by defoliation of the dwarf shoots. This 
shedding, which frequently follows in a few days, is not to be 
regarded as the effect of immediately preceding unfavourable 
climatic conditions. It is, in fact, one result of the formation 
of cork at the base of the dwarf shoots, which are subsequently 


pushed off when t^rowth is resumed. Seedlings affected b)- 
the blight usuall}- perish, and can onl}- recover when about 
half of the leaves remain green and escape fresh infection. It 
is decidedl)- inadvisable to make use of diseased yearling seed- 
lings for planting. Neither is it advisable to use diseased 
pines two years old and upwards, because they are usuall}- 
so weakened by transplanting that thc)^ soon perish. Diseased 
plants on a regenerated area ma}-, under favourable circum- 
stances, recover from the disease. This, however, never happens 
when the mycelium of the fungus has spread from the leaves 
into the tissues of the stem itself In particular, if the medulla 
of the plant has become brown owing to the presence of the 
mycelium, death supervenes, even although the buds look quite 
healthy in spring. 

Should diseased leaves exist in the crowns of old pines, 
infection ma}- be easil}- induced b}- falling leaves. The young 
plants are infected either by the dehiscence of the apothecia of 
the diseased leaves that fall on them, or b}- the spores that are 
conv-e}-ed to them from the diseased leaves in the descending 
rain-drops. On this account it is not generall}^ advisable to 
form pine seed-beds under the drop of old pine-trees. 

Infection, in most cases, accompanies wind and rain which in 
blowing over an infected area catches up numerous spores, and 
bears them to sound plants. The experience that the disease is 
most prevalent on ver}- young plants, and in the case of older 
ones onl}- to a height of about two feet from the ground, is to be 
explained b}- the fact that onl}- the air-currents that are close to 
the ground have the chance of catching up the spores of the 
fungus and of depositing them upon plants. 

In order to raise health}- plants, it is advisable to form pine 
seed-beds in dicot}'ledonous woods, or, at least, at as great a 
distance as possible from young woods affected b}- the disease 
of leaf-shedding. Nurseries for seedlings and transplanted trees 
that have ever shown the disease should onl}- be used for fresh 
sowings after all diseased plants in the nurser}- itself, and in its 
neighbourhood, have been destro}-ed. 

If one is compelled to form seed-beds in unhealth}- districts, 
one should select such situations as do not adjoin, at least on 
the west side, young diseased woods. If the choice exists, it is 

I 2 


advisable to form the nursery in such a position at the edge of 
the wood that the west winds that impinge upon it shall first 
have blown over a wide extent of open country. The seed-beds, 
which are not to be made too large, should be enclosed on the 
side towards the wood by a perfectly close board fence 6h feet 
high. If spruce nurseries are available, containing dense and 
high beds of plants running from north to south, the pine seed- 
beds may be laid down between the beds of spruce, so that 
the latter form a protection against the spores that are borne by 
the west wind. Burying pines in deep trenches during winter 
often results in complete smothering of the plants owing to the 
exclusion of atmospheric oxygen. On the other hand, a light 
covering of leaves in winter affords good protection against 
contact with the spores. 

In protecting the areas under regeneration against fungal 
leaf- shed, regeneration by groups, under certain circumstances, 
gives the best results. Blanks in close pine woods may be very 
successfully restocked even where the disease destroys every- 
thing on larger clear-felled areas. This is undoubtedly due in 
the first place to the protection afforded against the spore-laden 
wind. In arranging the direction of felling one must take all 
possible care to prevent the west wind from blowing over large 
infected areas before it reaches the part of the wood that is being 
regenerated. Very extensive seed-fellings, when they adjoin 
each other, further the epidemic distribution of the disease in 
any case. Where seed is sown or trees are planted in stripes, it 
is a good plan to plough the stripes from north to south, and to 
throw the furrow slice on to the west side. If the furrows run from 
west to east, the west wind, blowing along them, is sure to carry 
the spores from diseased plants to sound ones. Where the 
spruce and Douglas fir thrive, these trees may be planted in 
stripes running north and south, partly at the edge of the wood, 
and partly at fixed distances throughout it, to act as screens and 
prevent the disease spreading. This must be done at least ten 
years before the final felling of the pine wood. 

Areas that are completely overrun by this disease should be 
planted with Weymouth pines, or some other disease-resisting 
species, according to the character of the soil. 

The W'eymouth pine suffers here and there from a leaf- 


disease which is due to an alUed parasite, Hystenuiii bracliy- 
sporuui. I am not j'et able to determine whether Hystcriuin 
laricimiui, which has been observed in great abundance 
on larches in certain districts of the Alps, is also a true 

The sub-family of the Pezizecu is to be distinguished by cup- 
shaped or saucer-shaped sporophores, which produce the 
hymenial layer free on the upper surface. 

PEZIZA (helotium) willkommii ^ * 

The fungus which induces the larch-blister is the cause of one 
of the most destructive and widely distributed diseases of the 
larch. It was first described by Willkommr who, however, 
made a mistake in its identification, and called it CoTtiaini 

Cortzaun, in fact, bears onl}- a superficial resemblance to 
Peziza, and belongs to the Basidioniycetcs. On the strength of a 
macroscopic similarity, also, it was next said to be Pesiza 
calyciiia, till I recognized that in this fungus we had to do with a 
new and still unknown species. The ascophore is at once 
distinguishable from that of P. calycina by its very short cup- 
stalk. So much by way of explaining the regrettable confusion 
of names. 

The larch is a forest tree which thrives splendid!}- throughout 
the whole of Germany, suffering but little from frost, at least 
not more so than other indigenous trees. Originall}', how- 
ever, its distribution was confined to high Alpine regions, 
because only there could it offer successful resistance to its 
enemies. Amongst these enemies are to be classed a number 
of insects, notably the Larch moth, ColeopJiom laricclla. This 
insect is also found in Alpine regions to a considerable height 
(over 4,000 feet), and so widely is it distributed, and so 

1 R. Hartig, Ujttersiichimge7i aiis dem Forstb. Inst., I. pp. 63 — 88. 

- Willkomm, Mikroskopische Feinde des Waldes, II. pp. 167 et seq. 

* [Though often overlooked, this fungus is quite common on the diseased 
Larches in EngLand and Scotland, with all the characters and relations to the 
"blisters" described by the author. Phillips, op. cit. p. 241, gives it as 
LachneUa calycina, and makes no note of its relation to the disease.— Ed.] 


numerously represented, that it is at first surprising why it does 
hardly any damage there. This is easily explained from the 
fact that at high elevations the transition from winter to 
spring is very rapid, and the development of the leaf-fascicles 
occupies but a short time. On the plains the larch begins to 
display green buds even towards the end of March, but their 
further development is often retarded for a long time, until, 
in the beginning of Ma)% the growth of the leaves pro- 
gresses more rapidly. This is the dangerous period for the 
larch, because when the caterpillars awake from hiberna- 
tion they begin to devour the green buds, and when growth 
proceeds slowly these are largely consumed, and the trees 
are, for the most part, defoliated. On the other hand, when 
the leaf-fascicles develop rapidly, a small proportion of the 
foliage suffices to feed the caterpillars. In Alpine regions 
the short spring saves the larches from complete or excessive 
defoliation, which, especially when often repeated, results in the 
crippling and death of the trees. The Larch Aphis also, 
C/iennes Laricis, damages the foliage of the larch to no small 
extent, though not nearly so much as the moth. The disease 
which is induced by P. Willkoinuiii differs entirely from the 
crippling which larches experience as a result of the attack of 
the moth, aphis, &c. This parasite is indigenous to high Alpine 
regions, where it produces the same disease that has resulted in 
the destruction of innumerable woods in Germany, Denmark, 
and Scotland. In its native habitat, however, it is only under 
special conditions of environment that it destroys whole woods. 
In order correctly to appreciate this point we must first review 
the course of development of the parasite. 

The spores — which originate in cup-shaped fructifications to 
be afterwards described — soon germinate in the presence of 
sufficient moisture, with effect not on an uninjured tree, however, 
but only on a wound. Such wounds are very often due to hail- 
stones, or to the dwarf-shoots being devoured in spring — as was 
mentioned above — or they are formed in the upper angle of 
the base of a branch (Fig. 58, U) owing to its depression under 
accumulations of snow or hoar-frost. From such wounds the 
vigorous, copiously ramifying, septate mycelium spreads in the 
soft bast, partly between and partly in the cells advancing in 



the sieve-tubes, and killing and browning the tissues. The 
mycelium also grows into the wood, and even penetrates as far 
as the medulla. 

That portion of the cortical tissues which has been killed 
during the first year dries up and appears as a depression, 
especially after growth in thickness has been resumed by the 
healthy part of the tree (Fig. 58). 

In summer thegrowth of the mycelium ceases, and an unusually 

Fig. 58. ^A canker-spot that has been re- 
cently formed in the upper portion of 
the stem of an eight-year-old larch from 
the Tyrol. Infection has occurred above 
the branch, b, where a crack has been 
formed in the tissues, owing to the branch 
having been depressed under a load of 
snow. Numerous immature ascophores, c, 
have already formed on the dead cortex. 

Fig. 59. — Cross section of a well- 
grown larch which has been at- 
tacked by/'. IVillkommii. Infection 
had occurred ten years previously 
at the dwarf shoot, a. Each year 
the mycelium advances in opposite 
directions, in spite of the fact that 
a layer of cork, b b, is formed at 
the beginning of each growing 
season along the boundary of the 
living tissue. In the immediately 
preceding year a very small quan- 
tity of wood had been formed. 

broad layer of cork is formed for the protection of the tree 
along the boundary between the sound and diseased tissues. 
These layers of cork (Fig. 59, b b) which form between the dead 
and living tissues induce external rupturing of the cortex 
at points along the boundary of the canker-spot (Fig. 60), 
the result being that turpentine flows from the interior of 
the tree. Year by year the canker-spot enlarges along its 
whole periphery, rather more rapidly, however, longitudinally 
than horizontally, and it is probably the vital activity of the 



cortical tissues which in summer causes a temporary inter- 
ruption to the progress of the parasite. In autumn the 
mycelium again succeeds in entering the Hving bast, either 
through the cambium region or by way of the wood, so that, 

as a matter of fact, the la}'er of 
cork is only of slight service. In 
proportion as the passage of the 
plastic substances is confined to 
one side of the tree, growth of 
the wood and bast is stimulated 
at that part (Fig. 59). Thus the 
conflict between parasite and 
host-plant may remain long un- 
decided, and in the Tyrol I found 
larches still alive with blisters of 
a hundred years' standing. 

Should the parasite advance re- 
latively quickly, and, at the same 
time, should the growth of the tree 
at the affected part be slow, then 
the canker- spot soon embraces 
the whole stem or branch (Fig. 59), 
and the tree dies above this spot. 
By artificial mycelial infection 
one may, almost without fail, 
produce a blister on any part of 
a sound larch. 

Soon after the death of the 
cortical tissues, the cushion-like 
stromata of the parasite originate 
in the form of small yellowish 
white pustules of the size of a 
pin-head (Fig. 58 r. Fig. 60 a). In 
the interior of these stromata, 
and partly on their surface as 
well, vermiform passages or roundish cavities are formed, the walls 
of which are covered with innumerable club-shaped sterigmata, at 
whose apex extremely minute cells originate. Whether these 
organs, w^hich appear to be incapable of germination, are 

Fig. 60. — A canker-spot of two 
years' standing, close to the collar, 
and hidden by the grass. On the 
upper portion, which is exposed 
to air-currents, the stromata are 
abortive ; but in the lower por- 
tion, which has been kept moist, 
they have developed to form 
vigorous ascocarps. 


abortive gonidia, or arc to be classed with spermatia, remains, 
in the meantime, undetermined. In this place it is specialh- 
important to emphasise the fact that they are incapable of 
assisting in the distribution of the parasite. 

The small stromata are very readily affected by a dr)' atmo- 
sphere and by air-currents, in which they quickly wither and die. 
They develop only when constantly surrounded b}- moist aiiV 
Under such circumstances the)' produce the well-known cup- 
shaped ascocarg^s (Fig. 60, b b). These possess a hymenium 
of a fine red colour. The hymenium consists of innumer- 
able asci surrounded by filamentous paraphyses. Eight colour- 
less spores arc formed in the interior of each ascus. The fact 
that the mycelium penetrates even into the wood, and kills it, 
explains why one or a few small blisters may greatly interfere 
with the growth of the whole stem. Numerous cup-shaped 
ascocarps ultimately make their appearance on the dead cortex, 
and these are met with even when blisters are absent. 

In muggy situations the larches soon become diseased, and die 
in a few years without any large blisters making their appear- 
ance. The cup-shaped ascocarps of the parasite appear upon the 
cortex. It looks as though the large quantity of water present in 
larches whose transpiration is interfered with greatly favours the 
development and spread of the fungus in the wood, and that the 
disease consequently spreads throughout the whole plant. 

The foregoing descriptive sketch of the results of my investiga- 
tions may suffice to explain the recognized facts connected with 
the occurrence and distribution of the disease. 

The larch-blister has been indigenous to high Alpine regions 
from time immemorial. It occurs, however, with marked in- 
tensity only in damp muggy valleys in immediate proximity to 
lakes {e.g. the Achensee in the Tyrol, &c.), though on plateaus 
it may also destroy a small tree here and there. Owing to the 
prevalence of air-currents, freely exposed ascocarps never ripen 
on plateaus and valley-slopes. The ascocarps ripen only on 
those blisters which are situated at the foot of the stem close to 
the ground, or on blistered branches that are in contact with the 
earth. This is owing to the surrounding high grass sheltering the 
young ascocarps against air-currents, and so keeping them moist. 

In the early decades of this century, when the larch was planted 


in various parts of Germany, the enemy was left behind in its 
native habitat, and the trees flourished to perfection. Prob- 
ably every old forester knows some groups of larches of the 
most stately growth which date back to that period. In conse- 
quence of these satisfactory results, the larch was generally 
planted throughout the whole of Germany. Most excellent 
results were obtained, even where the inferior quality of the soil 
held out but a poor prospect of success. 

But after woods of all sizes had been established from the foot 
of the Alps to the coasts of the North Sea and Baltic, the fungus 
spread downwards from the Alps, to find everywhere the most 
favourable conditions for its development. These consisted of 
dense young pure woods, groups that had been formed in re- 
planting up old beech woods, moist stagnant air, wounds caused 
by the moth, &c. Commerce also assisted to intensify the evil, 
diseased larches being sent out from the nurseries and trans- 
ported from district to district. 

Under these conditions the fructifications of the fungus 
attained to luxuriant development and ripened their spores on 
the blisters, while the spores found ample opportunity of germin- 
ating, and of infecting the trees in the close pure woods. To-day 
but few of the many promising young woods remain. The 
larches have maintained their ground best in those woods 
where a few were introduced as advance-growth. The air 
circulating in the freely developed crowns has not only kept 
the disease in check, but has also prevented the spores from 
ripening on diseased specimens. 

Supposing that we have to do with a diseased larch wood, it 
is first necessary to determine whether the damage is entirely 
due to the moth or whether it is a case of fungoid blister. 

Often enough both will appear in company. If it is simply 
a case of stunting in consequence of the attack of the moth, 
pruning away the branches till only the vigorous upper part 
of the crown remains may be permanently beneficial. The 
upper branches will grow vigorously, and may form a good, 
permanently healthy crown, especially as the moth is most 
destructive on the lower branches. 

If it is a case of fungoid injury, pruning may assist somewhat 
only if the bole as a whole, and especially the part in the crown, 


is sound If a tree is in vigorous growth, the smaller blisters 
low down on the stem, although they increase in size, will induce 
death only at an advanced age. 

Blisters on the branches are, in themselves, of less importance. 
They merely contribute to the danger of the further spread of 
the disease by means of spores. 

As regards the future cultivation on plains and at moderate 
elevations of this so essentially valuable tree, the following points 
may be noted in the light of what has been said. It should only 
be grown singly — that is to say, it should form but a small part 
of a mixture, and it ought, if possible, to be planted somewhat 
in advance of the other trees. It should never be planted in 
pure woods, and should always occupy an open situation. Where 
diseased woods are present in the immediate neighbourhood, 
it is better to abandon the idea of cultivating this tree. The 
greatest caution is to be exercised in procuring young trees 
from outside sources, and plants showing any signs of disease in 
the seed or plant beds must at once be removed and burned. 


On the light sandy soils of Germany, France, &c., especially 
in pine and other coniferous woods, one not unfrequently meets 
with numerous ascophores o{ R.iindulata growing on the ground. 
These bear a considerable resemblance to a morel (Fig. 61). 
In diameter they vary from two thirds of an inch to two inches. 
The broad ascophore (Fig. 61, a) is undulating and chestnut 
brown on the upper surface, diverse in shape, of a velvety lustre, 
and glutinous in wet weather. The under side (Fig. 61, h), which is 
destitute of a stalk, is pale yellow and woolly, and is frequently 
united to the subterranean mycelium by means of numerous loose 
mycelial strands (Fig. 62). If a section be made of the ascophore, 
it will be found that towards the upper surface the hymenium 
(Fig. 63) is composedofasci,each containing eight spores, amongst 
which filamentous septate paraphyses, clavate towards the apex, 
will be made out (Fig. 63, a). Besides these there are present 
numerous non-septate secreting-tubes {b), which project a little 
above the surface of the hymenium. These are filled with a 

I R. \{?LX\:\'g, Naturioisscnschaft : Zcttschriff, Awgwit 1892. 

* [This occurs on heaths, &c., in England.— ED.] 



brown secretion which pours over the surface as a slimy glutinous 
substance, swarming with bacteria. The bacteria also find their 
way between the paraphyses, so that it is scarcely possible to 
get a culture of spores that is free from them. It is these, too, 
which induce the rapid decay and solution of the entire ascophorc. 
The spores (Fig. 64, a) are spindle-shaped and pointed at both 
ends, and the wall of the spore is thickened at both of the ex- 
tremities. Before germinating, each 
spore generally contains two large 
drops of oil. 

Ascophores of Rhiai)ia were 
sent to me ten years ago from 
Silesia, with the remark that in a 
young pine wood where many of 
the plants had died the fructifica- 
tions of this fungus appeared on 
the surface of the ground in the 
neighbourhood of the dead trees. 

Fig. 61. — a, the upper side, and, /', 
the under side of a sporophore 
of J\. undulata; t: is a small fun- 

Fig. 62. — Section of a sporophore. 

My request that a few dead trees should be forwarded for in- 
vestigation was not complied with, so that it was only two years 
ago, on receipt of material from Herr von Bliicher, forester in 
Schwerin, that I found myself in a position to make a more 
intimate acquaintance with the parasite and its life-history. 

In the beginning of August 1890, Herr von Bliicher and Herr 
von der Liihe sent me numerous ascophores of the parasite, as 
well as diseased and dead conifers, along with information re- 
garding the occurrence of the disease at Schildfeld, near Bennin, 
in Mecklenburg-Schwerin, The diseased and dead plants were 



specimens four to ten years old of Abies pectinata, Tsnga Mer- 

tcnsiana, Pseudotsuga Douglasii, Picea Sitkcvnsis, Pinus Strobus, 

and Larix curopcca. 

The part of the wood that was attacked extended to about 

2I acres. In the whiter of 1889-90 the wood which then existed 
— namely, a thin stocking of pines, larches, 
and spruces about fifty years old — was 
stubbed, and in the spring of 1890 the 
area was replanted chiefly with three- to 
four-year-old plants, which were inserted 
partly in pits and partly in notches. 

In the month of June disease had 
appeared among the plants. The leaves 


Fig. 63. — H ymenium , 
consisting of a, 
paraphyses ; b, se- 
creting-tubes ; c, 
asci, which contain 
eight spores each. 

Fig. 64. — a, spores of Rhizina ; l>, ditto, 
twenty-four hours after sowing ; c, 
ditto, twenty-four hours later ; (/, the 
spore c more highly magnified. 

rapidly died and fell off, and the fungus appeared to be gra- 
dually spreading over the area. The ascophores were found 
almost exclusively at a distance of about ten inches from the 
plants, but on the surface of what had been the pit. But 
between the plants also, on the bare ground covered with raw 
humus, numerous ascophores were met with. The soil was 
sandv in character, covered with raw humus and bilberry-bushes. 



The above report refers onl}- to conifers attacked b}' the 
parasite. Professor Crie of Rennes was good enough to send me 
more than once the roots of diseased plants of Castanea t'esca. 
On one of these I found RJiizina iindidata luxuriantly developed. 
On removing a diseased or dead plant from the ground, one 
will find that a large quantity of sandy soil is firmly held 
amongst the roots by means of numerous fungus-filaments, but 

that no outpouring of resin what- 
ever is visible (Fig. 65). 

On the roots being isolated 

fi and carefully examined, it will 

be found that peculiar m}'celial 

Fig. 65. — The roots of a silver fir 
which has been killed by Rhizina. 

Fig. 66. — Mycelial growths resembling 
Rhizoctonia, which are met with on 
the roots of plants infested by 
Rhizina, magnified by 3. Mycelial 
strands protruding from the cut sur- 
face in moist air. Natural size. 

bodies resembling Rhizoctonia project from the cortex. At a 
distance of two to three fifths of an inch these begin to ramify, 
and ultimately divide into filamentous mycelia (Fig. 66). If 
one cuts off a root and makes a culture in a moist chamber, it 
will be found that such mycelial bodies will form in large numbers 
on the cortex, or on the cut surface of the wood. These ramify 
in the usual way and end in a fine point (Figs. 66 and 6j). 
and are always brilliantly white in colour. 

A microscopic investigation will reveal the cause of this 
colour, which is due to drops of ethereal oil adhering in great 
numbers to the external filaments, or to the apices of the fine 



hair-like filaments which stand more or less at right angles to 
the m)-celial strands (Figs. 68 and 69). 

These short simple or compound hairs produce a large drop 
of ethereal oil at the apex, which finall)- rup- 
tures the elastic cell-wall at the end of the hair, 
and flows out. The hair thus comes to have 
a funnel-like aperture at its apex (Fig. 69, e). 

I am not aware of attention having pre- 
vioush' been called to the formation of 
ethereal oil in the form of drops by fungi. 
The oil is immediately dissolved by alcohol. 
As the mycelial filaments in the periphery 
of the strands contain numerous small oil- 
drops, it would appear that this ethereal oil 
is also exuded from the lateral walls of the 
hyphas, although it is quite possible that this 
has gradually found its way thither from the 
apex of the hairs. When the filamentous 
m}xelia that envelop the soil-particles are 
examined, it will be found that most of the thin threads possess 
numerous clamp cells, and are somewhat brown in colour. 

Fig. 67.— Mycelial 
strands of Rhiziua 
which have been 
cultivated in moist 
air. They are part- 
ly separated from 
the wood. 

Fig. 6S. — A mycelial strand 
bearing; hairs. 

Fig. 69. — a, a mycelial filament with an oil- 
drop attached ; b, ditto, with an oil-drop at 
the apex ; c, a hair with a large oil-drop ; 
d, a bifurcated hair from whose apices the 
oil-drops have become detached ; c, apex 
of a hair viewed from above. 

Although I have much diffidence in maintaining that this 
feature, which otherwise is peculiar to the Hyvieuouiycetes, is 


characteristic of this parasite, still I cannot doubt that these 
filaments with clamp cells belong to it. However, I will not 
maintain this as an absolute fact, especially as clamps do not 
occur either in the interior of plants or in the mycelia pro- 
duced by germinating spores of RJiizina. 

My first cultures were undertaken on August 19th, 1890, with 
fresh spores, which I sowed partly on a gelatine extract of 
fruit and partly on humus sandy soil. These produced no 

On repeating the culture with numerous spores on Septem- 
ber 1 8th, only a single one germinated. On the other hand, 
germination was general in twenty-four hours in a seeding on 
gelatine extract of fruit which was undertaken on November 
18th. These germinating spores are shown in Fig. 64, b. The 
extraordinarily thick germ-tube proceeded from the lateral 
wall of the spore, and from the first its diameter was as 
great as that of the spore itself. After only forty-eight 
hours the germinating spore had reached the stage which is 
represented, slightly magnified, in Fig. 64, c. 

The stout much-branching mycelium is septate, and resembles 
in every respect that which is found penetrating the healthy 
cortex of slightly or much-diseased plants. Under such circum- 
stances it grows between the cells of the parenchymatous 
tissues, while in the soft bast its progress is partly intercellular 
and partly intracellular, the sieve-tubes being frequently packed 
full of a dense filamentous mycelium. In the process of time 
the mycelium kills the tissues of the cortex and soft bast, whose 
elements become brown and completely dismembered, or, in other 
words, isolated. The development is so luxuriant that it forms, 
in certain places, a pseudo-parenchymatous fungus-tissue, consist- 
ing of vesicular swollen cells. This however is speedily destroyed 
as soon as the tissues between the wood and periderm become 
almost completely decayed. In this process of decay very 
minute organisms resembling Micrococcus play an important 
part. When employing a high power, the whole field of view 
sometimes swarms with these minute cells, whose diameter does 
not exceed i to i"5 micromillimeters. These originate (Fig. 70) 
on very small stalks resembling sterigmata, which project some 
from the lateral walls and some from the apices of the fila- 



mentous mycelia. Subsequentl}' they appear to increase by 

It is very desirable that foresters, especially in sandy districts, 
should direct their attention to the occurrence and biology of 
this parasite. 

The term "Soil Canker" has been used for twenty years to 
designate all those diseases in young and old woods where no 
indications above ground can be referred to as a cause. Such 
diseases have their seat below the surface, and gradually 
spreading from the first point of 
attack they occasion blanks and 
gaps in woods and nurseries. 
During the last twenty years I 
have described a whole series 
of parasitic fungi which induce 
such diseases. These include 
Agaricus inelleus, Trametes ra- 
diciperda, Polypoms vaporarius, 
Rosellinia guerciua, DcviatopJiora 
necatrix, and PJiytopJitJiora oui- 
nivora (in the narrow sense). 
To these must now be added 
RJiizina nndulata. 

The various species of Vac- 
cinecE are attacked by parasites 
of the genus Sclerotinia.^ The 
gonidiophores appear in spring 
on young leaves and stems, which 
consequently become brown, in 

the form of a mould-like covering which emits an almond- 
like perfume. The insects that are thereby attracted con- 
vey the gonidia to the stigmata of the flowers of the 
Vaccinea;. A sclerotium is formed in the berries, which be- 
come brown, dry, and " mummified " and drop off, and from 
them there develop in the following spring one or two 
long-stalked chestnut brown cup- like ascocarps. The ejected 
ascospores infect the young shoots, and again produce the 
gonidium-bearing form. 

1 Woronin, Uebcr die ScIc7-otienkra7ikheit der Vaccimeribecrcn, 1888. 


Fig. 70. — Mycelium ol KJiizina from 
the cortex of the silver fir. a, a 
filament of average thickness ; 
b, very thin filaments ; c, gonidia 
resembling Micnxoccus. Magni- 
fied 1500 times. 



Sclerotinia Vaccinii is parasitic on Vacciniiun Vitis Idcsa ; 

S. Oxycocci, on V. Oxycoccos ; S. baccanini, on V. Myrtilliis ; 

S. niegalospora, on V. idiginosnui. 

Of still greater importance, from an agricultural point of 

view, is Peziza ciborioides {Sclero- 
tinia Trifolionim), the clover-can- 
ker, or the sclerotium disease of the 
clover. This parasite is interesting 
from the fact that on clover-plants 
infested by the mycelium sclerotia 
from O'l — I cm. in size are formed, 
and these produce ascocarps in the 
following year in July or August. 

A similar course of development 
is found in Peziza Sclerotionivi 
{Sclerotinia Libertiana), which pro- 
duces the sclerotium disease of the 
beetroot and carrot. 

The best known is Peziza 
Fuckeliatia, through its gonidium- 
bearing form Botrytis cinej'ea* the 
vine-mould, which finds its way on 
to various plants in forcing-houses 
and conservatories, producing a 
loose grey mycelial covering and 
killing the twigs. 

For some years a species, Botrytis 
Douglasii} has proved injurious to 
the Douglas fir, which is now gener- 
ally cultivated in Germany. In 
seed- and plant-beds especially, 
where infection by lateral contact 
is such an easy matter, one often 
notices that the young incompletely 

developed shoots die and become brown. The shoots of the 

Fig. 71.— Branch of the Douglas 
fir, the youngest shoot of which 
has been killed by B. Donglasii. 
The apex of the shoot of the 
previous year has also been 

1 Botrytis Douglasii n. sp., C. Freiherr v. Tubeuf, Bcitriige zur Kenntniss 
der Baumkrankheite7i. Berlin, Springer, 1888. 

* [Several forms of Botrytis are common and destructive parasites in our 
green-houses and gardens. — Ed.] 


previous year may also die back for a certain distance 
(Fig. 71). 

One afterwards observes, both on the leaves and on the 
twigs, small black sclerotia not larger than a pin-head. In 
a moist chamber these germinate and produce the gonidiophores 
of Botrytis. The gonidia germinate easily, and infect the tender 
shoot of the Douglas fir. Tubeuf's researches show that silver 
firs, spruces, and larches were also infected by this fungus, 
and it remains to be determined whether diseases in the forest 
may not also be induced by it. 


In the case of the parasites that belong to this sub-family 
of the Discoinycetes, there is no proper fructification. The 
hymenium is a flat layer which occupies the surface of the 
plant, and consists of free tubes which develop among the 
epidermal cells, or between the epidermis and cuticle. 

All the species induce characteristic hypertrophy of the 
part of the plant that is attacked. 

This is a widely distributed parasite, which is familiar enough 
by causing the formation of the so-called " Mock," " Pocket," 
" Starved," &c., plums. Its mycelium persists from year to 
year in the soft bast of the branches of Priums doniestica, 
P. spinosa, and P. Padus, in which it grows intercellularly, 
gaining access to and contorting the young leafy shoots. The 
same is true in the case of the flowers, where malformation of 
the ovary is recognisable even in the beginning of May. 
Proceeding from the soft bast the mycelium spreads through 
the fleshy parenchyma of the fruit, where, on the one hand, 
it prevents the formation of the stone and seed, and, on the 
other, induces elongation and the well-known deformation of the 

1 Sadebeck, U7tte7'siichungen iibcr die Pilzgattieng ^'- ExoasciisP Hamburg, 

- De Bary, Beitriige ziir JMorphologte dcr Pilze, I. p. 33. 

* [Common in this country. Sadebeck has just pubUshed an exhaustive 
monograph on the whole group of parasitic ExoascccB (Hamburg, 1893), 
revising the classification, and clearing up many doubtful points.— Ed.] 

K 2 


fruit. Numerous mycelial branches penetrate between the 
epidermis and cuticle, where, by the formation of transverse 
branches, they form short chambers. In this way an almost 
uninterrupted layer of fungal mycelium is formed under the 
cuticle. Each fungus-cell next grows outwards to produce a 
short cylindrical ascus, and the cuticle after being detached 
from the epidermis is ruptured, and the ascogenous layer 
becomes completely exposed. 

Each ascus becomes separated from the basal part, or "stalk," 
by a transverse septum ; and, by free cell-formation, six to eight 
roundish spores are formed in its interior, to be afterwards 
ejected through the ruptured apex. The spores either germi- 
nate forthwith, or multiply by budding, and form a kind of yeast. 

The pocket-plums decay owing to the concurrence of numerous 
saprophytic fungus-forms. 

Exoascus deformans is closely related to the foregoing species, 
but lives partly in the leaves and shoots of Persica vulgaris and 
Amygdalus comnmnis, and partly in the leaves and shoots of 
Primus avuun, P. Cerasus, P. Chanicccerasiis, and P. domestica. 
On these trees, according to the investigations of Rathay,^ it 
causes the so-called witches' brooms. Whether the Exoascus 
that occurs on cherries is really a new species {^Exoascus Wies- 
neri), as Rathay assumes, or whether the distinctions that have 
been noted are not perhaps due to differences in the host-plants, 
must remain doubtful until infection-experiments have been 
carried out. Peculiar crumpling is induced in the leaves, similar 
to that which is sometimes caused by Aphides. The branches 
that have been taken possession of by the fungus anastomose 
freely, and usually exhibit decided negative geotropism, while 
the basal portion is often hypertrophied. These constitute 
the thunder-brooms and witches' brooms. Towards the base, 
the branches of these witches' brooms are often double the 
thickness of the branches from which they spring. Towards 
the apex, on the other hand, they become normal. A possible 
explanation of these phenomena is that, as the mycelium grows 
more slowly than the young shoot, it finds immature tissue only 

1 Rathay, Ueber die Hexenbesen der Kirschbdume und iiber Exoascus 
Wiesneri, Rath., im Sitzber. d. Wien. Akad. d. Wzssensch., vol. Ixxxiii., March 



at its base, which, under the influence of the parasite, enlarges or 
increases abnormally. On the other hand, the mycelium arrives 
too late at the apex of the shoot to be able to exert a similar 
influence there. 

Exoascus InsititiiS produces witches' brooms on Pritnus in- 

Exoascus bnllatiis induces bladder-like swellings, which after- 
w^ards become mealy underneath, on the leaves of pear-trees. 
In the case of the hawthorn it produces formations like witches' 
brooms, which bear leaves of a reddish colour. 

Exoascus ahiitorqiius {Asconiyces Tosqui)ietii) often appears im 
great abundance both on the 
leaves of Alnus glutinosa and 
on the scales of the female 
catkins of that tree and of 
Alnus iiicana. Not only does 
it cause the leaves to become 
crumpled and corrugated, but 
also to increase in size in 
every way. On the cones of 
the alders it produces pocket- 
like outgrowths, which, when 
fresh, are of a brilliant red 
colour, and remind one some- 
what of the pockets of plums 
(Fig. 72). 

Exoascus flavus {Sadebeckii) 
causes the formation of yellow blotches, in this case also on the 
leaves of A. ghitmosa and A. incana. 

Exoascus epipJiyllus, which infests the leaves of Alnus incana 
and A. glutinosa, is only with difficulty to be distinguished from 
the former species by its broader stalk-cells. It induces sinuous 
crumplings of the leaves, the outgrowths usually appearing on 
the upper side. 

Exoascus borcalis produces witches' brooms on Alnus incana. 
These are very numerous near Munich and at other places in 
Bavaria. It is probably identical with E. cpipJiyllus. 

Exoascus turgidus {Taphrina betulina) very often produces 
witches' brooms on the birch. 

Fig. 72. — Malformation of the fruit of 
Alnus imanu induced by Exoascus. 



Fig. 73. — A leaf of /'. niora 
affected by Exoasais Pof-iili. 

f IG. 74. — Malformation 
of the fruit of P. 
trennila, due to E. 

Fig. 75. — -A witches' broom of the hornbeam induced by Exoasctis Carpini. 
Half natural size. 


Exoasciis Betithe {Ascojiiyces BetiilcE) produces bladder-like 
outgrowths on the upper side of the leaves of the birch. 

Exoascus carnca produces globular bladder-likc swellings on 
the leaves of the birch. 

Exoascus aureus* {Tap/iriua aurea, T. Populi) produces golden 
yellow outgrowths on the leaves of Populus nigra (Fig. 73) 
and pocket-like outgrowths on the ovary of P. treimda and 
P. alba (Fig. 74). 

Exoascus Carpiiii produces witches' brooms on the hornbeam 

(Fig- 75)- 

Exoascus ccerulcscens {^Ascomyces ccBruksceiis) produces bladder- 
like swellings on oak-leaves. 

Exoascus Ulmi produces outgrowths on the upper side of elm- 


The number of those fungus-forms with all of whose stages 
of development we are not yet acquainted is an extremely 
large one. In particular a large number of fungi are known 
to us with whose gonidia — whether on sporiferous hyphae or in 
closed organs (pycnidia, spermogonia) — we are familiar, but of 
whose ascophores we are ignorant, so that we are unable 
systematically to classify them. 

A few of the more important species that occur parasitically 
on trees, especially on forest trees, may be referred to here. 

Cercospora acerina.i The Maple-Seedling Fungus 

In rainy years a disease is sometimes conspicuously prevalent, 
both on maple seedlings in the nursery and on those which have 
sprung up naturally. It is to be recognized by the cotyledons 
and first leaves, and also by the shoot axis, becoming black 
and decomposed, or, if less severe, merely by black blotches 

1 R. Hartig, Ufitcrsuchujij^efi, I. p. 58. 

*[I have found this species deforming the ovaries of poplars in Surrey.— 

t [No group of fungi offers more opportunities to the investigator anxious 
to add to our knowledge of pathogenic forms than the numerous " imperfect " 
ascomycetes so common on our trees, &c. — Ed.] 


appearing on the leaves. Even with the naked eye one may 
often recognize a grey covering on the diseased leaves. 

On more thorough investigation we perceive a luxuriant 
mycelial growth in the tissues of the diseased parts, from which 
innumerable short gonidiophores grow outwards. These pro- 
duce tufts of long curved multicellular gonidia, which germinate 
in moist air even in a few hours, and push their germ-tube 
directly into the epidermis of the maple-leaves, which conse- 
quently become brown. 

The mycelium, which is intercellular, swells up to form 
large brown mycelial resting-cells and cell-plexuses, which 
contain oil-drops. These persist during the winter, and carry the 
disease over to the following year. The fungus is also able to 
live saprophytically on humus in the soil. 


The disease induced by this fungus, which has often been met 
with all over Germany, appears most frequently in seed- and 
plant-beds which are stocked with spruces and silver firs. I 
described it in the AUgevieine Forst- unci Jagd-Zeitung, 1883, 
where I advanced the view that it was due to the formation 
of ice and the consequent crushing of the cambium. As I 
expressly stated, the truth of the hypothesis which I there 
advanced had still to be determined. Von Tubeuf has now 
proved that here, as in so many cases, we have to do with a 
parasitic disease. In summer one notices in nurseries of the 
spruce and silver fir that a number of plants first become pale 
and then die. If the plants are pulled up, it is seen that the 
cortex on the parts immediately over the ground is withered, 
but that, farther up, the stem is swollen as a natural consequence 
of continued growth (Fig. "](>). 

When the wood dries up or dies at the point where death of the 
cortex first took place, the plant must perish. On the rind, at the 
place where contraction is visible, one finds the mycelium of the 
fungus, and numerous gonidial cushions which develop partly in 

1 C. V. Tubeuf, Beitriige zur Ktnntniss dcr Baui/ikrank/ictte?i, pp. 40 — 51, 
Plate V. Berlin, Springer. 1888. 

Fig. 77. — Gonidiophoie of P. Hartigii. 
(After Tubeuf.) 

Fig. 78. — A branch of silver fir infected by 
Fig. 76.— a young spruce which has been infected P. abietitta. Numerous black tubercles 

close to the ground by P. Hartigii. 

are visible on the dead cortex. 


spheroidal p)xnidia, and partly on flat stromata which arc 
disposed in the tissue of the cortex. 

The characteristic gonidia (Fig. jj), which are situated on 
short or long stalks, are at first hyaline, thin, oval, and uni- 
cellular, but afterwards become four-celled owing to repeated 
transverse division. The two middle cells are large and dark, 
the small stalk-cell and the terminal cell remain colourless. The 
latter pushes out a branched filament which, however, must not 
be confounded with a germ-tube. It is only one or other of the 
three lower cells that germinates, most frequently the lower of 
the two brown middle cells. 

On account of the general distribution of this disease, and the 
consequent loss incurred, it would appear advisable carefully to 
root out and burn all diseased and dead plants that may be 
found in nurseries. 

Similar pathological symptoms have also been observed on 
young beeches, ashes, and maples. I should be glad to receive 
such plants, in order to prove whether parasites are the cause of 
disease, and, if so, to determine the species. 


A disease which has not hitherto been described, but which is 
extremely common on young and old trees in the Bavarian 
Forest, is due to a parasite which may temporarily be called 
PJioina abietina. The disease may be recognized by both small 
and large branches of the silver fir becoming pale and withered ; 
in fact, I have occasionally observed diseased spots two inches 
in diameter on the cortex of silver firs as thick as one's arm. 
As a rule the disease appears only on branches or on the main 
axis of the younger classes of silver firs, and attracts atten- 
tion by the cortex dying right round the branch, as is shown 
in Fig. "j"^. 

Numerous small black pycnidia break through the epidermis 
and appear on the dead cortex, either as small roundish bodies 
or as many-chambered, irregularly shaped, black sclerotium-like 
tubercles (Fig. 79, a). Numerous unicellular, colourless, abruptl\- 



Fig. 79.- 


spindle-shaped gonidia (Fig. 79, b), which at once germinate in 
water, develop on the h}-menial layer that lines the walls of the 
cavities of these organs. 

Although I have watched the disease every year since 1885, 
and have sought for the ascophores, I have hitherto been unable 
to find them. I may, however, remark that in almost all my 
cultures on silver fir branches a luxuriant growth of the 
ascocarps of Peziza calycina has appeared upon the cortex on 
both sides of the diseased part. This fact, however, is not 
sufficient proof of a connection between 
these two fungus-forms. Attempts to 
produce the one form from the other 
by cultural experiments have so far 
proved abortive. 

The pycnidia ejaculate the gonidia 
probably for the most part during wet 
weather in summer and autumn. 

It does not appear necessary that 
mechanical injury of the cortex should 
precede the entrance of the parasite — 
at all events, I have never been able 
to observe such. On old trees a large 

proportion of the twigs and branches are often brown, a state of 
things that struck me at once on my first visit to the Bavarian 
Forest. In the Black Forest also, and at several places in the 
Bavarian Alps, the disease is to be met with. In the case of 
the thicker branches nutrition through the wood may still be 
continued for several years after the cortex has died. For this 
reason growth in thickness above the dead part is distinctly 
visible, and causes the cortex to rupture at the boundary of the 
living and dead parts. When the wood covered by the dead 
cortex dies and dries up, the passage of water ceases, and the 
branch dies above the seat of the disease. 

Should the fungus attack one side of the branch only, the 
dead cortex is exfoliated, and the formation of callus commences 
along the healthy margin. 

of p. abictiim which has 
ruptured the periderm ; 
magnified twenty times. 
/', gonidia magnified 420 




Plane-trees i^Platamis) suffer very frequently from a disease 
which is characterised by the leaves acquiring brown blotches 
and dying. From the middle of May onwards one observes 
that death sets in at certain places, and continues along the 
nerves of the leaf. Small black spots may then be observed 
appearing on the dead parts, which are the gonidial cushions of 
Gloeosporiuin nerviseqiLUivi. 

Unfortunately we still know very little regarding the develop- 
ment of this fungus, for even trials at infection have not yet 


For a number of years a disease of the black (Austrian) pine 
has been observed in the south of Norway and throughout the 
whole of Germany. This disease has constantly been on the 
increase, but has not yet been thoroughly investigated. It is 
now a considerable number of years since Dr. C. v. Fischbach 
sent me 'diseased branches, and an opportunity for observing 
the disease was afforded in the forest division of Freising, near 
Munich, but the investigation yielded no satisfactory results. 
This disease may find mention here, especially since Dr. Brunc- 
horst's description is now available. 

The most vigorously growing Austrian pines show a pale- 
ness in the leaves of the previous year's shoots, whose buds, 
instead of shooting out, die off. The disease spreads from the 
tissues of the shoots, having its inception in the cortical tissues. 
Here infection is very often brought about, as it appears to me, 
through the agency of a small plant-mite, which bores through 
the epidermis into the cortical tissues of the shoot to a depth 
of I — 2 mm. Infection may, however, also take place easily 
enough at the base of the leaves, where the epidermis is thin. 

1 Dr. Fr. v. Tavel, Bot. Zeit., iS86, No. 49. 

- Dr. C. V. Fischbach, Eiiic ncue Kraitkhcit der Schwarskiefer. Central- 
blatt fiir das gesammt Forstivesen, 1887, p. 435. Dr. Brunchorst, Ueber eifie 
jicicc verheerende Krankheit der Schwarsfohrc. Bergen, 1888. 


Black pycnidia, with gonidia similar to those of Fusidium, 
develop at the base of the dying leaves, and on the wounds that 
result from the separation of the leaf-fascicles. 

Dr. l^runchorst has not yet been able to observe perithecia, 
nor has he hitherto succeeded with infection-experiments. Not 
only has the death of single pines been noticed, but in many 
cases, especially in Norway, the destruction of large woods has 
been recorded. 

It is strongly to be insisted upon that whenever this disease 
appears in young woods of the Austrian pine, all diseased shoots 
should be cut off and burned. 


In the neighbourhood of Munich the branches of the English 
tnaple {Acei- caiiipestre) suffer from a disease which kills them 
off before the young shoots develop in spring. In the middle 
and lower parts of the crown especially, it frequently happens 
that more than half of the previous year's shoots perish. On 
these it will be found that the periderm has been ruptured 
by oblong cushion-like fungus-bodies. 

The disease almost always confines itself to the youngest 
shoots, the two-year-old shoots being infected only in very 
exceptional cases. Infection takes place in May and the 
beginning of June, when the young shoots are still tender and 
unprovided with periderm. When the spores of the parasite 
(Fig. 80, 6) come into contact with a young shoot, they germinate 
within a few hours. The spore represented in the figure had lain 
in water for five hours only, at the end of which time it showed 
large germ-tubes at both ends. The mycelium bores into the 
cortex, and takes possession of the shoot for a distance of 2 — 4 
inches, but does not kill it in the same year. Even when the 
leaves are shed in autumn there are no external symptoms of 
disease. In spring, the buds of the diseased shoots swell up, 
as a rule, but soon wither. At that time the mycelium is to be 
found growing vigorously, not only in the diseased cortex, but 
also in the medullary rays and the vessels of the wood. It 
grows both intercellular and intracellular, and pushes numerous 

1 R. Hartig, Forstl. N'aturwiss., Zeitschrift, August 1892. 



short stout lateral branches, which resemble haustoria, into the 
interior of the parenchymatous cells. 

Colourless stromata of a fleshy, pseudo-parenchymatous 
structure (Fig. 80, j) form in the cortex beneath the periderm. 
In cross section these measure 0'3 — 0'6 mm., while longitudinally 
their length varies between i and 4 mm. In the month of May 
the periderm ruptures longitudinally, and reveals the sporogenous 

Fig. 80. — Septoglceum Hartigianum. 

.ayer as a greyish green cushion, surrounded by the edges of the 
elevated periderm (Fig. 80, 2). The surface of the colourless 
fleshy stroma (Fig. 80, ^) is formed of cylindrical sterigmata, 
which frequently enlarge somewhat towards the base. These 
sterigmata measure 30 — 35 micromillimeters in length and 
6 — 7-5 micromillimeters in breadth. The gonidia, which are 
formed at the apex of the sterigmata, measure, when ripe, 
from 24 — -36 micromillimeters in length and 10 — 12 micromilli- 
meters in breadth. In shape they are irregularly oblong-oval, 
and truncated at both ends (Fig 80, 5). 


In the great majority of cases the gonidia appear to be doubly 
septate, ahhough occasionally one meets with singly septate 
examples, and even unicellular gonidia are not unknown. They 
are of a pale brown colour, and germinate in a few hours, 
pushing out a thick germ-tube from both ends (Fig. 80, 6). 

The parasite is distributed in May and the beginning of June 
by means of the gonidia, which are washed by the rain from 
the higher diseased shoots on to the young shoots of the lower 
part of the crown. In other cases the wind may carry them on 
to distant maples. 

This destructive parasite of our gardens and parks can only 
be combated by removing the diseased shoots from the crown in 
the beginning of May. 


Both in young spruce woods and in the seed- and plant-beds 
of the nursery a disease, which on a cursory examination may 
easily be mistaken for damage by frost, very frequently attacks 
the young shoots. 

In the month of May, when the young shoots are still succu- 
lent and delicate, the disease generally manifests itself (see 
Fig. 81, a) by the leaves at the base, or it may be the middle, of 
the shoot becoming brown and soon dropping oft". At first the 
apex of the shoot remains quite green, but on lateral branches 
it droops downwards. The disease advances rapidly towards 
the apex of the shoot, and the base of the young leaves appears 
dark green (Fig. 81, d). When held up to the light, it will be seen 
that the internal tissues of the leaf are dead and shrivelled, and 
possess a reticulated appearance. Finally the leaves all drop 
off", or a few dead ones may adhere to the apex of the shoot, as 
indicated in Fig. 82, a. The axis of the shoot shrinks more or 
less according as death had overtaken it in an early or late stage 
of development. Very frequently the shoot becomes diseased 
and begins to shrink at the point where its base is enveloped by 
the scales of the previous year's terminal bud, and this often 
occurs in the case of shoots that are tolerably well developed. 
The base of the shoot shrinks, and its internal tissues are so dis- 

1 R. Hartig, Zeitsclirift fiir Forst- und Jagdivesen, November 1890, p. 668. 



organized by the fungus which causes the disease — as we shall 
see presently — that the shoot bends sharply over at that point 
under its own weight, as though it had been fractured at the 
base. Very frequently death spreads back from the base of the 
youngest shoot to the apex of the shoot of the previous year 
(Fig. 8 1, r), in which case the young lateral shoots that are 
situated there also succumb (Fig. 82, a). 

As already indicated, this may, in May or early in June, be 

mistaken for damage due 
to frost, although the latter 
is generally confined to cer- 
tain localities, appears sud- 
denly on a great number 
of shoots, and is character- 
ised by all the affected 
leaves dying simultaneous- 
ly. The disease at present 
under discussion has no re- 
lation whatever to frost. 
Shoots that have succumb- 
ed to this disease also bear 
a certain resemblance in 
many cases to those dam- 
aged by Chermes abietis ; 
the galls which the latter 
induces at the base of the 
shoots being often so small 
as not to be visible on the 
upper side, and in both 
cases there is a deflexion of 
the shoot downwards. 
In the course of the summer the dead shoots display spore- 
receptacles (pycnidia) in greater or less abundance in the form 
of very minute black tubercles. These are so small as just to be 
visible to the naked eye, and are very often to be found only 
amongst the bud-scales at the base of the dead shoot. In other 
cases they are also to be met with higher up ; in fact, they are 
often specially abundant on the shrunken shoot-apex (Fig. 82, a)- 
They either rupture the epidermis of the shoot or occur on the 

Fig. 81. — (T, a young diseased spruce branch, 
the apex of which is still green and fresh. 
h, a leaf attacked by disease towards the 
base, twice natural size, c, apex of a 
two-year-old shoot, into which the disease 
has spread backwards from the younger 
shoot. The brown discoloration of the 
cortex and pith is indicated by shading. 



leaf-scar on the long pulvinus, where they present a bud-like 
appearance (Fig. 82, b). It also frequently happens that black 
pycnidia develop on the few dead leaves that have adhered 
to the shoot (Fig. 82, a).. a 

These pycnidia, which are 
uni- or multicellular, pro- 
duce numerous small go- 
nidia (stylospores) at the 
apex of subulate sterig- 
mata which spring from 
the inner wall. These 
stylospores, which are bi- 
cellular, colourless, spindle- 
shaped, and some 13 — 15 
micromillimeters in size 
(Fig. 82, r), appear as white 
tendrils on the pycnidia in 
the month of May, at which 
time they produce the dis- 
ease, if borne by the wind or 
rain to the still tender and 
non-cuticularized shoots of 
the spruce. 

On May 6th, when growth 
was active, I infected young 
spruces by taking a drop 
of water, in which stylo- 
spores were suspended, and 
placing it by means of for- 
ceps amongst the leaves in 
the middle of a shoot and 
partly amongst the bud- 
scales at its base. Infection 
succeeded in every case, 
and eight days later death 
appeared amongst leaves at 
the infected points, and 
soon spread in all direc- 
tions. On the 13th of May, 


82. — a, a spruce-leader of which the 
youngest shoot, the apex of the older 
shoot, and the two lateral branches have 
been killed. /', pycnidia projecting from 
the cortex and leaf-scars, magnified five 
times, c, formation of spores in the inte- 
rior of a pycnidium, magnified by 400. d, 
spores germinating in water. e, spores 
germinating in nutritive gelatine. 



a drop of water containing spores was placed amongst the 
scales at the base of a shoot some three inches long, on 
a twenty-year-old spruce. About twelve days later the shoot 
was so much diseased as to droop in the manner shown in 
Fig. 81, a. 

Spores sown in water on May 6th showed the first symptoms 
of germination in eighteen hours (Fig. 82, d). When sown in 
nutritive gelatine the spores and germ-tubes were rather more 
vigorous (Fig. 82, e), an extremely luxuriant mycelium with 
spherical and clavate segments being developed both on a 
microscope-slide and in a test-tube. Twelve days after 
sowing, pycnidia with ripe stylospores developed on this 

The culture, which was continued till August 12th, produced a 
dense mycelial growth, with pycnidia but no perithecia, so that 
at that time I abandoned the hope of obtaining the latter. An 
investigation of the diseased shoots showed that the stout vigor- 
ous mycelium developed between the cells in the green leaves, 
and immediately induced death in adjacent cells. In the shoot 
axis it penetrated all the tissues, growing as both intra- and 
intercellular filaments in the pith and cortex, while in the wood 
it was specially abundant in the annular and spiral vessels. The 
development of the parasite is confined to the short period 
between the beginning of May and the early part of June. 
Whether the pycnidia will form early or late in summer appears 
to depend on the humidity of the air, or, in other words, on the 
occurrence of dry or wet weather. As already stated, they may 
ripen in fourteen days, provided the conditions are favourable, 
as m the case of artificial cultures. 

Until we obtain the perithecia we must rest satisfied with a 
provisional name for this new parasite. The character of the 
pycnidia and spores leads us to place this fungus in the genus 
Septoria, but as there is already a Septoria Pini I have named 
this fungus Septoria parasitica. Finally, it may be mentioned 
that in the spring of 1 890 I found this parasite on Picea Menziesii, 
and this makes it probable that it also occurs on other species 
of spruce. 



Seed-beds of the pine and spruce are frequently subject to 
disease in the months of May and June, so that, even where 
seed has germinated satisfactorily, the plants are more or less 
decimated, and the otherwise well-stocked drills show blanks as 
large as one's hand. During wet weather the young plants die 
and rapidly decay. At first only single plants are attacked, but 
soon the disease spreads more or less rapidly along the whole 
drill. If the weather is dry the diseased plants wither, their 
yellow colour attracting attention even from a distance. During 
summer, say from the middle of June, the disease ceases, and 
the dead plants disappear, to leave only the blanks, which are 
usually ascribed to the ravages of cockchafer grubs, crickets, 
surface caterpillars, &c. I have shown that in most cases these 
pathological phenomena are due to PhytoptJioj-a omnivora, which 
spreads not only by ^onidia but also by its mycelium, which 
traverses underground from root to root, and thus explains the 
rapid spread of the disease. 

In 1889 I received a number of diseased pine seedlings from 
Herr Mantel, forester in Grossostheim, with the information that 
for some years a disease agreeing with the symptoms just 
described had appeared immediately after the germination of the 
seed in his pine seed-beds on sandy land. Some plants which 
he sent to the forest school of Aschaffenburg were examined, 
when it was found that, instead of P. omnivora being the cause 
of the damage, it was some other unknown disease. 

An investigation of this new disease showed that it was 
induced by a different parasite. The mycelium of the parasitic 
fungus which caused the damage attacked not only the seed- 
lings of the pine but also those of the spruce, alder, birch, 
&c. In the seed-bed the plants were attacked either at the 
roots (Fig. 83, a) or at the hypocotyl {b), close beneath the 
surface of the ground. In very dense seed-beds and during wet 
weather the mycelium also spreads above ground, and infects the 
cotyledons and the highest part of the stem (Fig. 83, c). At 
the point where it comes into contact with the epidermis of 

^ Forstlich Naturwissensch. Zeitsch., November 1892. 

L 2 


the seedling the filamentous septate mycelium, which becomes 
somewhat brown with age, produces tortuous lateral hyphae, 
which ramify abundantly, apply themselves closely to the epi- 
dermis (Fig. 84, a), and exercise a solvent action on the delicate 
non-cuticularized epidermis. If one lifts up the hyphae it will 
be seen that the epidermis has been dissolved at the points 

of contact. Without doubt mycelial 
filaments also bore directly from these 
points right into the plant. 

The stomata form a means both 
of ingress and egress for the hyphae. 
Such a stoma is depicted in Fig. 85, 
which shows that the sides of the 
depression leading to the stoma, and 
the external walls of the epidermal 
cells, are dissolved at the points where 
the hyphae are or have been in close 

Fig. 83. — Diseased pine seedlings. 

a, a specimen with dead roots ; 

b, ditto with a dead stem ; 

c, ditto with dead leaves and 

Fig. 84. — A filamentous mj'celium whose 
lateral hyphpe, a, come into close con- 
tact with the epidermis ; b, a mycelium 
which has developed in a nutrient 

contact. These places appear granulated because the ash con- 
stituents of the cell-wall are either left wholly intact or are but 
partially dissolved. I have proved that under the action of the 
ferments exuded by wood-destroying parasites the cell-walls also 
display granulation during the last stages of decomposition, and 
for the same reasons as those just given. When a diseased 
plant is investigated on the first symptoms of attack, a vigorous 
growth of mycelium will be found in all the tissues. Although 
the green cells do not part with their chlorophyll till some time 
after death, they easily lose connection with each other, and the 



stem or root becomes limp. At this stage of the disease the 
interior of the plant is nearly full of a luxuriant mycelial growth. 
In a short time large numbers of bacteria appear in the tissues 
and induce complete decomposition, the mycelium of the para- 
site sharing the common fate. In plants that still appear sound 
towards the top, the stem or roots frequently retain only the 
epidermis and the xylem of the vascular bundles. I have 
infected vigorous pine and spruce seedlings, which were taken 
from dense seed-drills and planted in flower-pots, by laying 
one or more diseased plants amongst them. When a bell- 
glass was placed over the flower-pot all the plants were 
diseased or dead in four days. 
The mycelium enveloped the whole 
plant, and the disease began, for 
the most part in the upper portion 
(Fig. 83, c). On the other hand, if 
the flower-pot was left in a room 
uncovered, infection was induced 
only by the mycelium growing in 
or on the surface of the soil and 
attacking the roots or the lower 
portion of the stem (Fig. 83, a, b). 
All the plants succumbed in eight 
days, with the single exception 
of one seedling which remained 
healthy. No results followed trials 
at infection which I conducted in 
the end of June on vigorous seed- 
lings. As has already been proved in the case of other parasites* 
it is only when the epidermis is unprovided with a cuticle that it 
can be dissolved by the mycelium. It is known thz-t P . onuiivora 
also is only destructive in May and June. 

Innumerable gonidia develop in dense bunches on the 
luxuriantly branching mycelium, and especially in the stomata 
of the diseased plants. I have represented such a mycelial 
branch with gonidia in Fig. 86. When ripe they are more or 
less falcate, pointed at both ends, and generally consist of six 
cells. In germinating they usually produce two germ-tubes at 
or near the apices, as is shown in the lower part of Fig. 86- 

Fig. 85.- — Epidermis of a pine 
seedling showing a stoma. 
Solution has taken place 
wherever the filaments have 
been in contact. 



The shape of the gonidia makes it probable that the parasite is 
a species of Nectria. A few days after being sown in a gelatine 
extract of fruit they produce a luxuriantly branching mycelium, 
whose hyphae are much septated and anastomose irregularly 
(Fig. 84, ^). This mycelium produces either similar gonidia, or 
such as are somewhat smaller, less bent, and possessed of fewer 

On being transferred to black bread the mycelium grew so 
vigorously that the large glass jar in which the culture was 
conducted was completely filled with a white growth. In the 
flower-pots also, in which the infected conifer seedlings were 
growing, the mycelium developed so luxuriantly in the soil as 
to find its way out at the hole in the bottom and to form a dense 

mass between the pot and the table. 
This proves conclusively that the 
fungus, like most species of Nectria, 
can also exist as a saprophyte, and 
as such may live in the soil. 

Unfortunately I have not been 

successful in reproducing the peri- 

thecium form of the fungus. On 

a slide numerous spheroid bodies 

which were the first stages of peri- 

thecia or pycnidia formed in the mycelium, but they always 

failed to mature. On this account I am unfortunately still 

unable to refer the fungus to a species. 

As regards measures that may be instituted with the view of 
preventing the spread of the disease, attention should in the first 
place be directed to the diminution of surplus moisture. If 
such are present, we should also remove all objects such as latticed 
frames, branches, &c., that have been laid down to shelter the 
plants. As it is certain that the fungus remains in the ground 
from one year to another, one must take care, in laying down 
new seed-beds, to avoid situations where the disease was 
prevalent in the previous year. If this cannot be done, one 
should interstratify the upper nine inches of soil with brush- 
wood, dry turf, or some such material, and roast it, or at least 
raise its temperature to such a pitch that any spores present in 
it may be killed. Herr Mantel related to me how he had heated 

Fig. 86. — Immature, ripe, and 
germinating gonidia. 


the soil of certain beds by burning dry wood in parallel trenches 
a foot deep and about a foot apart. The fire was kept up for 
two days, so that the soil was practically roasted to the depth of 
more than a foot. Seed was afterwards sown on these beds as 
on the others. On the beds so treated the plants remained 
sound for two years, but in the third year the old symptoms 
reappeared. It is most likely that the fungus again invaded the 
ground by means of gonidia carried from adjoining beds. It 
would appear desirable, should circumstances make it advisable, 
that the above plan should be experimentally tested. One 
would thus discover whether infested seed-beds may continue 
to be utilized without incurring much expenditure, or whether 
it is better to change the ground. 


In Alpine districts a disease is very prevalent on Alnus 
viridis, which, superficially examined, reminds one strongly of 
the ravages of the larvae of CryptorhyncJms lapathi. Numerous 
stems and branches contract the disease and die. It is chiefly 
in August that the leafy branches become infected. The 
withering of the cortex attracts attention to the presence of the 
fungus, and directly afterwards small black tubercles appear 
on the dead tissues. The stage of the development of these 
pustules depends upon the length of time that has elapsed since 
the branch died. Thus, although the presence of the fungus may 
be detected on branches that are still living, it is met with in its 
highest development only on such as are perfectly dead. The 
progress of the disease down the tree is indicated by a sharp line 
between the diseased wood, which is brown, and that which is 
still sound. As the disease advances, other lateral branches 
become affected. An exceedingly vigorous and very tough 
mycelium is easily discoverable in the moribund wood, and 
especially in the vessels. The lenticular tubercles consist of 
black pseudo-parenchyma situated beneath the periderm. 
Owing to their rupturing the periderm at the highest point 

1 \'.T\ihe\xi,Forstltch-Jtatur'wis. Z^//Jt7w7//, October 1892. 



of their concave surface, a small roundish aperture is formed. 
Numerous perithecia, with very small tubes and hyaline spores, 
form in the dead cortex underneath the lower surface of the 


Fig. 87. — The upper left-hand figure represents a portion of a branch of Almis viridis. 
The periderm has been ruptured at four places by the stroma of Valsa oxystotna, 
on which the necks of the perithecia are visible. To the right are seen the asci 
and spores. The upper right-hand figure represents a piece of a smaller branch 
whose periderm has also been ruptured by the stromata, but on which the peri- 
thecia have not yet developed. The central figure shows the microscopic section 
of one of the stromata of the upper left-hand figure, and the lowest figure a similar 
section taken from the branch shown in the upper right-hand corner. 

lenticular tubercles. The perithecia push a long stout flask- 
shaped neck through the lenticular stroma. On account of the 
black necks finally protruding to some extent from the stroma, 
each tubercle shows a considerable number of them. 



The Basidiomycetes constitute the third group of fungi. In 
their case ah spores originate by abscission. 


The rust-fungi are true parasites, and develop their mycehum, 
which is usually intercellular, in the tissues of the leaves and 
cortex (and also, though less frequently, in the wood, e.g. Cole- 
osporiuin SenecioJiis) of phanerogams, and abstract their nutri- 
ment by means of haustoria from the interior of the cells. 
Their course of development is characterised by most species 
producing sporocarps, which are usually cup-like in shape. The 
bottom of these so-called secidia is lined by a hymenium 
consisting of numerous, usually club-shaped, basidia, each of 
which at its apex abjoints a series of spores which are usually 
reddish yellow in colour. These are united to each other by so- 
called intermediate cells, which are dissolved before the forma- 
tion of the spores is completed. The basidia that are situated 
at the periphery of the hymenium, instead of forming spores, 
grow together to form an envelope called the peridium, which 
opens at the apex or by a longitudinal fissure. The peridium 
may, however, be entirely absent. 

Before the formation of aecidia, spermogonia with spermatia 
usually originate, the latter probably playing the part of male 
sexual cells.-|- It is probable that the aecidium is the result of a 
preceding sexual act, and is therefore a true sporocarp, like the 
perithecium and apothecium of Ascomycetes. However, there 
are also rust-fungi in which the aecidium is entirely absent 
{CJirysomyxa Abietis). 

* [The British species have been worked up into a monograph by Plowright 
{BriL Uredinecc and Ustilaginece, Kegan Paul, 1889), who has also devoted 
attention to the experimental investigation of some of the heteroecious 
forms. — Ed.] 

t [Researches into the morphology and physiology of these and other 
organs of the Uredine£e by no means support this conclusion. Brefeld's 
statements, as well as those of De Bary, much as they differ in detail, point 
to the opposite view — that there is no probability of sexuality in this group. 
The curious and interesting homologies are well put forth by Von Tavel.— 


Besides the secidium, a kind of gonidium is almost always 
formed, which, being designed to carry the fungus over from one 
year to another, has great capacity for retaining its power of 
germinating. This is known as a resting-spore or teleutospore, 
and, instead of directly forming a mycelial filament, it first pro- 
duces a promycelium, on which a number of small cells, called 
sporidia, develop, and it is these which produce the disease by 
infecting new host-plants. The teleutospores are unable to 
produce infection, because they are usually in such intimate 
contact with the substance of the host-plant that their distribu- 
tion by the air is almost entirely precluded. The mycelium that 
is developed from the sporidia again produces spermogonia, and 
— after fertilization — sporocarps (aecidia). Thus an alternation 
of generations between the two forms of aecidia and teleutospores 
is presented, which, however, in the case of many rust-fungi, is 
further complicated by the fact that a form bearing teleutospores 
is not directly produced by the germinating aecidiospores, but 
that numerous generations of gonidia of another kind — namely, 
uredospores — often originate. These at once germinate, without 
forming a promycelium, and reproduce the form bearing uredo- 
spores. During summer they serve for the rapid distribution of 
the fungus, till the teleutospores are produced by the mycelium, 
as usually happens in autumn. The cycle of development of 
many rust-fungi is interesting, from the fact that not only the 
uredo form but also the aecidium form may possess a facultative 
character ; that is to say, these forms may develop only under 
certain favourable circumstances, and in the absence of such 
conditions they may be entirely omitted without the existence 
of the parasite being thereby imperilled. 

The generation which forms the aecidia, and that which pro- 
duces the teleutospores, are either to be found on the same plant 
(autoecious parasites), or, with the alternation of generation, 
there also occurs a change of the species of host-plant (heter- 
oecious parasites). In the case of the hetercecious rust-fungi the 
discovery of the related forms which belong to one and the same 
species of fungus naturally presents great difficulties. This is 
sufficient to explain why we are not at present acquainted with 
the aecidia of many teleuto forms, and, on the other hand, do not 
yet know to which teleuto forms many aecidium forms belong. 


On this account it will be necessary, as in the case of the 
Ascomycetes, to append a list of imperfectly known rust-fungi, 
to which, depending on the development form, the provisional 
names of Aicidimn, C<2oma, and Uredo are given. 

The rust-fungi are divided into several families, of which we 
are here interested only in the PuccinicB and MelampsorecE. The 
former are characterised by the teleutospores being situated 
singly, or several together on a stalk ; whereas, in the case of 
the latter, the teleutospores are united to each other in large 
numbers to form a firm palisade-like layer. 


The genus Piiccinia, which is rich in species, is characterised 
by the teleutospores containing two cells and remaining attached 
to the basidia, which at the same time serve as the stalk. They 
appear as small brown or black brown heaps of a round or 
oblong shape. 

Piiccinia graminis * is the commonest kind of rust amongst 
cereals, occurring everywhere, not only on our various kinds of 
grain but also on many grasses. The teleutospores, which are 
disposed in narrow ridges, hibernate on the common grasses, 
though, if they have been produced on the lower parts of the 
stems of cereals, they may also be left on the stubble fields. In 
spring, when the sporidia that originate on the promycelia gain 
a footing on the young leaves of the barberry, Berberis viilgaris, 
they produce the barberry fungus, ^ciditun Berberidis. The 
aecidium form — whose spores germinate on cereals and other 
species of grasses, and produce the wheat-rust, Uredo linearis — 
is distinguished from the black ridges of teleutospores of 
Piiccinia graini?iis, which occur later, by the reddish brown 

This destructive cereal rust may be most effectively combated 
by rooting out the barberry. This measure must not, however, 
be confined within narrow limits, because the spores of the bar- 
berry-fungus can be widely distributed by wind. 

*[This, the common rust of wheat and other grasses, is of classical 
interest, since it was this species in which De Bary first estabUshed the 
remarkable phenomenon of hetercecism. — Ed.] 


Piiccinia stricEforviis {strainifiis) produces a cereal rust on rye, 
wheat, and barley. It closely resembles the foregoing disease, 
from which it differs, however, by the ridges being smaller 
and less elongated, and by the very short-stalked club-shaped 
teleutospores remaining hidden by the epidermis. Theaecidium 
form is ^cidhun asperifolii, which develops on the leaves of 
AncJmsa officinalis, Borago, EcJiiuni, &c. 

Piiccinia coronata produces a rust on cereals, especially oats. 
Its teleutospores are provided at the shoulder with a girdle of 
punctiform thickenings of the spore membrane. The aecidium 
form y^cidiiim Rhanini is well known by the peculiar rich golden 
yellow swellings which it produces on the leaves, flowers, and 
stalks of Rhamnus catJiartica and R. frangula in which it 

Of the large number of species of Puccinia, the only other one 
that need here be noticed is Piiccinia Asparagi, which completes 
its course of developmen t on the asparagus alone. The asparagus 
rust, which may seriously devastate asparagus-beds, is best com- 
bated by burning the halms in autumn, and by the timely 
removal of the shoots that are first diseased. 


The species of this genus are distinguished from the species 
of Piiccinia by the teleutospores being stalked and consisting 
of a number of cells. The groups of teleutospores which 
develop on the under side of the leaves are preceded by uredo- 
spores, whose orange red powder often covers the under side of the 
leaves in large quantity. The course of development of the 
various species has not yet been sufficiently studied.* 

PJiragniidiiun incrassatinn, the rust of the bramble, induces the 
formation of red blotches on the leaves of Rnbus fniticosiis and 
R. ccesiiis, and the organs consequently die prematurely. 

Phraginidiinn Rnbi Idcei produces similar pathological sym- 
ptoms on the leaves of Rubiis Idceiis. 

Phragviidinvi siibcorticiun produces the rust of the rose. 

*[A11 occur in this country. — Ed.] 



The species of this genus with which we are acquainted arc 
perennial in the cortical tissues of various species of Jimiperus. 
They induce local increase in growth, which takes the form of 
peculiar swellings on the branches and parts of the stem that are 
attacked.* Each autumn the teleutospores are developed under 
the outer cortical layers, and in spring and early summer they 
break through the cortex in large numbers and appear as fructifi- 
cations which are conical or sausage-shaped, yellow or brown, and 
mucilaginous or cartilaginous in texture. These fructifications 
consist of the very long filamentous basidia whose outer wall 
has been converted into mucilage, and of the two-celled resting- 
spores which they bear at their apex. The formation of the 
promycelium and sporidia takes place in the mucilaginous mass, 
which in the end is completely dissolved by rain-water. The 
sporidia gain a footing on the leaves of various pomaceous trees, 
where they produce the aecidium form of the genus RcEstclia. 

It appears to me desirable that the forms which are already 
known and described should be subjected to further examination, 
because the few test trials that I have undertaken have at once 
led to results which are at variance with what has been accepted. 
I append here a short description of the three recognised species, 
without, however, being able to vouch for its accuracy on the 
strength of my own investigations.-|- 


Teleutospore layers on Jiinipenis connminis. They are hemi- 
spherical or conical, golden yellow, later swelling up to very 
large, variously shaped (spherical, pear-shaped, ovate, &c.) bodies. 
Spores spindle-shaped, some brown with a thick endosporium, 
75 microm. long and 27 microm. broad ; others yellow, with a 

1 Oersted, Bot. Zeit., 1865, p. 291 and elsewhere. 

* [These are often called Cedar-apples in America. See Farlow, Mem. 
Boston Soc. of Nat. Hist., 1880. — Ed.] 

f [A good deal of work has been done on the various and very confusing 
species of late years, most of which occur in this country. See Plowright, 
/.^,and Von Tubeuf {Cent. f. Bakt., 1891, and Zeitschr. f. PJlansenkrankh., 
B. II. p. no). Also Farlow, t.c, &c. — Ed.] 


thinner endosporium, about 66 microm. long and 17 microm. 
broad. The aecidium form has been observed as Rcestelia 
corniita on Sorbiis Aiicuparia, S. tonninalis, Aronia, and other 
pomaceous plants. The secidia are situated on orange yellow 
or red swollen blotches, which are united in various numbers 
into round or oblong groups. The peridium, which has the 
shape of a very long-necked bottle, is yellowish, or yellowish 
brown, twisted like a horn, up to 8 mm. long, open at the 
shoulder, serrulated, and either not or only ultimately slightly 
and irregularly lacerated. 


Teleutospore-layers on Jiiniperus commiinis. They are 
cylindrical, tongue- shaped or band-like, often bifurcated, 
twisted, and bent, somewhat cartilaginous in texture, yellow. 
and up to 12 mm. long. Spores spindle-shaped, contracted at 
the middle, bright golden brown, 70 — 120 microm. long, and 
14 — 20 microm. thick. The aecidium form, RcBstelia lacerata, 
is met with on species of CratcBgus, and occurs abundantly 
in smaller or larger groups on orange yellow swollen blotches 
— -though frequently covering large areas, especially on fruit — 
and is usually accompanied by contortions and other deforma- 
tions. Peridia, flask-shaped when young, later cylindrical cup- 
shaped, dirty white, rupturing longitudinally to various depths 
into numerous erect or somewhat outwardly inclined lobes. 


Teleutospore-layers on Juniperus Sabina, J. virginiana, J . 
pJicenicea^ J. Oxycednis, and Finns halepensis. When fresh they 
are abruptly conical or cylindrical, often slightly compressed 
laterally and expanding somewhat towards the top, sometimes 
pectinate, red brown, 8 — 10 mm. long. Spores broadly 
elliptical, either not contracted at the middle or contraction 
scarcely observable, chestnut brown, 38 — 50 microm. long, and 
23 — 26 microm. thick. The aecidium form, called RcBstelia cancel- 
lata, is found on Pyrus communis^ P. Michauxii, and P. tomentosa. 
In shape the secidia are like very short-necked bottles, about 
2 — 2\ mm. high, and several are situated together on orange 


yellow, roundish or irregular, cushion-like swollen blotche?. 
Pseudo-peridium yellowish white, closed at the shoulder, 
ruptured on the side by numerous longitudinal fissures, which 
extend to the surface of the leaf. These longitudinal fissures 
are bridged over by short transverse rodlets, whence the whole 
peridium appears grated. In this connection I may remark that 
I have repeatedly observed the pear-rust in great abundance 
in places where no examples of the above-mentioned host- 
plants of the teleuto form were to be found within a wide 


To the three above-mentioned species a fourth falls to be 
added, whose aecidium form is very abundantly met with in 
the Bavarian Alps on Sorbiis Aria and 6". CJiamcBinespilus, and 
which as ^cidium pejicillatum has already been described as an 
independent form (Fig. 88). 

In equal abundance one meets in the same region with a 
teleuto form on Jmiipenis coniiminis which does not agree with 
any of the above-named species, but whose connection with the 
aecidium form on Sorbiis Aria has been proved by infection- 
experiments in the garden of the Munich institute of forest 

The teleutospore-layers appear in May on Juniperiis com- 
inunis, as hemispherical orange yellow to yellowish brown masses, 
which, as in the case of Nostoc coimminis, are mucilage-like 
in texture and capable of swelling (Fig. 89, a a). They 
easily drop off when the branches are shaken, and then pale 
yellow smooth scars remain, which are often i cm. in diameter 
(Fig. 89, b b). The spores are all about the same size, being 
approximately 40— 45 microm. long and 20 — 25 microm. broad. 
The two short abruptly conical cells, whose height is about 
equal to their greatest diameter, are provided with smoky- 
grey walls. Some of them coalesce all along their base, others 
are to a certain extent separated owing to contraction — in fact, 
it not unfrequently happens that the two parts of a teleutospore 
become completely disunited. Most of the cells possess three 
germ-pores, which, when situated near the transverse septum, 
frequently alternate with those of the second cell (Fig. 90). 



The secidia appear on Sofbus Ai-ia, S. Chamcsmespilus, Pyrus 
Malus, and Sorbiis tormmalis (?). 

The cushion-like stroma, on which the aecidia are often 
arranged in circles, is very thick, and luxuriantly developed. 
The pseudo-peridia are somewhat cup-shaped, and are split 
up as far as the base into a large number of filaments i mm. 
in length, which bend out somewhat. The aperture of the 


88.— /Ecidia of 

trenielloides on 

leaf of Sordiis 


Fig. 89. — G. trenielloides on 
J. communis, a a, teleu- 
tospore-layers ; b b, scars 
which are left after the 
mucilaginous mass has 
dropped off. 

Fig. 90. — Teleutospores of 
G. trenielloides. a, basi- 
dium ; b, spore that has 
not yet germinated ; c, a 
similar spore, showing 
contraction along the 
centre ; d, ditto with the 
cells separated ; e, a 
germinated teleutospore 
with promycelium and 
sporidium ; /, basal view 
of a teleutospore showing 
three germ-pores, the 
one from which the germ- 
tube is issuing being 
closed by a lamella. 

jEcidium is distinct, and is black in colour owing to the dark 
spores. Material sent by Herr Nawaschin, of Moscow, shows 
that this species also occurs in Russia, where the teleuto form 
develops not only in the cortex but also on the leaves of 
Junipcnis communis, forming an oblong cushion-like stroma 
which extends over about half of the leaf. In that country the 
aecidia occur on the leaves of the apple-tree. 




The fungus which attacks the 
cowberry, and its aecidium form, 
yEcidiiun coluniuarc, which pro- 
duces the columnar rust of the 
silver fir, are indigenous wher- 
ever silver firs abound. The 
first-mentioned form, indeed, is 
also met with in districts from 
which the silver fir is absent, and 
this furnishes a proof that the 
oscidium form possesses only a 
facultative character. 

Specimens of Vacciniinn Vitis 
Idi^a that are attacked by the 
parasite are at once distinguishable 
from healthy plants by their man- 
ner and habit of growth. Whereas 
the latter rise but a short distance 
above the ground, individuals that 
are infested by the fungus grow 
quite erect, display an unusually 
vigorous height-growth, and de- 
velop two shoots even in the same 
year. The diseased plants, singly or 
in groups, tower above the healthy 
plants, whose height they exceed 
in some cases by a foot. At the 
same time they exhibit a striking 
appearance, the greater part of the 
stem being swollen to the thickness 
of a quill, while only the upper 
part of each shoot retains its normal 
dimensions (Fig. 91). At first the 
thickened spongy part of the stem 
is of a white or beautifully rosy 
red colour, which soon, however, 
changes into brown, and later be- 
^ Hartig, Lehrbuch der Baumkrankkeiien, ist edition, pp. 56 et seq., Table II 


Fig. 91. — A plant of \'. Vitis Idaa 
which lias been infected by M. 
Goeppertiana. a, the infected 
stem containing the mycelium. 
The new shoots /', in the year 
succeeding that in which the plant 
was infected, undergo abnor- 
mal thickening under the in- 
fluence of the mycelium ; the 
apex alone retaining normal 
dimensions. c, the youngest 
shoot ; d, a dead portion. 



comes almost black. The lower leaves of each shoot are dwarfed, 
while the upper ones develop normally. If one infects a healthy 
cowberry plant with the aecidiospores of the columnar rust of the 
silver fir — which will be described presently — the stem remains 
unaltered during the first year, although the mycelium spreads 
in the tissues of the cortex. Next year, however, the young 
shoots are affected in the manner just described. The my- 
celium grows into the young shoots, where, by the exuda- 
tion of a ferment, it 
p^p^-_,_^ stimulates growth in 
all the cortical cells. 
This effect, however, 
can onlybe produced 
so long as the cells 
of the new shoots are 
still young. But on 
account of the slow 
upward growth of 
the mycelium in the 
shoot, it only reaches 
the apex at a time 
when the cells of 
the cortex are com- 
pletely matured, and 
w^hen, consequently, 
it is no longer able 
to stimulate to in- 
creased growth. 

Fig. 92. — Cortical parenchyma and epidermal cells 
from the stem of V. Vids Idcea. The mycelium 
grows in the intercellular spaces, and pushes short 
branches, which swell at the apex, against the outer 
wall of the cells. A delicate prolongation at the 
apex of these branches penetrates the cell-wall, after 
which a sac-like haustorium develops in the interior 
of the cell. Underneath the epidermal cells the 
hypha; enlarge in a clavate manner, a a. Haus- 
toria, /', and teleutospore-mother-cells, c c, develop 
in the epidermal cells. Magnified 420 times. 

The mycelium, 
however, pushes up 
as far as the topmost bud, which may be stimulated to shoot 
out even in the same year as that in which it is formed. The 
mycelium, which is perennial, is intercellular, and abstracts 
nutriment from the parenchymatous cells by means of haustoria 
(Fig. 92). It ultimately reaches the epidermis, underneath which 
it swells up in a clavate manner (Fig. 92, a a). 

Haustoria {b) are also pushed into the epidermal cells, which 
may at once be distinguished, by their shape, from the young 
spore-mother-cells {c c), which are also developed there. 



From four to eight, usually six, such mother-cells are produced 
in each epidermal cell. These increase in size till they occupy 
the whole space, and then divide each into four teleutospores, 
which are arranged side by side in a palisade fashion (Fig. 93,(7). 
During wet weather in May of the following year, each teleuto- 
spore germinates /// situ to 
produce a promycelium {li), 
on which the sporidia deve- 
lop on short sterigmata (r). 
Should these reach the silver 
fir they penetrate the young 
leaves by means of a germ- 
tube, and four weeks later 
the mycelium produces two 
rows of secidia on the un- 
der side of each leaf, which 
are characterised by an ex- 
tremely long peridium (Fig. 
94). The peridia burst open 
in various ways at the apex, 
and allow the ?ecidiospores to 
escape (Fig. 95). These are 
characterised by the unusual 
length of the intermediate 
cells, which separate the in- 
dividual spores from each 
other. The jecidiospores ger- 
minate when they reach the 
epidermis of Vacciniiivi Vitis 
IdcEa, and produce either a 
tube which remains of uni- 
form thickness, though it 
sometimes branches, or a 
germ-tube which enlarges in 

a sac-like fashion towards the extremity. Infection is accom- 
plished by means of a fine hypha which springs from the germ- 

The infected leaves remain green for a considerable period, 
and only fall off in the course of the summer. Even in 

Fig. 93. — Epidermis and cortex of Jl 
Vids Idira containing ripe teleuto- 
spores of M. Goeppcrtiana, some of 
which are germinating. The mother- 
cells, each of which forms four teleuto- 
spores, are usually found six together 
in an epidermal cell, a. A germinat- 
ing teleutospore produces a promy- 
celium, /', on which, after the formation 
of three transverse septa, four sporidia, 
(■, usually develop on short sterigmata. 
jNIagnified 420 times. 

M 2 



August I have found green leaves beset with the withered 

Serious damage is done only when the young silver firs are 
situated amongst badly infested cowberr}'-bushes, and when the 
greater part of the leaves contract the disease. The aecidium 
form possesses a facultative character — that is to say, it may be 
absent without endangering the existence of the parasite whose 

Fig. 94. — a, the branch of a silver 
fir on the under side of whose 
leaves two rows of the aecidia 
ofyi/. Goeppe7iiana {ALcidiuin 
cohtinnarc) have developed ; b, 
the nscidia magnified. 

Fig. 95. — An recidium of yl/. Goeppei-- 
tiana, a, in the tissues of a leaf of 
the silver fir ; /', the strings of 
cecidiospores with intermediate 
cells ; c, germinating cecidiospoi-es. 

sporidia are capable of germinating on, and of directly infecting 
cowberry- plants. 

Where damage is to be apprehended from the columnar rust 
of the silver fir during the regeneration of a wood, one may reduce 
the chances of an outbreak by rooting out diseased cowberry- 
plants. On account of their striking appearance, these arc not 
difficult to find. 


Under the name Melampsora popidiua, the Poplar-rust, are 
denoted the fungus-forms belonging to this genus which are 
met with on various species of poplars, and which await more 
thorough and exact investigation. 

Forms are met with on Popiihis tremnla whose cushion-like 
* [Occurs in England, and reqtiires investigation. — Ed.] 



stromata are distinguished by their smaller size from those which 
occur on P. balsaniifera {JSI. Balsamifera Thinn.), and it would 
also appear that the form M. populina Jacq. which is often 
met with in luxuriant development on P. nigra is distinct from 
the first two. Owing to the development and increase of the 
uredospores in the course of the summer, the foliage may 
appear quite golden yellow in August, and poplars sometimes 
suffer so severely from this 
rust that even in September 
the trees may be entirely 

The teleutospore- layers 
are primarily concealed by 
the epidermis of the leaf, but 
ultimately appear above the 
surface as smooth brownish- 
yellow cushions, which after- 
wards become dark brown 
(Fig. 96) ; while the yellow 
uredo-layers, after rupturing 
the epidermis, may be recog- 
nised as loose clusters of 

It would appear desirable 
that these various forms of 
poplar-rust should be made 

the subject of more exact investigation, seeing that their jecidium 
forms have not yet been determined with certainty. 

I have, so far, investigated only the Melampsora that affects 
Popnlns tremnla. Even as early as 1874^ I drew attention to 
the fact that, with scarcely an exception, aspens present 
in young Scotch pine woods are infested by Caoiiia piiiitor- 
qniivi, and that some connection might possibly exist between 
Ccvouia and some fungus that occurs on the aspen. 

I indicated that such a connection was doubtful in the 
case of Melmnpsora Treimdce, because this fungus occurs in 
districts where Cczoma pinitorqiiiim is unknown. In the in- 
terval, however, Rostrup proved experimentally the connection 
^ Wichtige Krankheiteii dcr Waldbiittmc, p. 91. 

Fig. 96. — Aspen-leaf showing the teleuto- 
spore-layers of M. Tremuhr. 


between these two fungi, and this I was able afterwards to 
confirm. At the same time I proved that M. TreimilcE produces 
Cceorna Lands on the larch. 

Then .Rostrup also obtained Caonia Merairialis by infection 
with M.,.rTr£;mii Ice. Rathay believes that he has also obtained 
y^cidiuin Ciematitis on Clematis v it alba by infection with spores 
oiM.populina. . 

As regards CcEoma piiiitorqitmn apd C. Laricis, I obtained the 
aecidia of both by infection with sporidia from the same aspen-leaf, 
and, further, I have infected Pimis with teleutospores of Melamp- 
sora which I had raised by sowing CcBoma Laricis on the aspen. 

Thus, although it appears to me that the identity of these 
two species of Ccconia has been conclusively proved, still it 
would be desirable to have this confirmed. Even more pressing, 
however, is the solution of the. problem whether CcEonia Mer- 
airialis also originates in the same species of Melainpsora, or 
whether various species occur on the aspen to which these 
aecidium forms belong. Further, it is necessary to discover 
whether the species present on P. nigra, P. alba, and P. bal- 
saniifera are identical with those on the aspen ; and, finally, it 
remains to be determined whether the cccidium forms possess a 
facultative character, as appears to me most probable. I have 
described below the two diseases produced on conifers by M. 
T renin I a;. 

First Form on Pinus sylvestris rt'//// Caeoma pinitorquum. The 
Pine Shoot Twist Disease. Melampsora Tremulse pinitorquum 

This disease is distributed throughout the whole of Ger- 
many, being most prevalent in the north, where it has proved 
exceedingly destructive, especially from 1870 to 1873. It may 
attack young pine seedlings even at the stage when they are 
just appearing above ground. In such a case longish pale 
yellow sporogenous layers rupture the epidermis and appear 
upon the surface of the stems or leaves. The disease is most 
frequently observed in pine woods from one to ten years old, 
infection being brought about by the teleutospores of M. 
Tremnla; which develop on aspen- leaves that are lying about on 
the ground. The disease ma}- be recognized by the fact that in 
the beginning of June, sometimes even in the end of May, at the 



season when the apex of the green leaf-fascicles on the young 
shoots is projecting somewhat from the leaf-sheath, pale yellow 
patches f to li inch long and )- to f inch broad appear upon the 
green cortical tissues of the shoots (Fig. 97), and on these, with 
the help of a pocket lens, numerous small rather deeper yellow 
tubercles, the spermogonia, may be made out. These are 
formed partly in the epidermal cells, and partly between them 
and the cuticle, the latter being raised up to cover the spermo- 
gonium (Fig. 98). The CtroinaAd-y^r originates in the second 
or third row of cortical cells, and is formed by the inter- 
cellular mycelium growing outwards from the interior of the 
stem to produce a sporophore in that 
region. The secidiospores are afterwards 
distributed by abscission from the apex 
of basidia in the usual way. While the 
formation of this internal sporogenous 

Fig. 97. — Apex cf a young 
pine-shoot showing the 
sporogenous layers of 
C.piniiorquinn through 
the ruptured cortex. 
Natural size. 

Fig. 98. — Transverse section of a sporogenous layer 
of C. pinitorquuiii before the cortex has ruptured. 
Two tubercular spermogonia are visible in the 
epidermis. i 

layer is proceeding, the surface of the part of the cortex affected 
is constantly assuming a deeper golden }'ellow colour, while, 
at the same time, it forms a cushion-like elevation which 
results in the development of a longitudinal fissure (Fig. 97) in 
the external layers of the cortex through which the spores are 
shed. The tissues of the cortex beneath the sporophore after- 
wards die as far in as the wood, and a callus is formed, under 
favourable circumstances, in about a year. 

During the development of the spores, and for some time 
afterwards, the young shoot continues to elongate normally, 
except at the seat of the disease. The result is that the diseased 

1 68 


shoot bends over a little at the place occupied by the spore-layers. 
In many cases, however, the ultimate contortions, which have 
earned for the parasite the designation Pine Twister, C. 
pinitorqimm, are due to the weight of the young shoot, whose 
apex, in the case of lateral branches that are considerably 
damaged on one side, is bound to be depressed. Later on the 
apex again grows up, and thus S-shaped contortions arise. If 
the weather is normal, a few such sporophores are formed 

Fig. 99. — Apex of a pine which has been attacked by C. pinitorquitm. The leading 
shoot has been killed almost to the base. The branches of the whorl, as well as 
the main shoot-axis, show diseased spots and contortions which have existed for 
a considerable time. 

annually on the young shoots, while in very dry weather the 
sporogenous layers wither up as soon as formed, and no external 
damage is visible. Should May and the beginning of June be 
very wet, the sporophores develop so abundantly and luxuriantly 
that the shoots, with the exception of the base, die off and dry 
up completely (Fig. 99). A badly diseased young Scotch pine 
wood appears in the end of June as if a late frost had killed and 
contorted all the young shoots. Next year the dormant eyes 



of the leaf-fascicles that survive at the base of the shoot develop 
into shoots, and these afterwards become diseased like the rest. 
The fact that a pine that is once attacked by the fungus suffers 
from the disease year after year during successive decades 
justifies the assumption that the mycelium of the fungus is 
perennial in the shoots. From the part of a pine wood that is 
first attacked — the focus of the disease — the disease continues to 
spread each year in a centrifugal manner. The point is to be 
emphasized that very young woods — those from one to three 
years old — suffer most from the disease. Pines that become 
diseased at a later period are sometimes so badly crippled as to 
hold out but faint hope of a healthy wood, but as a rule a dry 
spring occurs sooner or later which retards the development of 
the fungus, and so, a few years' mitigation of the disease being 
granted, the plants gradually recover, although they again suffer 
in unfavourable seasons. About the thirteenth year the disease 
spontaneously disappears. Clearing out the aspens from the 
young pine woods is the surest method of combating the disease 

Second Form on Larix europaea wit/i Caeoma Laricis.i Melampsora 

Tremulae Laricis 

The larch-Ieaf-rust is distributed throughout the whole of 
Germany, and is frequently so common that a large part of the 
foliage is destroyed by the fungus. 
It is often overlooked on account 
of the damage bearing a certain re- 
semblance to that due to Chernies 
Laricis. In the month of May 
numerous spermogonia first of all 
appear on the leaves, amongst 
which the 6"<^^;//rt- layers break 
through the epidermis of the leaf 
in the form of long or short yellow 

After the spores have been shed 
the leaves wither and fall off. Felling the aspens in the neigh-*" 
bourhood of larch woods protects the latter against the disease. 

1 Wichtige Krankheite?i der Waldbdume, 1 874, p. 93, and Allgemeine Forst- 
joid Jagd-Zcitung 1885, p. 326. 

Yic. 100. — Larch-leaves attacked by 
CtCiV/tii Laricis. 




Several species of Melanipsora occur on the various willows, 
and these, until a short time ago, were grouped under the 
common collective name of M. salicina. By means of the form 
of the teleutospores and urcdospores, Thiimen was the first to 
distinguish a number of species, which ought to be thoroughly 
investigated for the sake of verification. 
Rostrup - has, in the interval, succeeded 
in obtaining the scidia of two species, 
which will now be more particularly 

The uredospores appear sometimes as 
early as the end of May or the begin- 
ning of June, in small reddish yellow 
clusters on the lower surface, more 
rarely on the upper surface, of the 
leaves of Salix pruinosa, S. daphiioidcs, 
S. viniinalis, and other willows. The 
disease spreads rapidly, partly owing to 
the internal growth of the mycelium, 
which penetrates the cortex of the 
shoots by way of the leaf-petioles, and 
partly by means of the uredospores 
which are carried by the wind. These 
germinate very quickly, and produce 
numerous new uredo-clusters, generalh' 
on the eighth day after they have been 
sown on a sound leaf. Leaves that are 
attacked soon become marked with black blotches, and drop off. 
Before the leaves fall off or die numerous teleutospore-layers, 
about the size of a pin-head, develop beneath the epidermis of 
the leaf (Fig. loi). These occur more especially in late summer 

^ Von Thiimen, Mittheilungen aus dem forstlichen Versiichswesen Oester- 
reichs, ii. p. 41 et seq. Hartig, Wichtige Krankheiteii der Waldbiiume^ 
pp. \\() et seq. 

2 Rostrup, Fortsatte Undersogeher over Snyltesvampes Angreb par 
Skoi'traeerne Kjobcnliaven., 18S3. 

Fig. ioi. — M. Hartigii on 
Salix p7-uhiosa. a, a living 
leaf with sporophores ; /', 
a leaf which has been 
killed at places ; c, sporo- 
phores on the stem close 
to the base of the leaf 


and in autumn. These small cushion-like bodies, which are at 
first pale brown and later very dark brown in colour, pass the 
winter in the tissues of the decaying leaves that are l}'ing on the 
ground, and produce promycelia and sporidia in spring. These 
sporidia are carried by the wind to the leaves of the young willow- 
shoots, and induce the disease afresh. They produce Ccuoma 
Ribesii on the leaves of Ribcs alpimiui, R. Grossularia, R. rnbnun, 
and R. nigriini. This aecidium form appears, however, to pos- 
sess merely a facultative character, for we annually meet with 
numerous instances of the disease, especially in autumn, even in 
places where no examples of Ribes are to be met with for long 

Hitherto I have met with serious infestations of the fungus 
only on Salix prtiinosa (syn. caspica, acntifolid), numerous osier- 
beds being entirely destroyed by repeated premature defoliation. 
The best preventive measures consist in raking together and 
burying or burning the fallen fungus-infested leaves from 
autumn till spring, and in careful attention to the osier-beds 
during summer. As soon as the rust appears sporadically it is 
advisable to cut off and bury, the infested shoots. In place of 
the glabrous-leaved Salix pniinosa, which suffers most from the 
fungus, the cultivation of the hybrid J?, pruinosa x daphnoides 
is to be recommended, the latter being pubescent, and thus better 
protected against infection. 


This willow-rust is very common on Salix Caprea, S. cinerea, 
S. aiirita, S. longifolia, S. repens, and vS. reticulata. It produces 
the cEcidia of Cceoina Evonyini on Evonynms. 

Then we also meet with Melajupsora epitca on vS. alba, 
S. incana, S. piopiirea, S. nigiicans, and 5". retiisa ; and J/. 
mixta on S. triaiidra, S. hastata, and 6". silesiaca. 

M. betulina occurs on various species of Betula. 

M. Caipini „ „ Carpinus Bctiilus. 

M. Sorbi ,, „ Sorbus Aiiaiparia ?a-\A S.torminalis. 

M. Arice „ ,, Sorlnis Aria. 

M. Padi ,, „ Primus Padus. 

M. Vaccina „ ,, Species of Vacciuiuni.* 

♦[Several of these are recorded for this country, and are much in need of 
thorough investigation. — Ed.] 



The genus Coleosporiiim is distinguished from the preceding 
one by the teleutospores being formed out of several super- 
imposed cells, each of which produces a unicellular promycelium 
with a single sporidium. 

According to the investigations of Wolf and Klebahn, three 
species of this genus produce aecidia on the leaves of the pine. 
These include not only C. Senecionis, which occurs on various 
species of Senecio (according to Wolf), but also C. Euphrasies 
and C. Tnssilaginis (according to Klebahn). The aecidium forms 
are known under the names of Peridermium Pini acicola or 
Peridennunn oblongisporinm, the pine-Icaf-rust.^ 

In the months of April and May the cxcidia may be observed 
on the one- and two-year-old leaves chiefly of younger pines, 
occasionally also of old trees. The sper- 
mogonia are found scattered amongst the 
reddish yellow vesicles (Fig. 102), which 
are only a few millimetres in size. The 
former become brown with age, and thus 
look from the outside like small black 
blotches. The mycelium develops in the 
interior of the leaf, where it passes the 
winter, and in the following year it may 
again produce aecidia without killing the 
leaf Seeing that the leaves which are 
infested by the aecidia do not die pre- 
maturely, or at least not to any great ex- 
tent, the injury done by this form of the 
fungus is insignificant. Discoloured spots are merely formed 
upon the leaves. 

Several species of Cronartiiiui produce aecidia and spermo- 
gonia in the cortical tissues of pines. These have hitherto been 
grouped under the name Perideriniuin Pini corticola* The 
disease, which occurs both in young and old woods, is very 
prejudicial to the health of the trees. How infection takes place 
— whether it must always be preceded by an abrasion of the 
cortical tissue, such as is induced by insects, woodpeckers, hail- 
' R. Hartig, Wichtige Krankheifen der IValdbaiinie, 1874. 
* [This disease occurs every year on pines in this country. — Ed.] 

Fig. 102. — The secidia 
and spermogonia of 
Perideriiiiiiin Pini 
acicola on the leaves 
of a pine. 



stones, &c. — remains for the present undetermined. Parts of 
the stem older than 20 — 25 years appear to be incapable of 
receiving infection. The mycelium of the fungus spreads by 
intercellular growth amongst the cells of the cortex and of the 
bast, from which it proceeds, by way of the medullary rays, into 
the wood to the depth of about four inches. 

Wherever the mycelium obtains access, the starch-grains and 
other cell-contents disappear, their place being taken by drops 
of oil of turpentine, which form on the inside of the walls, or 
saturate the wall-substance itself. The cells are, of course, 
killed, death however being unaccompanied by browning of the 
tissues. The whole stem, to a depth of some three or four inches, 
is completel}- saturated with resin, a section of wood, as much as 
one to two inches in thickness, being more or less translucent. 
As the mycelium penetrates the resin-ducts as well, killing the 
surrounding tissues, there is no doubt that a portion of the 
turpentine finds its way down from parts of the stem situated 
at a higher elevation. The 
assumption that direct con- 
version to turpentine of the 
cell-contents and of the wall- 
substance of the parench}'- 
matous cells also takes place 
is, however, justified by the 
complete resinous saturation, 
and by a frequent volumin- 
ous outpouring of turpentine 
from the cortex, which de- 
taches itself from the tree 
after death. 

Each year the mycelium 
spreads from the diseased 
part into adjoining tissues, 
the rate of progress being usually somewhat more rapid longi- 
tudinally than horizontally. In proportion as the mycelium 
spreads, so is the passage of the plastic materials confined to 
the sound side of the tree, in consequence of which the cam- 
bium in that region is stimulated to such a degree of activity as 
to produce exceptionally broad annual rings. Fig. 103 exhibits 
the cross section of a stem which, when fifteen )-ears old, was 

Fig. 10 

Transverse section from the 
upper part of the stem of a pine which, 
seventy years previously, had been in- 
fected at a by P. Pini corticola. The 
crown of the tree died in the year im- 
mediately preceding that in which the 
section was removed, and at that time 
the only portion of alburnum that was 
not saturated with resin or attacked by 
the fungus was the portion marked b. 
The portion of the wood saturated with 
resin is shaded in the figure. One tenth 
natural size. 



infected at a, but which, with the part of the crown situated 
above the point of infection, only succumbed in its eighty-fifth 
year. The top of a diseased tree is specially liable to die during 
a dry warm summer, when the wood, having for the most part 
undergone resinous degeneration, is unable to allow sufficient 
water to pass to compensate 
for the rapid evaporation from 
the crown. 

For the most part ascidia 
are formed only in that re- 
gion of the cortex which has 
become diseased during the 
preceding year. They break 

Fig. 104. — Portion of the stem of a 
young pine showing the vesicular 
CEcidia of P. Pint corticola break- 
ing through the cortex. At three 
places which are darker shaded 
spermogonia are situated beneath 
the periderm. Natural size. 

Fig. 105. — A pine-branch which has been 
infested by P. Pini corticola for several 
years. The branches on the left side, 
which were the first to be attacked, are 
already dead. From these the mycelium 
has spread backwards on to the main 
branch and the other lateral branches. 
One fifth natural size. 

through the outer dead cortical layers in the months of May and 
June as hemispherical, oblong, sausage-shaped, yellowish white 
vesicles filled with reddish yellow powdery spores (Fig. 104). 
Amongst these one recognizes with difficulty the flat spermo- 
gonia which are about the size of a pea. These consist of 
innumerable fine sterigmata which are situated between the 
deepest periderm-layer and the living cortex, being arranged at 
right angles to the former. The small spermatia are abjointed 
from the apex of these sterigmata. 

Branches and twigs of the crowns of the older classes of trees 


often die in a few years, after which the parasite frequently 
spreads downwards from the base of the branch to the main 
axis of the stem (Fig. 105). Even should the latter die the tree 
will still remain alive, provided there are branches and twigs well 
provided with leaves beneath the canker-spot. These branches 
constitute a kind of new leader, and the dead crown forms 
the " resin-top " or " resin-leader " (" bird-resin "), which was 
regarded by Ratzeburg as the result of injury caused by the 
Pine Beauty Moth, and designated as " spear-top." 

The disease has also been described by this observer as 
'' moth-wither," or, in other words, as the result of the attack of 
Phycis abictdla ( Tinea sylvestrella, Ratz). 

Three species of pine-blister-rust are to be distinguished in 
the cortex of trees, viz. : — 

(i) PeridermiiLin Strobi, which occurs only in the cortex of 
PiiiiLS Strobus. The aecidium form is Cronaytium ritbicila. 

(2) Peridcniiiuin Conuii, which occurs in the cortex of Piniis 
sylvestris, has for its aecidium form Cyonartinni asdepiadeuin on 
A sdepiiis Vincctoxicum. 

(3) Peridermium Pini. This is probably the most destructive 
species, and it is to be regretted that so far the plants have 
not been determined on which the teleutospores are produced. 
Until we discover the teleuto form preventive measures must be 
confined to felling pines that arc attacked. 

The genus Chrysomyxa is closely related to the preceding one, 
in that here also each teleutospore consists of a row of cells, the 
terminal one of which produces a multicellular promycelium 
with four sterigmata and sporidia. The sporophore consists of 
a dense orange yellow cushion-like body of varying shape. The 
urcdo and aecidium layers are similar to those of the genus 

This is a disease of the spruce which is met with through- 
out the whole of Germany, with the exception of the higher 
Alpine regions. It occurs on old as well as on young spruces, 

^ Reess, Bat. Zeit., \2,6s, Nos. 51 and 52, and \\-\\\komm^ D/c mzk?-os- 
kopischen Fcinde des Waldos, 1868, pp. 134 — 166. 

^' O-^^Ji^.^::^ u,^c0jJ ^ tVt.74.^>^ 



and is frequently so prevalent on the youngest shoots as to kill 
a large proportion of the leaves, which subsequently drop off. 

The fungus is autcEcious, the uredo and jecidium layers being 
entirely absent. Its teleutospore-layers alone are developed on 
the spruce-leaves. In the month of May the sporidia germinate 
on the delicate leaves of the young shoots, inside which they 
produce the mycelium, which contains numerous drops of yellow 
oil. By the end of June the part of the leaf that is infested may 
be recognized by its pale yellow colour. The diseased portion 
may occupy the base, middle, or apex of the leaf As autumn 
approaches it constantly becomes deeper lemon yellow in 
colour, while the rest of the leaf remains green. In autumn 
formation of the teleutospore-layers begins on both the 
under sides of the leaf These take the form 
of oblong, somewhat prominent cushions, 
which are at once to be recognized by their 
more golden yellow colour. In this condi- 
tion the fungus hibernates on the tree, and 
in the following spring the teleutospore-layer 
still further develops (Fig. 106), until finally 
the epidermis ruptures longitudinally and ex- 
poses a golden yellow stroma. The pnDmy- 
celia with their sporidia next develop from 
the cells of the teleutospores, as is shown 
in the case of C. Rhododendri in Fig. 107, 
and as this occurs in the month of May, at a 
time when the young shoots of the spruce arc 
forming, the sporidia have the opportunity 
of germinating directly on the young leaves. 
It is probable that spruces wdiich arc very backward in growth 
at the time when the spores ripen escape infection, and this 
explains why many individuals of a wood remain entirely free 
from the fungus while others are very badly attacked. Such 
cases have frequently inspired non-professional minds with the 
belief that the fungoid disease is dependent upon a pre- 
disposition to disease on the part of particular spruces. After 
the sporidia drop off the teleutospore-layers wither, while the 
leaves themselves soon afterwards die and fall from Uhe tree. 
As a rule the tree suffers but little from the loss of leaves. 

Fig. 106. — A spruce- 
leaf attacked by 
C. Abietis, the 
golden yellow 
spore-layers of 
which have not 
yet ruptured the 


because a sufficient supply of foliage is always left on the older 
parts of the branches as well as on the youngest shoots. As the 
meteorological conditions are not always equally favourable for 
the germination of the sporidia, and as the teleutospores may 
germinate at a time when most of the spruces are either too far, 
or not far enough, advanced in development to be infected by 
the sporidia, it very seldom happens that the disease maintains 
its intensity throughout a long series of years. With the 
exception of a spruce wood in the Saxon Erzgebirge, I have 
never come across very serious damage due to Chrysomyxa 
Abietis. Certain years occur, and not uncommonly, when the 
disease is very scarce, and when the spruces are in a position to 
clothe themselves with a full stock of leaves. On this account 
I am not able to agree w^th Willkomm, Frank, &c., in their 
recommendations regarding measures for combating the fungus, 
because felling the diseased trees, and such like measures, would 
have worse consequences than the disease itself. 

It may be not uninteresting to note that in the severe winter 
1879-80 it was noticed that in many districts the diseased leaves 
dried up so that the development of the fungus was prevented. 
And, further, it not unfrcquently happens that Hysterhun macro- 
sporum is encountered along with Chrysomyxa, the result being 
that the development of the leaves is interfered with, and they 
acquire black blotches. ^2»^CU^. 1 1 


The Rust of the Rhododendron is of special interest in that it / j. f 
is heteroecious, developing its teleutospore- and uredo-layers in » 
clusters in the form of small roundish and oblong cushions ' *M4aa*»< 
on the leaves of rhododendrons, while the aecidia [ALcidmni 4m»*-*-'*' 
abietimim, the Spruce-Blister-Rust) develop on the leaves of the Ou/u/t^ 
young shoots of th^ spruce. -VfcxJ* /^•* 

The occurrence of the disease on spruces is consequently ^ 
dependent on the presence of rhododendrons,- although, ofC^^^V-**' 
course, the spread of the sporidia, by means of wind and rain, >sc«sr /* 
from high elevations into the valleys or otherwise is not im- ^I 1 
possible. De Bary, to whom we are indebted for our know- rf^ 

1 De Bary, i5^/. Z«V., 1879. ^ ^^ 

- In the Alps R. hirsutuin and R. ferruginium are the common species. ^ -^ Liy 




ledge of the biology of this parasite, has, however, proved that 
the aecidium form may be dispensed with. Where spruces arc 
absent the sporidia germinate directly on the leaves of the 
rhododendron to produce the uredo layers, and these serve 
to maintain and spread the fungus during summer, until, in 
autumn, teleutospore-layers are again formed on the leaves of 
the youngest rhododendron shoots. These hibernate, and in the 
following spring the germination of the teleutospores results in 
the rupturing of the leaf-epidermis (Fig. 107). 

At first the development of the parasite in the spruce-leaf 
resembles that of Cluysoniyxa Abietis, but even in July and 

Fig. 107. — Teleutospore-Iayer of C. Rhodo- 
dendri on R. lii^sntum. The development 
of the promycelia has caused the epidermis 
of the leaf to rupture. 

Fig. 108. — Spernio- 
gonia and secidia 
of C. Rhododetidri 
on a spruce-leaf \ 


August numerous small spots, the spermogonia,/!, are to be 
observed on the yellow parts of the leaves. Shortly afterwards 
one sees the yellow vesicles of the aecidia breaking through the 
epidermis, and these bear a close resemblance to those produced 
by the pine-blister-rust on the leaves of pines (Fig. 108), In 
August and September, when the peridia burst at the apex, the 
secidiospores are liberated in such numbers that if a diseased 
spruce is shaken the air is filled with a dense cloud of spores. 
In the course of the same year the diseased leaves die and drop 
off. This distinguishes the parasite at once from Chrysomyxa 
Abietis, which, in an immature state, hibernates on the tree. On 
lateral branches it is usually only the leaves that are situated on the 
upper side that are diseased. The leaves on the under side, being 
preserved against infection by those above, escape the disease. 



This parasite induces the same pathological symptoms in the 
spruce as the preceding one, but it differs in producing its teleu- 
tosporcs and uredospores on the leaves of Ledum palustre* 
Letters received from Russia mention that the fungus occurs in 
extraordinary abundance in that country, and I have also recently 
had it forwarded from the district of Konigsberg. It has also 
frequently been observed in other parts of Germany, with the 
exception of the south, but of course only where Ledum occurs 
in the immediate neighbourhood. 

In the case of the parasites about to be described, the aecidium 
forms alone are as yet known, so that the investigation of the 
course of development of what are probably all heteroecious 
forms of fungi is work for the future. 


From amongst the aecidium forms of whose teleutospore-forms 
we are so far ignorant, attention will here be directed only to 
those species which occur on forest trees. 


This fungus is parasitic on the silver fir, where it produces the 
so-called witches' brooms and canker-knobs which may be seen 
an)-whcre in Germany where woods of silver firs occur.^ As I 
have always noticed small wounds on one- or two-year-old 
witches' brooms close to their base — near the point, namely, where 
they have developed from a bud — it may in the meantime be 
assumed that infection occurs at such wounds. The mycelium 
of the fungus, which stimulates growth in a very marked manner, 
is perennial in the cortical and bast tissues of the stem, and even 
penetrates the cambium and the wood. Should infection occur 

1 De Bary, Bot. Zeif., 1879. 2 /^/^^ 1867. 

[3 Very common also in many parts of Britain, Ci^. the south-west of 
Scotland. — Tratts.'] 

*[An Ericaceous plant found on moors, &c., in N.E. Europe. — Ed. ] 

t [The cankers occur in various Silver Firs grown in English gardens, 
though they often seem devoid of the Witches' Brooms and secidia. I have 
seen both, however, on Abies Pinsapo and other species. — Ed.] 

N 2 

I So 


at a part of a stem or branch where there are no buds capable of 
developing, the stimulated growth of the cambium induces the 
formation of a knob-like swelling at that place, which is due 
both to the increased formation of wood and to the more vigorous 
development of the cortex (Fig. 109). With the spread of the 
mycelium the swellings or canker-spots increase in size, and if 
present on the stem of 
a vigorous tree they may 
attain to large dimen- 
sions. At such places 
the tissues of the cortex 
and bast soon become 
fissured (Fig. no), and 
dry up here and there 
as far in as the wood, so 
that in course of time a 

Fig. 109. — Swelling on the silver 
fir, but without the formation 
of a witches' broom. Natural 

P'lG. no. — Longitudinal section of a silver fir 
showing a swelling,one third natural size, which 
had originated by infection thirty-one years 
previously, when the stem was four years old. 
On the right side the cortex, which has been 
dead for three years, has withered and dropped 
off. At the infected part the growth of the 
cortex and wood has been stimulated. 

door is opened for the entrance of wood-parasites. One of the 
commonest of these is Polyporus Hartigii, which produces a kind 
of white rot. A species oi Agar i ens also — namely, A. adiposus— 
frequently appears as a wood-destroying parasite. In consequence 
of the decomposition of the wood, storms and snow often break 
the trees. One not unfrequently finds swellings which have 



originated without any connection with witches' brooms (Fig. 
109), and on such swelHngs no formation of spores ever occurs. 

More frequently infection occurs on or in the immediate 
neighbourhood of a bud. When the bud proceeds to grow it 
forms a young witches' broom — ^that is to say, a shoot whose 
cortex is stimulated to growth by the advancing mycelium, and 
whose young leaves are so affected by the parasite that they 
remain small, are somewhat round in cross-section, and show 

Fig. III. — Branch of a silver fir, with a witches' broom two years old, a:. The 
mycelium in advancing through the tissues of the branch has so stimulated a 
dormant eye that it has developed into a shoot a year later, b. The portion of 
the branch that has been invaded by the mycelium is much swollen. 

scarcely any chlorophyll. They remain yellowish, and in the 
beginning of August two rows of aecidia appear on their under 
side, which open and shed their spores in the end of the 
month (Fig. in). Soon afterwards the leaves die and fall off. 
The witches' broom is consequently deciduous. Each year 
the mycelium advances into the new shoots, to produce the 
phenomena already described. The twigs of branches which 
show this peculiar symbiosis * anastomose abundantly, and for 
the most part incline upwards, so that they appear amongst 
the ordinary branches of the silver fir like perfectly independent 

* [Symbiosis signifies the fact that two organisms are living together— a 
dual existence.] 


organisms, somewhat after the manner of the mistletoe. The 
myceHum also spreads slowly backwards in the cortical and 
bast tissues, so that a swelling or canker-spot, such as I have 

already described, is formed on 
the stem or branch to which the 
witches' broom is attached (Fig. 
112). This swelling increases 
independently, even after the 
witches' broom, as such, is dead, 
an event which is sometimes de- 
layed for twenty years or more. 
In young woods all trees that 
show cancerous swellings on their 
boles should be removed in the 
thinnings, even in cases where 
they belong to the larger class 
of trees. 

Fig. 113. — i^cidia Fig. 114. — Theouter 

Fig. 1X2. — The appearance of a witches' 
broom, seven years old, in winter, 
when, being deciduous, of course it is 
leafless. Above the point from which 
the witches' broom springs the fir 
branch has almost ceased to exist. 

of ^. strobiliiiiini 
on the upper side 
of a scale of a 

side of the scale 
of a spruce-cone, 
showing two pale 
patches which had 
previously been oc- 
cupied by the seci- 
dia of A. conoriim 

This aicidium form develops its mycelium in the green living 
carpellary scales of the spruce. It destroys the organs of the 
flower, and produces dark hemispherical brown aecidia, which are 
densely crowded, for the most part on the inner side of the 
scales, though to some extent also on the outer side. These 
aecidia usually rupture transversely (Fig. 113). When such 
cones fall to the ground they arc easily recognized by the fact 

^ Reess, Die Rostpilsforuien der dei/tsc/ien Coniferen. 



that they open even in damp weather, whereas sound cones 
remain tightly closed. This cone-disease occurs throughout the 
whole of North Germany, reaching as far south as the spurs 
of the Alps. 

This cone-infesting fungus differs from the former by the fact 
that only two large aecidia are situated on the outer side of 
each scale of the spruce-cone. After the pale peridia have 
ruptured and the spores have been scattered, pale spots are left 
on the scales (Fig. 1 14). 

When this rust-fungus, which is common on spruces in 
Sweden and Finland, attacks a young shoot, the whole of the 

Fig. 115. — A spruce-branch, one shoot of which, a, has developed normally, while 
two shoots, d, have been attacked by A. coniscans. All the leaves of the 
diseased shoots are short and fleshy, and bear secidia on all their four sides. The 
lower sides, c, and the upper sides, d, of a diseased leaf showing the cecidia, 
the peridia of which are still present at c, but have mostly disappeared at (/. 

leaves are affected. They become occupied by a peridium, which 
ruptures in places or along the whole length of the leaf, and 
displays the golden yellow aecidia underneath. Such leaves 
make the shoot look like a fleshy cone (Fig. 115). These " cones " 
are eaten in Sweden under the name " Mjolkomlor." 

^ Reess, Die Rostpilzformen dcr dcutschcn Coiiifereti^ p. 100. 

1 84 



This disease closely resembles the vesicular or columnar 
rust, y^cidinm cohininare {Melanipsora Goep- 
pertiajid), from which it is distinguished by 
the presence of numerous spermogonia, and 
by the absence of a peridium. It occurs 
on the lower side of the leaves of the 
silver fir, usually in the form of linear 
sporogenous layers, which are situated on 
both sides of the mid-rib (P'ig. Ii6). It is 
very abundant in the Bavarian Alps and 
in the woods near Passau, and is probably 
to be met with wherever the silver fir is 
The damage which it causes takes the form of the diseased 

leaves falling off in the first year, but the injury is comparatively 


Fig. ii6. — A fir-leaf 
showing C. Abieiis 


Most of i\\c: HfJiieiioniycetcs are saprophytes, and develop their 
mycelium in soil that is rich in humus, or in the interior of the 
dead parts of plants, and especially in dead wood ; while the 
sporophore, which is often of large proportions, appears on 
the surface of the ground or on the outside of the plant, Onl)- 
relatively few of the Hymenomycetes are undoubtedly parasitic 
in character, and, in the case of many, more exact investigation 
must determine whether they are to be classed as parasites 
or saprophytes. The peculiarity in the production of the spores 
consists in their being simultaneously formed in fours at the 
apex of basidia, and that these basidia constitute a more or less 
dense layer (hymenium), which may occupy a part or the whole 
surface of the hymenophore. 

^ Reess, Die Rostpilzforinen der dcKtschcn Coni/eren, p. 115. 

* [These include the " Mushrooms" and " Toadstools "' in the wider sense: 
we are still in want of a good English general term for them, and the trans- 
lation of the German Htit-pilze does not really meet this need. — Ed.] 




The genus Exobasidiiim induces the formation of character- 
istic galls on the leaves, flowers, and stems of various ligneous 
plants. The basidia originate on the mycelium, which is chiefly 
intercellular, and force their way outwards between the cells 
of the epidermis, on the surface of which they form a hymenial 
layer. No special sporophore, in the narrower sense of the term, 
is produced. 


This parasite produces swellings on the leaves, flowers, and 
stems of Vaccinhim Vitis Idcva, V. Myrtillus, and V. riliginosuin. 
These are partly of a beautiful white colour and partly of a 
bright rosy hue, and arc to be distinguished from the swellings due 
to Melampsora Goeppertiana hy their being dusted over with the 

Fig. 117. — A shoot of V. J'ltis Idtca, 
with the hymenium of E. Vaccinii 
on the leaves, a a, and in the stem. 

Fig. 118. — "Alpine-rose apple" on 
Rhododendron h irsntti?n. 

white spores, whereas, in the case of the latter, the lustrous 
epidermis hides the sporogenous layer ; and further by their 
occurring more frequently on the under surface of the leaves or 
on the racemose inflorescence than on the stem (Fig. 117). A 
microscopic examination at once reveals the fact that the long 
somewhat bent spores are situated on four delicate stcrigmata at 
the apex of the clavate basidia. 

This fungus, which was formerly described as a separate 

^ Woronin, I'erhandliingcn dcr 7iatiirf. GesellscJiaft zic Freibtiri:;, 1S67, I\'. 


species under the name Exobasidiiivi RJiododendri, produces the 
familiar " Alpine-rose apples " (Fig. Ii8) on the leaves of the 
Alpine rhododendrons. These bear a strong resemblance to 
many of the galls on oak-leaves which are caused by various 
species of Cynips, and are to be met with in all Alpine districts 
where rhododendrons occur. 


T. radiciperda is undoubtedly the most dangerous of all the 
parasites met with in coniferous woods, not only because it 
produces the worst kind of red-rot, but also on account of its 
being the most common cause of gaps in both young and old 
plantations. I have had the opportunity of observing it on 
various species of pines, especially P. sylvestris and P. Strobjis, 
and also on other conifers, notably Picea excelsa, Abies pectinata, 
^.wd, Jiiniperiis comniunis. It is true that I have also occasionally 
met with its sporophores on the roots of old stools of Betiiia, 
and on beeches that have been damaged by mice, still I doubt if 
it occurs on dicotyledons as a parasite. 

Not unfrequently the disease appears in plantations which are 
not more than five to ten years old, though it also occurs in 
woods of a hundred years' standing. Here and there indi- 
vidual trees showing luxuriant growth suddenly become pale, 
and die. We shall afterwards see that identical pathological 
symptoms are displayed by trees infected by Agariais inclleus. 
In the neighbourhood of a tree that has been killed — no matter 
whether it is left standing or has been felled — other trees soon 
die, and so in the course of years the death-circle constantly 
extends outwards. Large gaps and openings are thus formed in 
woods which were previously quite close. At first one generally 

' R. Hartig, Zersetzungserschei)iu7igen des Holzes,'^'^. 14 et set/., Tables 
I. — IV. Under the name Polyporus annosus, Fr., a number of different 
species of fungi have been described, Tramefcs radiciperda amongst the 
others. This mode of description has, however, been accepted as sufficiently 
accurate even in the second edition of Fries's Systetna, which appeared some 
years after I had described T. radiciperda. The name T. radiciperda is thus 
entitled to priority, and is also to be preferred, as it prevents any confusion. 

* [Brefeld, Unters. aus dcm Gesammigeb. der Mykol. VIII., re-names this 
Heterobasidion annosum, and describes its second kind of spores — conidia.- - 



observes only one or a very limited number of diseased spots in 
a wood, but when these have been allowed to extend for some 
\'cars one notices new seats of disease establishing themselves all 
over the wood. 

When the dead trees are examined about the roots, one finds 
the sporophores, with their snow-white hymenial surface, appear- 

FiG. 119.- — The sporophores of T. 
radiciperda on the roots of a 
spruce. Natural size. 

FiCt. 120. — The mycelium of T. 
radiciperda on the root of a 
spruce. The outer bark-scales 
have been removed from the 
lower portion so as to show 
the felted mycelium, a a, while 
in the upper portion only 
cushion-like mycelial masses, 
h, project from between the 
scales. Twice natural size. 

ing between the bark-scales as very small yellowish-white cushion- 
like structures (Fig. 119). These coalesce with similar adjoining 
cushions, and in exceptional cases attain to a diameter of twelve 
to sixteen inches. In the case of pines the sporophores are 
usually to be found on the stool close under the surface of the 
ground, though sometimes also on the deeper roots, while in the 
case of the spruce they are almost always to be found only on 
the roots. Between the bark- scales one finds the ramifying 
felted mycelium, which is distinguished from that of Agariais 
inellens by its extremely delicate te.xture (Fig. 120). It scarcely 


attains to the thickness of the finest tissue paper, and only where 
it penetrates between the bark scales does it swell up to form 
yellowish-white bodies varying in size from a pin-head to a pea. 
Decomposition (red-rot) spreads from the roots up into the 
interior of the stem to a considerable distance. It is only in 
the case of the Scotch pine that the rot does not ascend into 
the stem from the stool. 

Shortly described, the biology of the parasite is as follows. 
The spores, which are formed in the hymenial layer of the 
subterranean sporophore, do not as a rule spread from the place 
where they originate, unless they are brushed off by a passing 
object. As sporophores are especially liable to be formed on 
diseased roots at the point where they abut on mouse-holes, 
it appears to be a likely supposition that the mice, or other 
burrowing animals, carry away the spores on their fur, possibly 
for long distances, and afterwards rub them off on healthy roots. 
The spores soon germinate in warm humid air, and the m}-- 
celium, entering between the bark-scales, will probably reach 
the living cortex at some point or other. From this time its 
course of development is in two directions. It forces its way 
into the wood, in which it very rapidly travels up the stem. The 
contents of the parenchymatous cells are killed and turned 
brown by the action of the ferment that is exuded by the 
mycelium, while a violet colour in the wood is the visible 
symptom of this stage of the decomposition. With the 
disappearance of the protoplasmic contents of the cells the violet 
colour is replaced by pale brownish yellow, except for a few 
dark blotches which persist. The latter afterwards become sur- 
rounded by a white zone, and simultaneously the wood gradually 
becomes lighter and more spongy. Ultimately numerous holes 
are formed, the tissues become dismembered, sodden, and pale 
brownish yellow, but never dark brown. 

The hyphae of the fungus travel upwards in the lumina of 
the elements of the wood, and easily pierce the walls of the cells. 
As they send off lateral branches they also invade the cells of 
the medullary rays, as well as adjoining tracheides. As already 
mentioned, the first perceptible change in the wood occurs in the 
contents of the living cells, which become brown and partly 
disappear. This is succeeded by the conversion of the lignified 


cell-wall into cellulose, a change which begins on the side next 
the lumen, and advances outwards. The cellulose is soon 
completely dissolved, and at last the delicate skeleton of the 
middle lamellae also disappears. At certain points this process 
proceeds with great rapidity. Here and there, for instance, one 
finds that the tracheides, in immediate proximity to the medullary 
rays, are filled with a brown fluid, which has probably originated 
in the latter, and which discolours and nourishes the mycelium 
to such an extent that a brown " mycelial nest " is formed. So 
energetic is the action of the ferment in the neighbour- 
hood of these bodies that the encrusting substances entirely 
disappear from the adjoining tracheides, which, to the dis- 
tance of several millimeters, are completely transformed into 
cellulose, and thus become colourless — in other words, white. 
Almost immediately after being converted into cellulose the 
middle lamella disappears entirely, and the individual elements 
of the wood thus become isolated, so that, when disturbed by a 
needle, they fall apart like the strands of asbestos. Gradually 
they are dissolved, and holes, which are constantly increasing in 
size, are formed in the crumbly wood. 

While the mycelium thus decomposes the wood, sometimes to 
a height exceeding eight yards, the parasite advances much 
more slowly in the cortex, where its presence is betrayed b}- three 
distinct phenomena. From the point of infection the mycelium 
spreads both towards the root-apices and towards the stem. It 
kills the cortex, and consequently the root, and when, after some 
years, it has reached the stem, it spreads from the stool on to 
roots that have hitherto remained sound. As soon as these are 
also attacked by the disease, the tree dies. 

A second function of the mycelium that grows in the cortex 
consists in the formation of sporophores, which appear here and 
there between the bark-scales of the roots or stool. These lead 
to the production of fresh seats of disease in the plantation, as 
has been already described. 

A third function is concerned with the spread of the disease 
subterraneously owing to infection by the mycelium. Where a 
diseased root comes into contact with the sound root of an 
adjoining tree (Fig. 121), or where the two are positively grown 
(grafted) together — as may very frequently be observed in a dense 

I go 


wood — the mycelium, which appears as a small cushion between 
the scales, grows into the cortex of the neighbouring tree. It is 
easy to induce infection artificially by taking a piece of cortex 
containing a portion of living m}xelium still capable of growth, 
and binding it firmly to the cortex of the root of another tree. 

Owing to the mycelium spreading subterraneously from tree 
to tree, these well-known gaps, which increase in size each year 
by the death of the marginal trees, appear in woods. At one 
time no reason whatever could be assigned for the peculiar 
behaviour of these gaps. On account of the decomposition of 
the wood proceeding rapidly and advancing far up the stem, and 

as it is succeeded by the 
death of the tree, the dis- 
ease is to be classed with 
the most dangerous forms 
of " red-rot." It is very 
abundant in the pine 
forests of North Germany,* 
and quite as much so in 
the spruce woods, especi- 
ally where these are situated 
in hilly districts. There is this difference to be noted, however, 
that when pines are killed by the parasite it is usually only their 
roots that are affected and rotten, the stem, with the exception 
of the stool, showing no signs of decomposition. The wood of 
the stool is generally strongly impregnated with resin, and I 
believe I am right in concluding that it is the abundant resinous 
contents of the pine, which are especially prominent in the lowxr 
part of the stem, that form a barrier to the upward growth of the 
mycelium of the fungus. In the case of the spruce, on the other 
hand, and of the Weymouth pine, w^iich is poor in resin, decom- 
position of the wood spreads high into the stem. 

It appears to be necessary to keep a watchful eye upon 
coniferous woods at all stages of their growth, so that diseased or 
dead trees may be instantly removed. In the case of the older 
woods, one may isolate the diseased spots by surrounding them 
with narrow trenches, and by severing all roots that may be 
encountered. In order to attain this end we must, of course, 
* [It is also by no means uncommon in this country. — Ed.] 

Fig. 121. — The thinner root has been killed by 
T. radicipcrda, and the stronger one has 
been infected at the point of contact. The 
disease has spread as far as the dark shading. 
One eighth natural size. 


form the trenches at such a distance from the diseased gaps as 
to warrant the assumption that all trees already infected are 
included in the isolated area. As a rule it suffices to include the 
trees nearest to the margin of the gap. If the workman notices 
that a dead root crosses the trench, then it will be necessary 
at that point to divert the course of the trench farther into 
the wood, otherwise the labour will be in vain. Although this 
operation is a certain preventive when it is properly performed, 
its careful supervision is so difficult when conducted on a large 
scale that I am doubtful if a general adoption of the practice is 
to be recommended in commercial sylviculture. The objection 
that the sporophores develop in the trenches does not appear 
tenable, because it is a simple matter to examine the trenches 
once a year and to remove such sporophores.* When the fungus 
appears at many points in a wood, even the most careful isolat- 
ing is of no avail. The gaps should either be filled up with 
dicotyledonous trees, or if, for any reason, this is deemed im- 
practicable and conifers must be employed, then the young trees 
must be carefully watched so that infected plants may be rooted 
out and the disease be promptly checked. 


This parasite is exceedingly abundant in the pine woods of 
North Germany.-f- In South Germany, where it is less common, 
it is met with chiefly in spruce woods. It also occurs in the 
spruce woods of the Harz, the Thuringian Forest, and Schleswig, 
as well as in the larch and silver fir woods of the Riesengebirge. 

It produces a so-called bark-shake, ring-shake, or heart- 
shake, which nearly always commences at a branch, and there- 
fore usually in the crown of a tree, 

The brown woody sporophores, which attain an age of fift}' 
years, vary in shape between an incrustation and a bracket. In 
the case of pines and larches they occur only on the part of a 
stem where a branch has fallen off (Fig. 122), while in spruces 

^ R. Hartig, Wichtige Kratik]ieiteti def Waldbdunie, p. 43. Zcrsetztings- 
erschei7iungen des Holzes, p. 32, Tables V. and VII. 

*[If Brefeld's account of the conidial fructification is correct, this maybe 
a more difficult matter than appears. — Ed.] 

"t" [Also occurs in this country. — Ed.] 



and silver firs they may spring directly from the bark as 

The spores which are annually produced in these sporophores 
are scattered by the wind, and should they gain a footing on a 
fresh branch-wound which is not protected by a covering of 
resin they push their germ-tube into the stem, and the mycelium 

Fig. 122. — Part of the stem of a pine bearing the sporophore of T. Piiii. a, healthy 
alburnum ; b, wood saturated with resin in the neighbourhood of the sporo- 
phore ; c, decomposed wood ; d, the canals in which the spores are produced ; 
e, older canals which have become filled up by fungus-tissue ; /", the upper 
surface marked by zones. One half natural size. 

spreads partly upwards and partly downwards. The younger 
class of trees enjoy immunity from infection, because in their 
case wounds are very quickly protected by an exudation of 
turpentine. From the time when the heart-wood becomes 
comparatively dry, turpentine ceases to exude from the central 
part of a branch-wound, and this consequently becomes liable to 
attack from the spores of the fungus. This accounts for the 



disease not usuall}- appearing upon trees younger than fift}' 

The mycehum shows a preference for growing longitudinally in 
the stem, while its distribution horizontally is most pronounced 
in some particular annual ring. On this 
account decay often takes the form of ring- 
shake — that is to sa}-, it is most pronounced 
in peripheral zones which encircle a part or 
the whole of the stem. At first the wood 
becomes somewhat deeper red brown in 
colour, and then white blotches or holes 
appear here and there. In the case of 
the pine especiall}^ these are largely con- 
fined to the spring wood of some par- 
ticular annual ring, in which the}' enlarge 
parallel to the longitudinal axis of the 
stem, the result being that the resinous 
zone of autumn wood ma}- persist alone 
for a long time, until it also is destro}-ed 
b}' decomposition. 

A resinous zone is formed along the 
boundary between the alburnum and the 
decomposed wood, and this interferes with 
the outward progress of the mycelium of 
the fungus. In those specimens which I 
have examined, this zone is absent onl}' in 
the comparatively non-resinous silver fir 
and in spruce-branches, and thus, in their 
case, the fungus is able to reach and pene- 
trate the cortex with ease. The action of 
the ferment of the parasite produces white 
spots, similar to those that have been de- 
scribed in the case of T. radicipcrda. The 
lignin is extracted from the cell-walls, and 
pure cellulose is left. The middle lamella 

is completely dissolved as soon as the lignin disappears, and 
thus the tracheids become isolated before being finally dissolved 
(Fig. \2i,a a to /;). The lamella which is nearest to and bounds 
the lumen persists the longest, and before it is dissolved the 


Fig. 123. — A tracheid 
of P. sylvestris which 
has been decomposed 
by T. Pini. The prim- 
ary eel I -wall has been 
completely dissolved as 
far as a a. In the lower 
portion of the figure the 
secondary and tertiary 
walls consist of cellu- 
lose alone, in which 
granules of lime are 
distinctly recognizable, 
/' ; filaments penetrate 
the walls and leave 
holes behind, d, c. 


ash-constituents of the wall impart to it a finely gi-anulatcd 

The fungus is only in a position to produce sporophores when 
it has developed luxuriantly in the interior of the tree. In that 
case the mycelium pushes its way outwards at a place where the 
base of a dead branch opens a passage through the alburnum. 
At such a place the sporophores are produced, and should these 
be forcibly removed a number of new ones are, as a rule, formed 
in a short time. 

It is thus seen that nothing is gained by removing the 
sporophores, but trees infested by the fungus should always be 
removed in the thinnings and other fellings. We thus remove 
the danger of infection, and utilize the trees before they are 
rendered entirely valueless by the advancing decomposition. It 
frequently happens that, although fungi are visible on the upper 
region of the tree, the disease has not extended down to the 
lower and valuable part of the stem, so that, after cutting off the 
upper portion, some good useful timber is available. If one 
waits till the final felling before removing the fungus-infested 
trees, all that one gets is very worthless rotten wood. Of course 
a stop must also be put to pilferers breaking or sawing off 
green branches, as this practice increases the probabilities of 
infection. Old branches that have died naturally cannot be 
attacked by the fungus. 

This disease is most prevalent in the neighbourhood of towns 
and villages, where the pilfering of branches is common, and in 
woods that are much exposed to the wind, and where, conse- 
quently, branches arc frequently broken off. 

When I first described this parasite of the silver fir and spruce, 
I made the remark, " Whether this is a new species distinct from 
Polyponis fiilvJis can only be decided by a careful study of the 
allied species of this genus. In the meantime it may pass 
under the name of P.fiilvus."^ Since then it has been recog- 
nized as an undoubtedly new species, and has been introduced 
into the literature of the subject by Herr Allaschen under the 
name P. Hartigii. 

1 R. Hartig, Die Zc}-set::u/!gserschcininio-e?i dcs Hohcs, p. 40. 


This fungus producesa kind of white-rot in silver firs and spruces, 
and is very frequently encountered in association with ALcidmm 
elatiiunn. Apparently infection is most liable to occur naturally 
on those parts of the cancerous swellings where the cortex has 
ruptured and exposed the wood. The mycelium, which is at 
first very vigorous, is yellowish in colour, and produces numerous 
short lateral branches, which are twisted in a worm-like manner, 
and are apt to fill up the cavities of the bordered pits of the 
tracheids. This vigorous mycelium gives off a few exceedingly 
delicate lateral hyphae, which bore through the walls, in which 
they form very minute holes. Only in the later stages of de- 
composition is the disappearance of the middle lamella effected, 
after which the inner walls are also dissolved, having first been 
greatly attenuated and then temporarily isolated. At this stage 
the mycelium is of extraordinary fineness. The wood of the 
silver fir appears yellowish, clear oblong patches being observable 
if carefully looked for on a smooth surface. The vigorous 
yellow hyphze induce the formation of narrow dark lines at the 
boundary of the sound wood. 

As the silver fir cannot form a strongly resinous zone, it is 
unable to prevent the progress of the mycelium into the 
youngest layers of wood. The mycelium consequently grows 
outwards with ease into the cortex, and, having advanced far 
enough, it produces the sporophores on the surface. These are 
at first hemispherical, but in the course of years they become 
more and more bracket-like in shape. Externally they are 
yellowish brown on the hymenial surface, but elsewhere they are 
ashy grey, almost smooth, display no zones, and are beset with 
exceedingly minute punctures or pits. The interior, which is 
tawny and lustrous, shows distinct zones, except in the region of 
the pore-canals, which increase in length each year at their 
lower extremity. 

As it is found that silver firs with cancerous swellings sooner 
or later break at the diseased spot owing to snow or storms, 
it has become the custom in many districts — for instance, 
in the Black Forest in Wiirtemberg — to fell all cankered trees 
during the thinnings, even when such trees belong to the larger 
class. In this way the spread of Polyponis Hartigii can best be 

O 2 



This fungus produces an exceedingly characteristic form of 
white-rot in the spruce. In the Salzburg and Bavarian Alps, 
and in the spruce woods near Munich, it is the commonest form 
of decomposition in the spruce, and I have also noticed it in the 
Harz. Infection takes place, and the sporophores are produced, 
above ground. On account of their white colour the sporo- 
phores are conspicuous even at a distance. They are annual, 
more or less bracket-shaped, and frequently superimposed the 
one above the other and grown together. They are verj- 
watery, somewhat sodden on the upper surface, and destitute 
of zones. 

The colour of the wood is but little altered by the decompo- 
sition. It becomes brownish yellow, and horizontally disposed 
holes filled with mycelium appear in vertical rows in the spring 
wood. These holes, which stand i to ih mm. apart, impart to 
the wood an appearance which reminds one of the finest graphic 
granite. The wood constantly becomes lighter and more friable, 
but the peculiar appearance is retained till the end of the final 
stage of decomposition. 

Should the wood be exposed, without chying, when decom- 
position is beginning, the mycelium will grow outwards to form 
a white skin, the m}xe]ial strands of which are chiefly disposed 
in a horizontal direction. 

Growth and decomposition are in several ways characteristic. 
The hyphit, which in the first stage of decomposition are yellow 
and stout (Fig. 124, a, b), are replaced by more delicate fila- 
mentous mycelia as decomposition proceeds, until at last the 
hyphse which are formed can onl}' be seen b)^ the aid of a very 
powerful microscope. The mycelium has a marked tendency to 
grow to some extent in a horizontal direction, at right angles to 
the long axis of the elements (Fig. 124, t), the chief result being 
the formation of the above-mentioned horizontal holes in the 
wood. Why these are formed only at definite distances from 
each other I have not been able to determine. Dissolution of the 

^ R. Hartig, Zcrsetzungsei'scheimingen, pp. 54 et seq. 
* [This is quoted as a British species. — Ed.] 



cell-wall begins at the lumen and proceeds outwards, being pre- 
ceded by the conversion of certain layers of lignin into cellulose. 

Fig. 124.^ — Decomposition of spruce timber by /'. horealis. a, a vigorous mycelium 
in a tracheid containing a brownish yellow solution which has originated in the 
medullary rays. In b and c the mycelia are still brown in colour and very 
vigorous. At d and e the walls are much attenuated and perforated, and the 
mycelia, not being so well nourished, are very delicate. At /" the pits are almost 
entirely destroyed. At g and h only traces of the walls remain. The destruction 
of the bordered pits is to be followed from i to r. At i the bordered pit is still 
intact, at /■ one \\all of the lenticular chamber has been largely destroyed, its 
inner boundary being marked by a circle. At / one side of the bordered pit 
has entirely disappeared. A series of pits is shown from ui to n, in each of 
which only a single delicate wall has been preserved — namely, that which is pro- 
vided with the closing membrane. In preparing the section a crack has formed 
in this wall. From to r pits are shown where both of the walls have been 
partially or completely dissolved, and only at p and q does one perceive the 
thickened portion of the closing membrane. At s one can plainly see the spiral 
structure of both cell-walls, which when united form the common wall between 
two tracheids. At t mycelia are to be seen traversing the tracheids horizon- 

The thin middle lamella persists longest, being converted into 
cellulose and dissolved only after the internal portion of the wall 
has entirely disappeared. 



The decomposition produced b}- this and the following parasite, 
P. ScJnveinitzii, bears a very strong resemblance to that caused 
by the dry-rot fungus Memlius lacrymans. 

P. vaporaj'ius is exceedingly common on spruces and pines, 
both roots and wounds above ground being attacked. It very 
frequently effects an entrance through a wound due to the 
barking of red deer. The wood becomes reddish brown, dr}-, 
and fissured, and as time goes on the resemblance to half- 
charred timber becomes more and more apparent. When rubbed 
between the fingers it crumbles into yellow dust. The m}-- 
celium is specially liable to develop in cracks, or between 
the dead wood and the bark, in the form of snow-white 
much-branched woolly felted strands, similar to many of the 
mycelial growths of Meniliits lacryvians. Although I have 
made no direct observations on the point, still I think it prob- 
able that the mycelial strands which grow on dead roots and 
stools may convey the disease subterraneously to adjoining trees. 
The sporophores, which are pure white, form incrustations, 
but never brackets. These originate on decayed wood or 
dead bark, or on luxuriant mycelial growths or strands. This 
fungus very frequently appears on the timber of buildings 
where, on account of its luxuriant mycelial growths, which 
have sometimes a fasciated, sometimes a strand-like appear- 
ance, it is usually confounded with Mernliiis lacrynians, whose 
mycelial growths, however, always assume an ashy grey colour 
shortly after being formed. As regards its importance as an 
agent in inducing decay in buildings, I may refer to the remarks 
which I shall have to offer when discussing M. lacrynians. 


In describing this parasite in Zerset::nngserscJieiunngen dcs 
Holzes'^ I have called it Polyporus luollis. This mistake was 

^ R. Hartig, Zosctziingsci-schciniiugcn, pp. 45 ct scq., and Table VI I L 
- P. 49- 

*[\'ery common on dead wood, and I have found it on the decayed wood 
of a green-house. — Ed.] 
t [Quoted as British.— Ed.] 



due to my having access onl}- to old dr}' 
rendered the correct identification difficu 
Professor Magnus has correctl)- identified 
the fungus as P. ScJnveinit::ii. It appears 
on the Scotch pine, the Weymouth pine, 
and the larch. 

The decomposition which it produces 
very closely resembles that due to the 
preceding species, but in the present case 
the white branching mycelial strands are 
absent, the mycelium at most growing 
out of the fissures as a fine chalky in- 
crustation. The smell of the wood, which 
is very characteristic and intense, reminds 
one of the smell of turpentine, without 
however being perfect!}^ identical. 

The sporophorcs, which appear on the 
dead wood or project from the bark-fissures 
of living trees, take the form of reddish 
brown cushions, which afterwards assume 
a somewhat bracket-like shape. The porous 
layer, which is yellowish green when }-oung, 
assumes a deep red colour if ever so slightly 

As decomposition advances the tracheids 
exhibit spiral cracks and fissures (Fig. 125). 
Apparently these cracks are due to the 
shrinkage of the wall-substance,which always 
remains fairly dr}-. It is owing to these 
cracks that the wood is so easil}- pul- 

P. vaporarins also induces cracks and 
fissures in the cell-walls, but, instead of 
extending completely round the cell-lumen, 
these are small, and are arranged in large 

■ sporophorcs, which 
It. In the interval 

Fic'r. 125. — Tracheid o 
Finns destroyed by /'. 
Schweinitzii, The eel 
lulose has been largely 
extracted, the walls 
consisting chiefly of 
lignin. Cracks occur 
in the secondary wall 
when dry, while the 
primary wall, a h, re- 
mains intact. The 
spiral structure of the 
secondary wall is the 
cause of the fissures 
in the walls of adjoin- 
ing cells crossing at the 
bordered pits, c, and at 
the punctures, d , e. 
Where pits and punc- 
tures are absent the 
fissures ai'e simple, f. 

numbers in vertical 



This is one of the most widely distributed parasites of the oak, 
Robinia, alder, tree-willows, poplars, walnut, and pear. It also 
occurs as a parasite on the common larch. Infection takes place 
through a branch-wound, and the mycelium spreads rapidly in 
the wood, causing it to become red brown and dry. The wood 
reveals numerous cracks, into which the mycelium grows, to form 
laro-e sheets of felted hyphae. In the case of dicot}-ledonous 
trees the vessels become filled, in the early stages of decom- 
position, with a dense fungoid growth, so that, on a transverse 
section, they appear as white spots, and, on a longitudinal section, 
as white lines. The walls of the elements of the wood become 
brown and very rich in carbon, and shrink greatly, but on being 
treated with dilute caustic potash they swell up and become 
almost completely dissolved. The spiral cracks, which always 
ascend from right to left in the interior of the fibres, never 
extend into the middle lamella. 

Whenever old snags, or any kind of wound, admit of the 
mycelium reaching the surface, a group of sporophores is annually 
formed. These are succulent, of a pale sulphur-yellow colour 
beneath, and pale reddish yellow on their upper surface, and by 
their size and strikingly luminous colour they readily attract 
attention. The pileus is internally of a white colour and cheesy 
consistenc)'. The pores reveal a hymenial layer with clavate 
basidia. The mycelium of this fungus very frequently develops 
numerous round gonidia in the wood itself, and during my early 
investigations on this parasite I regarded these as belonging to 
a different species of fungus. It very frequently happens that 
before diseased trees are overthrown by storms, their tissues, on 
one side or other, die as far out as the bark, and the latter, 
withering, drops off and allows the red-brown decayed wood to 
fall out from the inside of the tree. Thus it is not impossible that 
the gonidia may be carried into the air along with the dust of the 
decayed wood, and so assist in the distribution of the parasite. 

1 R. Hartig, Zersetzungserscheimmgen^ pp. no et seq. De Seynes, 
Rccherches pour servir a Vhistoh'e naturelle dcs vegctaux inferieia's, 1888. 

* [Verj' common in this country. I have frequently collected it in Windsor 
Park and elsewhere. — Ed.] 



This is the parasite most frequently met with on the majority 
of dicotyledonous trees. My investigations on its destructive 
effects on wood have been conducted for the most part on the 

Infection may occur on branches or on bark-wounds, and the 
myceHum spreads rapidly in the wood. At first the wood 
assumes a deep brown colour, and this is succeeded by yellowish 
white decomposition, which is the commonest kind of white- 
rot in the oak. The yellowish white wood constantly becomes 
lighter and softer, and resembles in its properties the cellulose 
that is used in paper-making. The h}'phae, which are at first 
very strong and afterwards extremely delicate, completely fill 
up the elements, and induce a form of decomposition which is 
characterised by the inner layers of the walls being converted 
into cellulose and dissolved, before the middle lamella, which 
persists for a long time as a delicate skeleton, undergoes similar 
changes. It will thus be seen that the process of decomposition 
bears a close resemblance to that described under P. borealis. 
The sporophores, which usually spring directly from bark that 
is infested by the mycelium, are at first hemispherical, but 
afterwards become more or less hoof-shaped. Although they are 
familiar enough, it may be mentioned that they differ from 
those of P. Hartigii, which they resemble in external appearance, 
by exhibiting concentric zones, and frequently cracks as well, on 
their upper surface, while internally the layers of pores are also 
interrupted by the zones. 


This fungus of the oak produces a form of decomposition 
in which oblong yellowish or white blotches occur, surrounded 
by firm wood which displays the original colour of the duramen. 

^ R. Hartig, Zersetzungserschcinitngcn, pp. \\\ ct jit^'., and Tables XV. 
and XVI. 

^ Ibid., pp. 124 et scq.^ and Table XVII. 

*[One of the commonest fungi in Windsor Park and neighbourhood, 
especially on old Beeches. — Ed.] 

t [Common in Britain. — Ed.] 

202 diseasp:s of trees 

The white blotches consist of elements which have been con- 
verted into cellulose, and which have become isolated by the 
solution of the middle lamella. The yellowish parts, on the 
other hand, reveal a form of decomposition of the cells which 
is exceedingly like that due to P. igniarhis, and which is cha- 
racterised by the middle lamella persisting the longest. The 
white patches are the first to be dissolved, and thus holes, sur- 
rounded by very hard sides, are formed. When freel}- exposed 
to the air the wood assumes a cinnamon brown colour, and is 
replaced by a mass of firm brown hyphae. 

The large hoof-shaped annual sporophores arc of a cinnamon 
brown colour, and appear on the bark or on the spots previously 
occupied by branches. They possess so little durability that 
one but seldom meets with a perfect specimen. 

Should P. dryadeits and P. igniariiis simultaneously attack an 
oak, and should their hyphae come into contact, a peculiar kind 
of decomposition occurs along the line where the hyphae of the 
two species meet. The wood becomes yellowish white, the 
decomposition being similar in appearance to that which is 
induced by P. igniarhis alone. All the longer medullary rays^ 
however, are represented by snow-white bands, which, on being 
investigated, are often found to consist of nothing but unaltered 
starch-grains, while the cell-walls have been almost entirely 
dissolved, or have been converted into cellulose. 


A parasite is frequently met with on oaks and beeches whose 
yellowish white sporophore takes the form of an incrustation or 
bracket, and which is distinguished by the hymenium being 
disposed on downward-directed spines of unequal length. The 
hymenium, which is at first simple, periodically increases in 
thickness by the hyphae growing through the last layer to form 
a new hymenium. In the lower portion of the spines especially 
this process is repeated five to eight times, the result being that 
the spines increase greatly in thickness, and the hymenium 
displays five to eight layers. 

In this case also the decomposition, which spreads from 

^ R. Hartig, Zersetziingscrscheintingcn^ pp. 97 ct seq., and Table XII. 
*[This is also British. — Ed.] 


infected wounds on the stem, takes the form of a white-rot. 
The colour is )-ellowish ashy grey, alternating with stripes of a 
pale brown colour, which generally persist longest in the 
medullary rays. In the later stages of decomposition snow-white 
masses of felted mycelium occur where a zone of spring wood 
is much deca}'ed. 

The peculiarity of the action of the ferment consists in the 
inner layers of the cell-walls swelling up into a gelatinous mass, 
without being converted into cellulose, before they are completely 
dissolved ; the middle lamella being the last to disappear. 


A form of disease which is very common in the oak throughout 
the whole of German}- is known as " partridge wood," on account 
of the peculiar discoloration w'hich it induces in the wood, and 
which reminds one of the white-speckled feathers met with on 
certain parts of the body of the partridge. At first the diseased 
wood assumes a deep red brown colour, and then white blotches 
on a dark ground make their appearance which stand in a certain 
relationship to the large medullary rays. These blotches after- 
wards become transformed into sharply defined cavities with a 
white lining. As the cavities, which are separated from each 
other by firm brown wood partitions, increase in size, the wood 
looks as though it had been attacked by ants, and, as a matter 
of fact, the symptoms are often mistaken for the work of these 
creatures. It is to be noted that each ca\'it}- usuall}- remains 
distinct until the stage of complete decomposition is reached. 
In the wood of the oak the mycelium first induces the contents 
of the parenchymatous organs to become brown. Graduall}- 
proceeding inwards, the starch-grains fail to give a blue reaction 
with iodine, colourless granules persisting for some time in the 
central cells of the medullary ra}-s, until the}' also are at last 
destroyed (Fig. 126). 

Where the white blotches make their appearance, as also in 
the partitions of the white cavities, all the organs are converted 
into cellulose, and the middle lamellae being dissolved the 

^ R. Hartig, Zcrsctzungscrschcimingcii^ pp. 103 et seq. 

*[I do not know this as British, but a specimen of diseased wood sent 
from India was marked in exactly the way Hartig describes. — Ed.] 


individual elements of the wood become isolated (Fig. 126, e — /i). 
It is remarkable that the process of decomposition in the 
neighbourhood of the cavities undergoes a change when these 
have become enlarged. The latter no longer appear white but 
greyish yellow, and reveal abundant felted mycelia, which pierce 
the walls at numerous places. Instead of a conversion into 

Fig. 126. — Decomposition of oak induced by T. Peniix. a, tracheids containing 
some filamentous mycelia, and showing a few perforations on their walls ; b, 
wood-parenchyma containing starch which is partly undergoing solution, the 
outer granules being the first to disappear ; (", vessels containing hyphse of the 
fungus ; d, sclerenchymatous fibres containing fungus-filaments, and showing 
perforations in the walls ; c and /, tracheids which are completely isolated by 
the solution of the primary wall. The thickened rings of the bordered pits are 
also seen to be isolated between the tracheids. As the elements are isolated, the 
cracks no longer cross at the bordered pits, g, cells of wood-parenchyma which 
are completely isolated and almost completely dissolved ; h, a tracheid just 
before final solution ; ?', sclerenchymatous fibre much decomposed ; k, a 
tracheid whose wall has been dismembered by cracks before being dissolved. 

cellulose resulting, the wood-substance is dissolved, partly by the 
enlargement of the perforations and partly by the centrifugal 
attenuation of the cell-walls. 

The sporophores of the parasite occur as incrustations in 
fissures or other cavities in the diseased wood, or on the outside 
of dead branches. The incrustations, which vary in thickness 
from rrV to J inch, are brownish yellow in colour, and consist of 


a layer of hjq^haj disposed at right angles to the surface. The 
hyphas end in somewhat club-shaped basidia, which are covered 
b}' peculiar hair-like outgrowths. Only a certain number of the 
basidia produce spores (four in each case), those which remain 
sterile producing a new hymenium in a succeeding period of 
growth, and in doing so they anastomose here and there b}* 
lateral budding. On a transverse section a sporophore, depending 
on its age, shows more or less distinct strata, of which only the 
youngest possesses a pale colour, the others being of a deep 
brown hue. When dead the whole of the sporophore appears 
dark brown. 


A ver}' striking and characteristic form of decomposition in 
the oak is produced by S. hirsutiun. In practice such wood is 
called " yellow piped " or " white piped." Usually a brown 
colour first makes its appearance in certain concentric zones, 
which to begin with are confined to one side but afterwards 
encircle the whole of the stem, and later on a longitudinal 
section will show detached snow-white or yellow stripes which 
appear as white spots on a cross section (" fly wood "). When 
the oxygen of the air has free access, as in the alburnum, 
branch- snags, &c., the whole of the w^ood is frequently converted 
into a uniform yellow mass. It scarcely seems to admit of 
doubt that this fungus also plays an important part as a sapro- 
ph}-te, and finds its way on to branches that are dying naturally. 
In the white stripes the mycelium converts the wood into cellu- 
lose, and when the middle lamella disappears the elements 
become isolated. In the yellow parts of the wood, on the other 
hand, the solvent action proceeds from the lumen outwards, as 
in the case of P. igiiiarins, and this is not preceded by a con- 
version into cellulose. The sporophores, which usually develop 
on the bark, appear first as a crust, but afterwards their upper 
edge — which is brown, faintly zoned, and covered with stiff hairs 
— projects in a distinctly horizontal direction. 

' R. Hartig, Zersctziingserscheittiin^^en, pp. 129 ct scq.^ Tabic XVIII. 
*[\'ery common in England. — Ed.] 


The familiar tinder-fungus, which occurs on beeches and oaks, 
produces a form of white-rot, and its mycelium has a tendency 
to form luxuriant patch-like or skin-like growths in fissures of 
the decayed wood. So far it has not been made the subject of 
thorough investigation. 


Occasionally P. beiuliiuis is to be found abundantly developed 
on the birch. Its hirsute sporophore, which is white beneath and 
brownish grey above, is at first globular, but afterwards takes 
the form of an inverted bracket with a convex upper surface. 
The decomposition induced by the parasite is a form of 


This parasite produces a form of white-rot in the birch. Its 
sporophore appears on the surface of the bark as a dark brown 
porous incrustation. 

Numerous other species of Polyponis doubtless occur as 
parasites on the wood of trees, but these, so far, have not 
been subjected to investigation. 

The following fungi are also worthy of mention. Dcudalca 
quercina\ is met with everywhere on old oak-stumps, where 
it forms large bracket-like sporophores which bear the hymenium 
partly in pores and partly on lamellae. During decomposition 
the oak-wood assumes a grey brown colour. Having found the 
fungus vigorously developed on branch-wounds of the older 
classes of oaks, I suspect that it is also a parasite. 

Fistulhia hcpatica., the Beef-steak Fungus, produces a deep 
red brown decomposition in the wood of the oak. 

All the above-mentioned wood-parasites, which obtain an 
entrance through wounds above ground, can only be combated 
in one or other of the following ways. First, great care must be 

1 D. H. Mayr, Hot. Centnilblatt, 1885. ^ Ibid. 

* [Common in Britain.— Ed.] 

t [1 have frequently collected this in this (Cooper's Hill) part of England. 

J [Both this and the next occur in Surrey and elsewhere in Britain. — Ed.] 



taken to do nothing that will cause the formation of wounds 
in trees, of which more will be said in the section on wounds ; 
and, secondly, where wounds are intentionally produced on trees, 
as in pruning, the necessary prophylactic measures must be at 
once put in force, and in particular an antiseptic dressing in 
the form of a covering of tar should be provided. 

At the same time woods should be kept tidy and free from 
decaying wood, which may bear the sporophores of parasites, 
but this must not be taken to mean that all old oaks that 
are already decayed are to be felled without regard to other 
considerations. For the sake of effect the forester should allow 
old trees and picturesque bits of timber to stand where deemed 
desirable in the neighbourhood of frequented paths, even 
although the benefits of so doing may not be at once manifest 
in the shape of hard cash. 


This fungus belongs to the most widely distributed and 
destructive of parasites. It lives parasitically on alTEuropean 
conifers, besides destroying those that have been introduced 
from Japan, America, &c., and I have even recognized it in the 
fossil wood of Ciipressinoxylon. Amongst dicotyledonous trees it 
appears to occur as a parasite on Pninits avium and P. doiiiestica, 
while as a saprophyte it is to be met with everywhere, not only 
on the dead roots and stools of all dicotyledonous and coniferous 
trees, but also on the structural timber of bridges, conduits, mines, 
&c. It has frequently been asserted that it also occurs as a 
parasite of the vine, but I have had no opportunity to convince 
myself of the correctness of this view. Those rhizomorphs 
whose occurrence I have hitherto observed on the vine belonged 
to Deniatophora necatrix. 

The disease often manifests itself on plants only three to five 
years old, though it also destroys spruces, pines, &c., a century 
old. One recognizes it by removing the bark at the collar 

^ R. Hartig, Wichtige KrankJieitcii dcr Waldbaumc, 1874, pp. 12 et seg., 
Tables I. and II. R. Hartig, ZcrsctziingscrscJtcinuugcii^ pp. 59 ct seq.^ Table 
XI., Figs. 1-5. 

* [This is one of the commonest of British fungi, and its rhizomorphs and 
sporophores are well known. — Ed.] 



and on the roots, when a firm snow-white myccHum (Fig. i2y,cc) 
is observed, which, in the case of the older class of trees, some- 
times ascends under the bark while the tree is still alive to the 
height of ten feet or more. Brownish black lustrous strands, 

2-V to yV inch in thickness, which occa- 
sionally anastomose, are observed in 
greater or less abundance adhering to 
the roots. These are met with in con- 
junction with the sheets of white my- 
celium under the cortex, though some- 
times they merely embrace the roots 

A great deal of turpentine and resin 
frequently adheres to the outside of the 
stronger roots, and this, mixing with the 
particles of soil, forms a firm mass round 
the collar (Fig. 128). The diseased trees 
speedily succumb, and are seldom to be 
recognized more than a }'ear before their 
death by their pale colour or stunted 
shoots. If we carefully dig up a plant 
that appears to be perfectl}' healthy, in 
the immediate neighbourhood of one 
that is manifestly diseased or dead, we 
will as a rule discover on the roots one 
or more places of infection where a black 
rhizomorph strand has bored into the 
cortex (Fig. 127, a). When the cortex is 
carefully removed the strand will be ob- 
served expanding from the place of 
entrance into a snow-white bod}' (Fig. 
127, b), which spreads in the cortical 
tissues and causes browning and death, 
as far as it reaches (Fig. 127, c c). The 
mycelium that grows in the living cortex 
is characterised b}- its fasciated and skin-like appearance. It 
very easily resumes the round strand-like form, which may 
cither grow to the outside of the roots or proceed to develop 
between the wood and cortex. When, owing to the death of 

Fig. 127. — The living root 
of a spruce, showing two 
spots, a, h, where the 
rhizomorph has entered 
.ind infected the cortex. 
The cortex has been re- 
moved from the larger 
root, d to d, in order to 
show the mycelium, c c, 
which has gained an en- 
trance at h. 



the tree, the shrinkage of the cortex affords space for the de- 
velopment of these strands, they anastomose abundantly, hke so 
many twigs, and envelop the wood of the stem in a reticulate 
fashion. The rhizomorphs that spring from the roots progress 
underneath the surface of the ground, at a depth seldom exceed- 
ing four inches, and bore into any sound roots of conifers that 
they may encounter, and thus the disease is spread from tree 
to tree (Fig. 127). In autumn, from the end of August till 

Fig. 128. — A young pine which has been killed by 
A. tnelleiis. Numerous sporophores are seen which 
have broken through the cortex at the surface of 
the ground. Branching rhizomorph- strands are 
present on the roots. 

Fig. 129. — A sporophore 
of A. nielleus which has 
developed on a rhizo- 
morph, a lateral branch 
of which has produced 
only abortive sporo- 

October, the large familiar sporophores (Fig. 129) may be 
observed developing on the rhizomorphs which grow in- 
dependently in the ground, or projecting from the cortex, 
especially the collar (Fig. 128), of trees that have been killed by 
the parasite. For further details I may refer to what I have 
published in the works already alluded to. The white spores of 
this Hymenomycete, which are spread either by the wind or by 
being brushed off by passing objects, develop first of all a 
filamentous mycelium, and from this the m)xelium form 
designated Rhizoinorpha is produced, as is easily proved by 



sowing the spores in a decoction of plums. The pathological 
symptoms can onl\- be explained in the light of the peculiar 
organization of the mycelial growth that lives in the cortical 
tissues. The apex of the rhizomorphs (Fig. 130) consists of 
delicate pseudo-parench}-ma, which, elongating by the division 
and growth of the cells, produces delicate hyphae on the inside 
at a certain distance from the point, whereby a felted tissue, 
called the medulla, is produced in the interior. The outer parts 
of the pseudo-parenchyma (Fig. 130, c), on the other hand, 
coalesce to form the so-called rind (Fig. 131, d), which when 
young gives off numerous delicate hyphae, and these, taking 
advantage of the medullar}- rays, penetrate the wood, and 
especially the resin-ducts, should such be present. In the wood 
the growth is upwards. This filamentous mycelium, which 
progresses much more rapidh' in the interior of the wood 
than the rhizomorphs which grow in the cortex, completely 
destroys the parenchyma that exists in the neighbourhood of 
the resin-ducts, and to all appearance this is accompanied 
by a partial conversion of the cell-contents and the cell-walls 
into turpentine (Fig. 131). The turpentine sinks down under 
its own weight, and in the collar, where the cortex is 
withered, having been killed by the rhizomorphs, it streams 
outwards, pouring partly in between the wood and the cortex, 
and partly into the surrounding soil at places where the cortex 
has ruptured owing to dr}'ing. On this account the disease was 
formerly called " Resin-flux " or " Resin-glut." In the upper 
parts of the stem, where the cambium and cortex are still sound, 
the turpentine also flows lateralh', b}- means of the ducts of the 
medullary rays, from the injured canals towards the cambium and 
cortex. In the latter this accumulation induces the formation of 
large resin-blisters. Wlien, during the summer, the cambium is 
forming a new ring, the plethora of resin has the effect of causing 
the production of numerous resin-canals, which are unusually 
large and abnormally constructed, and these impart to the wood- 
ring formed during the }'ear of sickness a very striking and 
characteristic appearance. 

The mycelium gradualh- spreads from the cells of the 
medullary rays and from the resin-ducts into the vascular 
elements of the wood, where it produces a form of deca}- which 


may be termed a variety of white-rot. During the progress of 
the decomposition from the surface of the stem inwards a 
certain stage is reached, which is highly favourable to the 

Fig. 130. — Longitudinal section 
through the apex of a rhizomoiph 
from the outer hyphsc of which 
numerous hair-Hke filaments, a a, 
spring. In the interior the central 
cells enlarge greatly, /', at a short 
distance from the apex, while the 
cells of the hyphse situated to- 
wards the periphery, c, remain 
smaller, to form the pseudo-paren- 
chyma of the "rind" ; d d indi- 
cates the boundary of the mucila- 
ginous layer which envelops the 

Fig. 131. — Transverse section through the 
cortex and wood of a pine-root which has 
been killed by a rhizomorph. a, the dead 
tissues of the bast ; b, the dead cam- 
bium ; c, the medulla of the rhizomorph ; 
d d, the pseudo-parenchyma of the rind 
of the rhizomorph ; e e, filamentous 
hyphae which have grown from the 
rhizomorph into the wood ; /, dead im- 
mature wood-cells ; g, a resin-duct which 
has been completely destroyed, the paren- 
chymatous cells which surround it being 
also dissolved. 

development of the m}xelium. While previously it was simply 
filiform and furnished with numerous lateral hyphoe, it now 
develops large bladder-like swellings, and at the same time the 
hyphse change into a kind of large-meshed parenchyma, which, 
like the tyloses in the vessels of many dicot)^lcdonous trees, 

P 2 


completely fills up the lumina of the tracheides. On account 
of the mycelium assuming a brown colour when in this con- 
dition, it makes the portion of diseased wood which it infests 
appear to the naked eye like a black line. As this kind of 
mycelium soon dies off and is dissolved, being replaced by a 
delicate filamentous mycelium, it seldom happens that the zone 
which it occupies exceeds the breadth of 3 — 4 tracheids. The 
walls of the elements of the wood afterwards display a cellulose 
reaction, and speedily dissolve from the lumen outwards. 

On account of the trees drying up, after the rhizomorphs have 
spread from the point of infection on the roots into the stem, and 
again from the stem into the hitherto sound roots, decomposition of 
the stem usually ceases before the mycelium has advanced from the 
alburnum into the duramen. It is only in the stool and roots 
that decay rapidly spreads throughout the whole of the wood. 

The practical preventive measures to be enforced in the 
case of this parasite are the same as those which I have 
already recommended for Trainetes 7'adiciperda (see pp. 190-91). 


Although, strictly speaking, the diseases of felled timber 
should not be discussed in a te.Kt-book of the diseases of trees, 
still an abbreviated summary of the results of my investigations 
on this subject may not be altogether out of place.^ 

As regards the management of squared and round timber 
before it is utilised — that is to say, in the forest and during 
transport — one should in the first place take all reasonable 
precautions to see that, after felling, only sound wood is retained 
as structural timber. Of course it is always possible that now 
and again a log or beam will be retained that turns out to be 
diseased during subsequent manipulation. This may be due to 
the fact that a parasite which has entered through a branch- 
wound has not spread up or down to one of the sectional 
surfaces, so that it is impossible to recognize the destructive 
effects of the fungus when the timber is despatched. It is often 

' Der iichte Hausschwamui {Merulhes lacry7nans) (Berlin, Springer, 1885), 
and Die Rothsireifigkeit des Bate- und Blochholzes und die Trockeitfdule. 
Allg. Forst- und Jagd-Zeit., November 1887. 


the case, however, that the diseased portion of a tree which is 
easil}- recognized — as, for instance, OM'ing to brownness, &c. — is 
cut off till the saw-cut appears to the naked eye to be perfectly 
sound. The apparently sound portion of the tree is afterwards 
disposed of, say as a log. Now, it may easily happen that the 
parasite has already penetrated into the portion of the tree that 
was regarded as free from attack, and consequently an infected 
piece of timber is sold as sound. Should such wood retain a 
portion of its moisture for a considerable period, the parasite 
will continue to grow^ until it destroys not only the wood that 
contained the filamentous mycelium at the time the tree was 
felled, but frequently also very considerable portions of the 
timber that was primarily sound. 

Polyponis vaporarhis, which occurs on spruces and pines even 
when alive, and which I have described at page 198, is the 
commonest and most destructive of these fungi. Frequently 
when investigating the destructive effects of " dry-rot " I have 
found the cause to be not Meniliits lacryi)ians but P. vaporariiis, 
whose mycelium forms snow-white sheets on beams and deals, 
and produces stiff strands several yards in length. Should 
timber which is infested by this parasite be applied to structural 
purposes, and should it not dry quickly enough, the fungus 
develops more or less luxuriantly, and in a short time com- 
pletely destroys all the wood-work. This fungus is apt to be 
speciall)- prevalent in cellars, and in the wooden floors of the 
ground flat of houses that are unprovided with cellars. 

Perfectly sound timber may, however, also be infected during 
the time it is lying in the forest. The danger is greatest in 
the case of peeled timber that is in immediate contact with 
the ground. Various wood-fungi, and amongst them Merulhis 
lacrymans, may induce disease in felled timber when it is stored 
for a considerable time on the ground in the forest. At the time 
of issuing my publication on Merulius lacrymans, I stated that 
it was doubtful whether this fungus occurs in the forest at the 
present da}'. Since that time I have received genuine speci- 
mens of J/, lacrymans, from Herr W. Krieger, Konigstein, 
Saxony. Peeled timber that is exposed to air-currents by being 
piled upon supports is much better protected, because the 
surface la^-ers soon dry, and render the entrance of the fungus 


impossible. In the case of peeled stems that are freely exposed, 
drought in a few weeks induces the formation of cracks in the 
alburnum. These occur about an inch apart, and penetrate to a 
depth of an inch or more. The rain-water enters these cracks, 
carrying with it any spores that it may contain. After pro- 
longed rain the wood swells owing to the absorption of water, 
and the cracks close. During wet years, or long storage of 
the timber, decomposition may begin even in the forest, the 
spores that have entered by the cracks germinating and causing 
the wood to become brown along both sides of the fissure. 

As a rule, however, spores that enter cracks in the alburnum 
do not germinate in the forest, because when the rain ceases the 
superficial layers of the wood quickly dry again, so that even if the 
cracks should have closed they subsequently reopen. Should such 
wood be removed from the forest to the building or saw-mill in 
a dry condition, it remains sound, even although the spores in 
the cracks retain their power of germinating for a long time- 
If, on the other hand, the wood is floated, so that it has the 
opportunity to become again fully saturated with water, a very 
undesirable pathological symptom makes its appearance, which 
is known to saw-millers, timber-merchants, &c., as " the red 
stripe," and represents the first stage of what is popularly called 
„ dry-rot." 

It is a familiar fact that there is no essential difference as 
regards durability, or resistance to the attack of M. lacryuians 
and other wood-fungi, between coniferous timber that is felled in 
summer and that which is felled in winter. The attempt that 
has been made to show that the destructive effects of M. 
lacrymans are modified by the varying chemical composition 
(as regards potash, phosphoric acid, &c.) of wood felled in 
summer and in winter must be described as total failure^. On 
the other hand, it is an undoubted fact that wood which is felled 
in summer suffers far more from dry-rot than that which is felled 
in winter. This apparent contradiction is easily explained. 
Winter-felling takes place in the lowlands and in the less 
elevated mountains. In these districts the timber is chiefly 
removed from the forest by land, after it has lain for a longer or 
shorter period with or without its bark. Such timber is either 
free from spores, or, should it contain spores that have entered 


by cracks formed in the alburnum during drying, it afterwards 
remains dry, and therefore sound, because the spores are unable 
to germinate in the dry wood. On all the higher mountains, on 
the other hand, felling takes place in summer. The wood is at 
once peeled and piled on supports, and in winter it is conveyed 
on the snow to the streams, to be sent off in rafts in spring. 
The timber is dried in the first summer — that is to say, directly 
after being felled and peeled — when cracks form through which 
the spores of fungi enter. During floating the logs become 
saturated with water, and the cracks close. When the wet logs 
arrive at the saw-mills they are piled up in thousands, to be 
sawn up in the course of the summer. The logs that are sawn 
up in May are, as a rule, perfectly sound, but from June onwards 
the number of " red-stripeci " specimens constantly increases, 
until in autumn it frequently happens that more than 50 per 
cent, of the logs are so decayed as to furnish but few serviceable 
boards. This is easily explained, if one considers that the 
saturated logs are prevented from drying owing to the way 
they are piled up on one another, and that the high summer 
temperature is suitable for the germination of the spores present 
in the cracks, and favours the destructive development of the 

The owners of saw-mills in the Bavarian Forest calculate that 
they lose 33 per cent, of their total timber by logs becoming 
red-striped. For some years I conducted extensive investiga- 
tions not only at Zwiesel in the Bavarian Forest, but also at 
Marquardstein and Freising, partly to determine the cause of 
timber becoming red-striped, and partly with the object of dis- 
covering a means of preventing the mischief. This is not the 
place to go into the details of these arduous investigations. 
I have shortly described the causes of the phenomenon above. 
As regards the prevention of the disease, it was found to be 
possible to obtain perfectly sound logs by protecting them 
against rain by a covering of boards or spruce-bark. Unfor- 
tunately this only induces another evil — namely, the excessive 
cracking of the timber, which means a very serious shortage in 
good boards. The rejected red-striped boards are used in 
houses for underflooring and for false floors. As it very often 
happens that the wood has not been sufficiently dried to kill the 


enclosed fungus-mjxelium, the latter continues to grow in the 
presence of moisture, and the wood is still further destroyed. 

Squared timber that has been floated suffers quite as much 
from red-stripe as that which comes straight from the saw-mill. 
As nowadays it hardly ever happens that perfectly dry timber 
is employed for structural purposes, there is great danger of the 
so-called " dry-rot " appearing in a destructive form. 

The greatest danger attaches to the ends of joists that are 
built into a wall. If the latter contains water, it is transmitted 
to the wood, so that joists which may have been fairly dry are 
again rendered so wet as to enable any fungus-mycelium con- 
tained in the cracks of the wood to develop and destroy timber 
that was perfectly sound when placed in the building. Should 
the ends of the joists have originally shown any appearance of 
red-stripe, the danger of total decay is of course increased. One 
ought therefore to endeavour, as far as possible, to avoid using 
red-striped joists, or at least their use should be confined to the 
highest story of a building, where the walls being thinner dry 
faster. Under any circumstances, however, one should never 
neglect to apply several coats of creosote (common coal-tar oil) 
or some special carbolic preparation to the ends of the joists for 
a distance of three feet, before they are built into the wall. Tar 
cannot be recommended, because it does not penetrate far into 
the wood, and it forms a covering which prevents the joists 
from drying. 

The other parts of the joists are not so much exposed to 
danger. Even when they are red-striped they usually dry so 
soon in properly constructed buildings as not to suffer further 
damage from any fungus that they ma}' contain, though of course 
their strength is reduced in proportion to the extent of the 

The name " dr}--rot " is unhappily chosen, in so far that it is 
characterized as occurring only in wet or damp wood, in which 
the fungi can find sufficient moisture for growth. Merulhis 
laciyviaiis, on the other hand, may destroy perfectly dry wood 
by imbibing and conducting the water requisite for growth from 
other parts of the building, and either parting with it to the 
woodwork or letting it escape in the form of drops or " tears." 
The disease has, in fact, acquired the name " dr}'-rot " because it 



is usuall\- only noticed in a building- when it, and consequently 
the woodwork also, has become practically dry. 

Frequently, however, dry-rot appears in new buildings to such 
an extent that not only the joists but also the boards of the 
false and true floors decay. When this is the case the cause 
is usually to be found in gross negligence on the part of the 
contractor. Most frequently the mistake is committed of placing 
wet deadening material (" pugging ") on the false floor and 
covering it over too soon, either with the boards of the sub-floor 
or of the true floor. I have thoroughly discussed the subject of 
deadening ma-terial in my work on M. lacryiiians. It must be 
as dry as possible, and free from humus or an)'thing that will 
condense moisture. Clean gravel or coarse dry sand suits best. 
Anj'thing of the nature of coal-dust should on no account be 

It is a great mistake to cover the floor too soon with oil 
paint or with parquet, because this prevents the evaporation of 
any moisture that may have been originally present in the 
boards, or that may have been imparted to them by the packing 
material. The water that is contained in the packing material 
and in the woodwork cannot afterwards escape upwards. All 
that is possible is an extremely slow evaporation downwards — 
that is to say, through the ceiling of the room beneath. Between 
the false ceiling and the matchboard ceiling the air becomes 
saturated with moisture, and this space offers conditions which 
are extremely favourable for the growth of fungi. The flooring 
boards, being saturated with moisture derived from the packing 
material, decompose under the action of the spores which are 
brought from the forest in the cracks of the wood. In two years' 
time, when the building has become perfectly dry, the moisture 
in the boards also disappears. The withdrawal of water induces 
very great shrinkage in the already decomposed w^ood of the 
lower side of the boards, while the upper side, being exposed to 
the air or protected by paint or varnish, is not similarly affected. 
The result is that the upper side of each board becomes convex 
in the middle, and the nails are easily wrenched out of the 
partiall}- rotten joists. Open joints are thus formed, which may 
be large enough to admit of the entrance of one's finger. -^ 

The repairs thus rendered necessar}' are very expensive, and 


give rise to vexatious litigation between the architect, builder, 
carpenter, and timber-merchant. Nor are the distinctions be- 
tween this form of dry-rot and that induced by Mentlius 
laaynians sufficiently appreciated as a rule, although the ravages 
of the latter may be easil)' recognized since the publication of 
my work on the subject. 

As a rule the term " dry-rot " is applied to those forms of 
decomposition in structural timber where the fungus that does 
the damage is invisible to the naked eye. This want of 
conspicuousness is accounted for by the fact that such fungi, 
instead of covering the wood or of filling up cracks in the timber 
or spaces between the woodwork and the walls with mycelial 
growths, distribute their fine hyphae in the substance of the wood 
itself. But in a series of fungi which destroy structural timber a 
luxuriant mycelial growth is produced outside of the wood, and 
it is to these that the term " House Fungus" is generally applied. 
These fungi vary exceedingly as regards appearance and life- 
history. Of them the most important and destructive is Menilhis 
lacryinans. Then we have also Polyporus vaporarius, which has 
already been described, and a number of other fungi which I am 
at present busily engaged in investigating. 

Space may also be found here for a few remarks on the 
soundness and quality of timber furnished by conifers that have 
been entirely defoliated by caterpillars, and especially by 
Liparis inonacJia and Gastropacha pini. When spruces or pines 
have been completely defoliated during spring or summer, the 
leafless branches as well as the top of the tree die in the course 
of the following autumn, winter, or spring, while the more 
valuable parts of the stem remain perfectly sound till the middle 
of the succeeding summer. As a rule it is not till the beginning 
of July that the inner cortex, especially on the south-west side of 
the ti'ee, begins to show brown patches and die. During the 
devastations committed by the nun moth in recent years, the older 
classes of spruces were almost all dead by autumn — that is to say, 
the cortex was brown. Underneath the dead cortex the wood 
of the alburnum became discoloured, and rapidly decomposed 
under the influence of numerous fungi. The timber of all those 
trees which were felled and immediately barked before the 
beginning of July of the year in which the havoc was committed 


was found to be of exceptionally high qualit)', and showed 
no blemishes. This was the case even after the barked trees 
had been piled for a whole year before being removed from the 
forest. Owing to the stores of carbo-hydrates having been 
nearly all used up by the cambium in the formation of the wood- 
ring during the summer of the first year, such timber offered less 
suitable conditions for the growth of fungi than the wood of 
trees that had not been defoliated. It was only after the 
cortex had died, or had been perforated by wood and bark 
beetles, that fungi could gain an entrance, when the high 
temperature and abundance of moisture offered favourable 
conditions for their growth. The low repute in which timber 
furnished by trees destroyed by insects is held is entirely due 
to the fact that such trees are often left standing in the forest 
till the cortex is dead. Trees that have been stripped of their 
leaves should therefore be felled and barked not later than the 
beginning of July of the year succeeding the defoliation. 

I now return to the consideration of the true dry-rot fungus, 
Meriilhis lacryinmis. 

Although this plant has been encountered at least once on the 
old stool of a conifer in the open forest, it is usually associated 
with man. It is probable, however, that it has hitherto escaped 
notice in plantations, and that it is more generally distributed 
than is usually supposed. Although it lives chiefly on coniferous 
timber, it also grows on oak, and the oaken boards of parquet 
floors are liable to be infected. 

The filamentous mycelium which is invisible to the naked eye 
grows inside the wood, from which it abstracts the proteids 
necessary for its growth. At the same time it dissolves 
the coniferin and cellulose of the cell-walls, and leaves behind 
a brown residue consisting of lignin, tannin, and oxalate of lime. 
So long as sufficient moisture is present these substances 
enable the wood to retain its original volume, but whenever 
water is withdrawn the wood becomes traversed by numerous 
fissures running at right angles to each other, and frequently 
breaks up into regular cubes. 

As the wood decomposes it becomes brown in colour, a result 
which is probably due to the higher oxidation of the tannin. 
Although soft when damp, the wood bears some resemblance to 


charcoal when di'}-, and may be rubbed down between the fingers 
into an impalpable yellow powder. An important property which 
it possesses is its great sponge-like power of absorbing water. 
This is chiefly due to the fact that, owing to the cell-walls having 
been perforated by the filamentous mycelia, the air is enabled to 
escape in front of the water which enters by capillarity. Thus 
it happens that when a house is attacked by M. lacryvians 
the woodwork is able to absorb water with great ease, and to 
transport it to considerable distances. Thus the capillarit}' of 
the diseased wood makes it possible for liquid water to be 
conveyed from the ground floor of a house to the upper stories, 
which it may render damp by evaporation. So far wood that is 
decomposed by M. lacrynians resembles that which is attacked 
by what is popularly called dry-rot. 

il/. lacrymans is, however, capable of growing out of the wood 
in which it feeds, if onl}- the surrounding air remains sufficiently 
humid to prevent the advancing m}'celial filaments drying 
up. Where, therefore, the air is stagnant and humid, the 
mycelium grows out of the wood, at first taking the form of a 
snow-white loose woolly growth, which spreads over the wood 
and covers its surface. These white fungus-growths also spread 
on to other objects from which they can obtain no nutriment, 
provided they are situated in the neighbourhood of the wood- 
work. Thus they creep up the walls, and spread over the 
damp ground, flag-stones, &c. Later on stouter branching 
strands of the same colour occur amongst the masses of floccose 
fungoid hyphae. These may attain to the thickness of the 
finger, and are of immense importance in the life-histor}' of 
M. lacrynians. 

Before proceeding to describe these stout strands, I may 
mention that the woolly mass of mycelia attains more consistency 
as it gets older, and forms a lustrous silk}- ash-coloured sheet which 
may be detached from the substratum. The ash}- grey colour 
of this mycelium enables us to distinguish it from that of 
P. vaporariiis, already described, which always remains white. 
/ The mycelial strands of M. /aery mans consist of(i) firm fibres, 

which make them to a certain extent untearable, (2) filaments rich 
in protoplasm, which in humid air may send out buds in all 
directions, and (3) organs resembling vessels with large lumina, 


which contain a plentiful supply of proteid substances. Not 
only water but also large quantities of nutritive substances are 
apparently conveyed in these vessel-like organs from the nutrient 
substratum — that is to say, the woodwork — to the more remote 
portions of the growing mycelium. Now, as these strands attain 
to a length of many yards, and by taking advantage of depres- 
sions in walls mount from the cellar to the ground floor, and 
from there to the upper stories, it is easily seen that the fungus 
may occur in parts of a building in which there is absolutely no 
woodwork, without having encountered any nourishment — that 
is to say, wood — on its way. Of course those strands do not 
advance as such. It is the delicate filamentous mycelium which, 
supplied with water and nourishment from the strands behind, 
and taking advantage of every crack and cranny, growls through 
walls, soil, &c. The chink in a wall which was entered at first 
by a delicate floccose mycelium later on contains a thick strand, 
which, however, has gradually developed from the former. 
Should the mycelium during its progress again gain access to 
woodwork, it destroys the latter, the delicate filaments entering 
and abstracting the nourishment, and thus gaining strength for 
more vigorous development. It is characteristic of AI. lacry- 
Dians that it is able to destroy even dry woodwork. This is 
rendered possible by the strands conducting enough water from 
other damp parts of the building to soak the dry wood, and 
thus make it suitable for attack. In muggy rooms, when wood 
is not available to absorb the water, the fungus parts with it 
in the form of drops or " tears," which has gained for it the 
name of the " weeping " house-fungus {lacrymans). 

If sufficient space be available, and as a rule in the presence 
of more or less light, though this is not absolutely necessary, the 
familiar sporophores are formed. Though they vary in form, 
these are usually of a flat saucer-like shape. The fungus-mass, 
which is at first white and loose, assumes a reddish colour in 
places, and displays vermiform folds, which soon become so 
covered with rusty spores that the whole surface is coloured 
deep-brown. The brown spores, which are so small that about 
sixty-five thousand millions can be contained in a cubic inch of 
space, display a germ-aperture in the thick wall at one end, 
which is closed however by a clear lustrous plug. 


The spores of M. lacrynians can germinate only when this 
plug has been dissolved or has disappeared in some way, and 
this seems to occur only under the action of some alkali. I 
succeeded with germination experiments only when I had added 
some ammonia or salts of potash or soda to the infusion in 
which the spores were placed. These salts are not to be 
regarded as nutritive in their effects, but merely as rendering 
possible the removal of the spore-pellicle that covers the germ- 
aperture. Every seed and every spore contains a certain quan- 
tit}' of nourishment which has been derived from the parent 
plant, and which is instantly available for use. Only when this 
has been used up during germination is further development 
dependent upon a supply of nourishment from the environment. 
I will not contest the possibility that now and then a spore 
of J/, lacrymans may germinate directly on wood, which 
of course contains minute traces of alkalis, still I have only 
succeeded in inducing spores to germinate on wood by adding a 
little alkali. This explains why injuries from M. lacrymans are 
specially apt to occur in places where urine, humus, wood ashes, 
coal-dust, and such like are present. 

Wood is the natural food of M. lacrymans, and in this respect 
there is no difference between summer-felled and winter-felled 
timber. The causes of the frequent complaints regarding 
summer-felled wood have already been discussed. 

Soil that is very rich in humus also offers nourishment to M. 
lacrymans, though only in small quantity. It is probable, 
although not certain, that when the mycelium is growing in 
contact with walls it dissolves and consumes minute quantities 
of lime, but in any case these are so small that no direct 
damage can be ascribed to this cause. 

When alive or still fresh, M. lacrymans has a very pleasant 
odour and delicate flavour, though this is succeeded by a some- 
what astringent taste. When sporophores, especially large ones, 
decompose, they disseminate a highly repugnant and very 
characteristic smell. There is no doubt that the gases generated 
by the decaying fungus are highly injurious to the health of 
human beings inhabiting rooms exposed to them. In addition 
to this, large quantities of water are evaporated from the fungus, 
and thus rooms are kept damp. 


Even under the most favourable circumstances, M. lacryinans 
can only appear after infection by spores or pieces of mycelia, 
and on this account it is important to determine how the spores 
or mycelia are distributed and carried about. 

I have already mentioned above that, under certain circum- 
stances, the spores may be brought with the timber from the 
forest. Such cases, however, must be extremely rare, at least 
under the conditions of forest conservanc}' that obtain in 
Germany, where large quantities of timber are seldom 
stored in the forest to admit of the development of M. 
lacrymaiis, which has hitherto been observed but once in such 
a situation. That timber may be infected and attacked by 
AT. laaynians during long storage in the forest naturally follows 
from what has been said. But as a rule infection occurs only in 
the towns, either in the wood-yards of carpenters, cabinet-makers, 
&c., or in houses. It happens often enough in wood-yards that 
the timber of old houses, which is still useful for certain 
purposes, is stored beside sound wood, so that the rain washes 
any loose spores and bits of mycelium on to the sound wood. 
Workmen, especially carpenters — who, let us say, have been 
executing repairs in a structure affected by ]\I. lacrymans — 
easily introduce the "spores into new buildings, by proceeding 
from the one to the other without changing or cleaning their 
clothes, boots, or tools. 

For M. lacrymans to appear it is not merel}' necessary that 
spores or mycelia should be present, but the conditions necessary 
for their development must also be favourable. The spores 
germinate onl}- in the presence of alkalis. This explains the 
disastrous consequences of emplo}'ing humus-substances or 
wood or coal ashes as packing materials, or allowing the work- 
men to pollute the building with urine. The further growth 
and vigorous development of the fungus are, however, most 
encouraged by the use of damp materials, e.g. damp wood, 
damp packing, damp stones, &c., because moisture is neces- 
sary for the gro\\-th of M. lacrymans as well as ever}- other 

This is no more the place to go into further details regarding 
preventive measures to be taken in building a house than it is 
to describe the measures to be instituted when J/, lacrymans 


appears in a structure. In the book which I have quoted I have 
thoroughly discussed all these matters. 

Amongst the saprophytic wood-fungi, Peziza centginosa 
excites a general interest. Although belonging to the Dis- 
coinycetes, it may be mentioned in this place, as it is to it 
that the so-called " green-rot " of wood is due. When much- 
decayed wood, of the oak and beech especially, less frequently 
of the spruce and birch, lies constantly soaked on the ground 
of the forest for a long time, it frequently assumes an intense 
verdigris-green colour. This is due to the wood being occupied 
by the mycelium of the above-named fungus, which, along with 
the saucer-shaped sporophore, is vividly green in colour. The 
green pigment, which may be extracted, is also present in the 
walls of the elements of the wood. 

On account of its indestructibility the green colouring matter 
finds employment in the arts, and recently experiments have 
been instituted to produce green-rot in wood on a large scale by 
artificial propagation. 

The so-called " blueness " of coniferous wood is due to a 
Pyrenomycete, Ceratostoma piliferiini {^Sph(zria diyina), whose 
brown mycelium enters the stem by the medullary rays, and very 
rapidly reaches the pith. It is specially common in pine woods 
on unhealthy trees, such as those which have suffered from 
caterpillars, or it may appear in a heap of damp fagots. 
Probably on account of deficiency of moisture it rather avoids 
the duramen, whereas the alburnum is often quickly occupied by 
the mycelium, and destroyed. 



Numerous wounds are produced annually in plants which are 
the result of normal biological processes. Thus leaves are shed 
in autumn, certain twigs are naturally cast off {e.g. in poplars 
and oaks), and the outer layers of the cortex die. The plant 
makes preparation some time in advance for all such wounds 
as occur naturally, so that at the moment when the wound is 
formed the process of healing may be regarded as completed. 
This preparation consists in a periderm being formed in the 
tissues along the plane which the surface of the wound 
ultimately occupies. In its origin and structure this periderm 
entirely agrees with the periderm of uninjured shoots, or with 
the peridermal covering that gradually forms on wounds which 
have resulted from an accident. In many cases a protective 
covering of gum is first spread over the wound, and later on the 
formation of a periderm is gradually accomplished. Only such 
wounds as are due to external mechanical causes, which have 
exposed the internal living tissues to the prejudicial influences 
of the environment, come into the category of pathological 


In order to understand the processes of healing and the 
production of new tissues, we must first cast a glance at the 
different kinds of tissues and their capacity to produce new 

On the young parts of plants the protective covering is 
represented solely by the epidermis, which usually consists of 
a single cell-layer. But before this has entirely lost its power 


of expanding, and has been ruptured by the growth in thickness 
of the stem, a new protective covering is formed beneath it, 
which protects the inner hving cortical tissues against drought. 
This periderm — on whose structure and characteristics it would 
be out of place here to enlarge — is formed from a layer of 
phellogen (cork-cambium), which results from the tangential 
division either of the epidermal cells w^iile still alive, or of a 
layer of cortical cells which is situated at a greater or less 
distance beneath the epidermis. The radially arranged cells, 
which are being constantly formed by division, die and become 
converted into cork, and thus a protecting envelope, more or less 
thick, is formed on the outside of the living tissues. By division 
of the phellogen-layer the envelope is constantly being renewed 
on its inner surface, whereas the oldest cork-cells on the outside 
are being lost by the exfoliation or detachment of compact 
layers of cork-cells. In the case of most trees bark is formed 
sooner or later, owing to the older layers of the cortex and bast 
losing their power of expansion.* When this occurs new cork- 
layers form in the interior of the cortex, and these separate the 
inner layers from the outer layers of cortex, immediately before 
the latter die, dry up, and rupture. 

It is evident that an injury to the dead periderm or bark 
is unaccompanied by any prejudicial results. The only way in 
which it can affect the growth of the tree is that by diminishing 
the pressure it stimulates the cambium to increased activity. 
Where the dead bark has been mostly removed in a broad 
zone from pines, for the purpose of laying on a ring of tar 
with the object of intercepting caterpillars, the trees during suc- 
ceeding years grow distinctly faster at the barked region than 
either above or below. In the event of the layer of living 
phellogen being injured, a new zone of phellogen and cork, 
which is continuous with the cork layer along the edge of 
the wound, is formed from the uninjured cells which are situated 
deeper in the cortex or phelloderm. 

The cortical parenchyma (Fig. 132, b c) which lies beneath the 

*[" Bark " is, therefore, all the dead tissue situated outside the phellogen: 
it may be represented by the corky layer of the periderm only, or may include 
this and dead tissues of the cortex, which the periderm has cut out as well. 



periderm possesses sufficient power of cell-division to enable it 
to keep pace with the increasing thickness of the stem. In the 
case of a wound, however, its reproductive capacity is confined 
to the development of a periderm close beneath the surface 
of the exposed tissues. This layer of cork, which is also formed 
along the boundary between the sound and dead tissues when 
plant-parasites induce diseases of the cortex, is called "Wound 

Fig. 132. — The formation of callus on the edge of a wound on an oak-branch, a 
periderm ; b, collenchyma ; c, outer cortex ; d, primary bundles of hard bast 
e, cortical parenchyma ; f, soft bast ; g, cambium ; h, wood ; /, " wound-cork ' 
formed by the outer cortex ; k, callus. 

Cork" (Fig. 132, i). Its formation does not depend on the 
season of the }^ear, for even in winter, should the weather be 
favourable, it is formed soon after the occurrence of a wound. 

Only that portion of the cortical parenchyma which is situated 
nearest to the cambium, or the soft bast, or in other cases merel}' 
the deepest-h'ing and }-oungest organs of the soft bast, take part 
in the reproductive processes that are about to be discussed. 

As wood consists for the most part of empty elements— viz. 
fibres, tracheids, and vessels — it possesses only a very limited re- 
productive capacity. The cells of the wood that retain vitalit}- 
consist of the parench)^ma of the medullary ra)'s and the wood- 

Q 2 



parenchyma, but these are so surrounded by the elements 
above mentioned that they are scarcely able to exercise even 
the limited reproductive capacity which they do possess. This 
capacity is exhibited in but two forms — first, in the production 
of tyloses or " filling cells " in the vessels of the wood whenever 
these are injured, and, secondly, in the development of so-called 
" intermediary '' tissue (" cementing tissue ") during the process 
of engrafting.^ When the cut surfaces of the scion and stock are 
bound together in a sufficiently fresh condition, any empty space 
which may exist between the two portions of wood becomes 
filled with parenchymatous tissue, which 
originates in the above-mentioned paren- 
chymatous cells of the wood itself. 

Wood that is exposed by a wound has 
the power of producing cortex and wood 
only if the cortex is removed during the 
season when the cambium is active, and 
the cambium layer or the young wood 
is protected against drought. In such a 
case the regeneration of the covering 
layers is effected. The region of the 
cambium, with its delicate cells and 
abundant protoplasm, consists, during the 
period from May to August, of initial 
cells, mother-cells that have been formed 
from these by division, and young em- 
bryonic cellular tissue (young bast and 
young wood) which is still capable of 
growth. When exposed to the air this region dries up very 
easily, and only during rainy weather, or when the air is saturated 
with moisture, does this tissue survive, and, by the transverse 
division of the elongated elements of the cambium, become 
converted into a healing tissue consisting of parenchymatous 
iso-diametric cells. 

Owing to energetic cell-division this gives rise in a few days 

to an investing layer (Fig. 133), which, under the influence of 

light, assumes a green colour. Frequently the cambium that 

covers the surface of a wound withers, with the exception of the 

^ Gcippert, Ueber iimere Vorgimge bei detii Veredcln, Cassel, 1874. 

Fig. 133. — Surface of a 
beech-stem from which 
the cortex has been re- 
moved, and on which 
an investing layer has 
been partially formed. 
Natural size. 



cambium of the medullar}- raj-s, so that the clothing of the surface 
of the wound is almost exclusively undertaken by the latter, 
giving the impression that the medullary ra}'s have grown out of 
the wood. The healing tissue, which is originally homogeneous, 
soon shows a certain amount of differentiation. The elements 
which abut upon the old wood change into wood-cells, while 
towards the outside a new bast region forms amongst the layers 
of cells that are assuming the form of parenchymatous cortical 

77 m 7 ''■'«' 

Fig. 134. — Cross-section of the stem of an oak which, two years before being felled, 
had ruptured at several places in the cortex in consequence of much-augmented 
growth. X and ;', two places where the cortex had ruptured ; a \.o b, new in- 
vesting layers formed by occlusion with their cortex, d ; c, callus ; e to e, 
lower surface of the loosened cortex, the cambium of which has also produced 
new growth. 

tissue. A portion of tissue between the wood and bast preserves 
the character of meristematic cambium, while a new epidermis 
forms on the surface of the cortex. 

In the accompanying woodcut (Fig. 134), which represents the 
cross-section of an oak whose bark became separated from 
the stem two years before felling, the portion of the surface 
of the wound situated between b and b has dried up. Beneath 
the shelter of the loosened cortex, e e, on both sides of the wound 
new healing tissues {a b) have been formed on the wood, and 
these have alread\' attained an age of two )-ears (1876-77). 


Should the loosened cortex be supplied with nourishment by 
remaining in organic union with the tree, new tissue may of 
course also be formed on its under surface, to which some 
cambium will also have adhered. In such a case the process of 
cell-division proceeds normally in the cambium, after it has been 
converted as explained above into short-celled cambium. It is 
in this way that the new tissues have been formed during the 
two years which have succeeded the loosening of the flaps of 
bark, e e (Fig. 134). 

The wood which is formed on the surface of the exposed wood 
of the stem and on the inner surface of the detached bast is 
distinguished from ordinary wood by its abnormal structure, 
and especially by the shortness of its cells and the absence or 
scarcity of vessels. H. de Vries/ who was the first to direct 
attention to this abnormality, designated such wood with the 
name " Wound-Wood." 

The formation of new cortex in the manner above de- 
scribed has been made use of on a large scale in the cultiva- 
tion of cinchona bark under Maclvor's system. Strips of 
cortex several yards long are separated from the wood along 
the cambium zone, alternate strips of the same breadth being 
left in situ. The whole is then covered with moss. The 
system can only be practised during the rainy season. The fresh 
growth contains twice as much quinine as the original bark.* 

When the cambium on the portion of a stem that has been 
deprived of its cortex dries up before it can produce an invest- 
ing layer, or should cambium be entirely absent from the sur- 
face of a wound, as, for instance, in the case of branch-wounds, 
&c., the only regenerative process that is possible is the 
formation of callus from the edge of the wound. 

Under the so-called Javanese method the cortex is removed, 
with the exception of a thin layer which contains the cambium 
and youngest bast. In a short time a layer of periderm forms 
beneath the surface, and prevents the loss of moisture. By this 
method, which may be practised at all seasons, the tree does not 
require to be bound round with moss. 

1 Hugo de Vries, Ueber Wundholz , Flora, 1876. 

* [This system of "niossing" has been much in vogue among the planters 
in Ceylon. — Ed.] 


The process by which callus is formed begins in the soft bast, 
and in the embryonic tissue along the edge of the wound — 
namely, the cambium (Fig. 132, £-). It is a purely mechanical 
process, and results from the bark-pressure on these tissues 
being reduced. The annual growth in thickness of the stem 
produces distension of the cortex and bast, which, however, 
is balanced for the most part by the living cells of these tissues 
dividing and growing, and so keeping pace with the increase in 
the periphery of the stem, while the dead external portions 
become fissured longitudinally. Nevertheless there is always 
a cer:ain amount of tension in the cortical mantle, whereby a 
considerable pressure is exerted on the cambium. Should this 
pressure on the cambium be locally reduced by a wound 
reaching to the wood, the processes of cell-division and 
growth are accelerated not only along the edges of the wound 
but also at greater distances. In Fig. 132 this is visible as far as 
^. Wherever the pressure has been reduced (in Fig. 1 34 this 
may be perceived at a distance of some inches from the points 
a a), the normal cambium changes into " wound-cambium " 
with short cells, which produces a luxuriant growth of " wound- 
wood," destitute of vessels and without distinct medullary rays. 
The process of cell-division proceeds most energeticall}- in the 
direction of the surface of the wound, where of course there is 
absolutely no counter-pressure, and one may perceive the 
cushion-like callus appearing between the wood and the cortex 
Either in the year in which it originated, or not till later, the 
wound-wood assumes a normal character, whereas the cortex of 
the callus remains thinner and more expansive for a series of 
years, and exerts less pressure than old cortex or bark. The 
increased rate of growth is consequently not confined to the 
first year, but is often maintained till the various callus-cushions 
which advance from the edges of the wound come into contact 
and coalesce. 

This coalescence is retarded, if not rendered absolutely 
impossible, in the case of trees which at an early stage clothe 
the callus with dead bark. 

Should the cortex of callus-growths that have come 
into contact be thin, living, and free from dead bark, it is 
squeezed out during further growth, so that cambium abuts upon 


cambium, and complete coalescence results.* Thick bark may 
retard this coalescence for many decades, as, for instance, in the 
case of the pine (Fig. 138). 

When one considers that the pressure exerted by the bark 
in consequence of the peripheral enlargement of the stem acts 
for the most part horizontally, like the pressure of a barrel- 
hoop upon the staves, it is evident that the formation of callus 
must proceed much more vigorously in the case of a longi- 
tudinal incision in the cortex than when the incision is a 
transverse one. This sufficiently explains why callus is most 
vigorously produced along the lateral margins of branch-wounds. 

Should an injury produce little or no reduction in the bark- 
pressure, as in the case of bruises caused, for instance, by one 
tree knocking against another during felling, the formation of 
callus is either absolutely prevented or proceeds with great 
slowness. The dead cortex, which, without becoming detached 
from the uninjured portion, retains its position on the bruised 
and lifeless spot, does not admit of a reduction of pressure along 
the edge of the wound, and consequently no formation of callus 
takes place. 

Finally, it may be mentioned that the shape of the wound 
may be recognized on the surface of the tree for many decades, 
the boundary between the old and new cortex being usually 
visible for a long time. 

It need hardly be mentioned that coalescence of the wood 
exposed by a wound, with the wood of the callus that is 
subsequently formed over it, is impossible, and especially so as 
the external wood-layers of the wound have previously died, 
dried up, and become decomposed to a greater or less depth. 

This leads us to the consideration of the changes that occur 
in wood which is exposed by a wound. In the case of those 
conifers which are furnished with resin-ducts, the surface of the 
wound is more or less perfectly protected, owing to the outer 
layers of wood becoming impregnated with resin. 

The resin-ducts, into which resin mixed with turpentine 
is shed from the surrounding parenchymatous cells which 

* [I have proposed to call all such cases of covering over of wounded sur- 
faces by the agency of a callus, "occlusion": the wound is said to be 
" occluded.'' — Ed.] 



produce it (resiniferous cells), are disposed in the wood both 
vertically and horizontally — that is, radiall}-. I was the first to 
show that the latter, 
which are known as 
medullar}'-ray canals, 
communicate freely at 
certain points with the 
vertical canals. This 
is owing to the fact 
that at those places 
where the two sets of 
canals come into con- 
tact the parenchyma- 
tous epithelial cells, 
instead of remaining 
coherent to each other, 
become widely sepa- 
rated (Fig. 135, 4 

By means of these 
intercellular spaces 
the resin of the ver- 
tical canals can with 
ease gain access to 
ihe radial canals, and 
should the latter be 
opened by a wound 
on the outside of the 
tree the resin is en- 
abled to flow freel)' 
out to the surface. 
This explains the 
abundant outpouring 
which takes place 
when conifers are par- 
tially barked in order 
to procure the resin. 

Under the oxidising influence of the air the resin that oozes from 
the wounded surface soon forms a hard incrustation ; and of 
course the partial volatilization of the turpentine also contri- 

FlG. 135. — Manner of communication between a ver- 
tical resin-duct, a, and a duct in a medullary ray, 
/', in the Norway spruce. The epithelial cells of 
both canals are for the most part empty and 
furnished with very thick walls ; the walls be- 
tween adjoining epithelial cells being abundantly 
pitted, ci. Only a small ]iroportion of these cells 
retain thin walls, protoplasm, and a nucleus, and 
seive for the preparation of turpentine, dd. At 
the point where the back of the vertical canal 
facing the reader, a, comes into contact with the 
horizontal canal behind, /', the epithelial cells of 
both canals are provided with very delicate walls, 
and are separated by large intercellular spaces, 
e e, the latter providing the means for the passage 
of the turpentine from the one canal to the other. 


butes to the induration of the exposed mixture of resin and 

If a conifer be felled or a branch removed, either during 
summer or winter, one very soon perceives an exudation of resin 
from the alburnum (" sap-wood ") of the cut surface. But in 
the case of the pine, spruce, and larch no resin exudes from the 
older parts of the wood, although these parts are frequently 
more resinous than the alburnum. I believe that this state of 
things may be easily explained by the fact that not only are 
the cell-walls of the alburnum completely saturated with water, 
but the lumina of the tracheids are more than half full of water. 
In spite of its volatility the turpentine contained in the resin- 
ducts is unable to distribute itself throughout the wet wood, 
and in the case of a wound is forced out of the canals. When 
the wood, with advancing age, loses its power of conducting 
water, and so becomes drier — no matter whether this is accom- 
panied by the formation of duramen (" heart-wood ") or not — 
there is nothing to prevent the turpentine spreading throughout 
the wood. Not only does it spread into the cell-walls and 
impregnate them with resin, but it is also deposited in the form 
of drops on the walls in the lumina of the tracheids, and in 
fact the lumina are not unfrequently^-completely filled with tur- 
pentine or resin. In this way old pine-wood is frequently so 
saturated with resin that sections as thick as one's finger become 
partially transparent. Should a .section be made of old wood 
that can no longer conduct water, there will be no exudation 
of turpentine, for the reason that it has become a part of the 
walls of the tracheids, or has been deposited in their lumina. 

This also explains why the alburnum becomes completely 
impregnated with resin when, in consequence of a wound, its 
outer layers are exposed and dry up. The water that is lost 
by evaporation is at once replaced by turpentine, which is 
conveyed in abundance from other parts by means of the resin- 
canals. The resinous impregnation of these outer layers forms 
a protection against further injury from the environment. 

The resinous saturation of the old stools of conifers, and the 
distribution of the turpentine in trees whose wood is being 
decomposed by parasitic fungi, are very peculiar. The 
turpentine moves from the decomposed parts to the boundary 


between the sound and diseased wood. One is inclined 
to assume that when the cell-walls arc destroyed by the 
m}xelium of the fungus the turpentine in the interstices of the 
micellee is again liberated and becomes volatile, and so 
penetrates such cell-walls as are either wholly or for the most 
part free from decomposition. As a matter of fact, those parts 
of the wood which are the last to be attacked by the parasite 
become completely saturated with resin, whereas mere traces of 
resin are to be found in the decomposed portions. Thus, when 
the alburnum has been destroyed, the duramen of old pine-stools 
is very resinous. So far there is no proof to support the view 
that the cell-walls are converted into resin during the decom- 
position of the wood. 

When wounds due to pruning, barking, &c., expose the wood 
of a dicotyledonous tree, the tree protects itself against the 
unfavourable influences of the environment in two ways. In 
the first place, the vessels become completely plugged up 
by tyloses,* which both prevents the entrance of rain-water 
and the evaporation of any water that may be present in 
these organs. In the second place, gums are formed in abun- 
dance in the neighbourhood of the wounded surface, and these 
fill up and close the lumina of the organs, especially the vessels, 
thereby protecting them to a certain extent against the 
prejudicial influences of the environment. It is probably to 
the direct action of the oxygen of the air that the brownness 
of the wood under the surface of the wound is due, tannin and 
its allies especially assuming a brown colour in the higher 
stages of oxidation. 

The foregoing protective agencies are, however, insufficient to 
afford absolute security to the exposed wood against decompo- 
sition and decay. On this account wound-diseases are much 
more liable to occur in dicotyledonous trees than in the resinous 

In the previous section attention has already been directed to 
wound-diseases due to parasites, and I shall again refer to this 
subject when dealing with the pruning of trees. But besides 

* [Tyloses are ingrowths of the cells surrounding a vessel, which push 
their way through the bordered pits into the cavity, and may there divide and 
grow further. — Ed.] 


the forms of decay in wounds which are induced b}' parasites, 
there are other forms of decomposition in wood in which parasitic 
fungi take no part. It is rather to the saprophytic fungi, in 
conjunction with atmospheric influences, that a variety of forms 
of decay in wood are to be ascribed. In the meantime I propose 
to apply the collective term " Wound-rot " to those various 
forms of decay which have not yet been explained.^ 

The many forms of decomposition which are embraced under 
this term have not yet been subjected to scientific investigation. 
Should a large portion of the stem become functionless and die, 
saprophytic fungi belonging to the Hymenoniycetes or Ascomycetes 
induce decomposition, especially when their growth is stimulated 
by the unrestricted entrance of rain-water. This state of things 
exists in the case of snags destitute of buds, the stools of felled 
trees, trees that have lost large patches of bark by game, 
sun-scorching, &c., and which soon die to a considerable depth 
owing to the effects of drought. When water and air find 
easy access to a wound, as in the case of root-wounds, and 
branch-wounds that have not been tarred, decomposition spreads 
fairly rapidly in the direction followed by the water in the 
elements, although this wound-rot certainly does not progress 
nearly so rapidly as that which is due to parasitic fungi. The 
so-called false duramen of the beech always proceeds from a 
wound, and not onl}' arc all the vessels filled with tyloses, but 
the tannin is also so changed as to produce brownness in 
the heart-wood. Saprophytic fungi slowly advance from the 
wounds, and produce decomposition in the false duramen. The 
sooner a wound is closed, either artificially or naturally, the 
better for the tree. When air and water are excluded, wound- 
rot advances so slowly as only to reach a depth of half an inch 
in a century, as is shown by the occluded branch-wound of an 
oak in my collection. 

The treatment of wounds follows from what has been said. 
Two objects have to be kept in view — first, the process of 
healing, and, secondly, protection against wound-diseases, both 
infectious and non-infectious. 

The most perfect form of healing — namely, the re-clothing of 
the wound with a new cortex — can only be looked for when the 
1 Zersef::i/ngsersrheininii;^en. &c., p. 63. 


injur}' is due to the separation of the cortex during the season 
of cambium-activity, and provided the cambium can be preserved 
against drought by the immediate appHcation of a bandage, 
which, however, must not come into contact with the cambium. 
The only practicable means consists in binding moist oil-cloth, 
straw ropes, moss, or such like round the stem. 

Should there be no prospect of a new cortex forming, every- 
thing should be done to favour the production of a callus. All 
dead and crushed portions of cortex which may press in- 
juriously on the edge of the wound should be removed with a 
sharp knife, only those portions of cortex which remain uninjured 
on the surface of the wound, and which are nourished through a 
connection with the edge, should be carefully retained. From 
these a callus is formed quite as quickly as from the edge of 
the wound proper. 

In order further to guard against wound-diseases, all loose 
portions of cortex along the edge of the wound should be 
removed, as moisture lingers for a long time between them and 
the wood, and is absorbed by the latter. The moisture itself 
hastens decay in the wound, and moreover it induces condi- 
tions that are favourable for the germination of the spores of 
infectious fungi, which thus gain an entrance into the interior 
of the tree. 

In the case of those conifers which arc supplied with resin- 
ducts, wounds need be protected only when a thick branch which 
possesses duramen is cut or broken off, or when the cortex 
has become detached by pruning or the barking of game during 
summer. The spruce is most exposed to wounds of this 

The wounds of dicotyledonous trees require protection at all 
seasons. In order to form a waterproof covering over the wound, 
grafting-wax is used by gardeners and coal-tar by foresters. 
I have never observed any injurious effect of the tar on the 
tissues, as has been repeatedly asserted by practical men ; in 
fact, I can affirm that it is only the ruptured organs and their 
walls that are penetrated and impregnated by the tar. Cells in 
the immediate neighbourhood of vessels, and libriform fibres that 
were filled with tar, remained healthy and perfectly sound after a 
number of years. 


" Preventitious " buds are also to be reckoned amongst the 
regenerative phenomena that follow on injuries to trees, and 
which compensate for portions that have been lost. Only a 
limited number of the axillary buds of a shoot develop in the 
following year to form new shoots. The majorit}' of these buds, 
and especially such as are situated in the axils of the bud-scales 
and of the undersized leaves towards the base of the shoot, 
remain imperfectly developed, and do not, as a rule, shoot out in 
the following year. It is these which constitute the dormant 
eyes, or " Preventitious Buds " of Theodore Hartig, so called 
because they are present on any given portion of stem from the 
first year of its existence. Only under certain circumstances do 
these burst forth into new shoots, e.g. epicormic * branches and 
the like. Preventitious buds is a term emplo}'ed in contradistinc- 
tion to adventitious buds, the latter indicating nciv buds that are 
formed under certain conditions. j- 

These axillary buds may remain alive for a hundred years and 
more, especially in the case of trees with a smooth rind, such as 
the beech, &c. 

It is only as regards apical growth that the preventitious buds 
(Fig. 136, a) are inactive, for they display a peculiar form of 
growth in length, which Theodore Hartig has called " Inter- 
mediary Growth." Each year the delicate vascular bundles, which 
extend from the medulla to the buds (Fig. 136, b), increase in 
length to the same extent as the portion of the stem on which 
the buds are situated increases in thickness. Such growth is 
perfectly analogous to the growth of the sucker-roots of Visaiin 
album, or to the growth in length of medullary rays. The bud- 
axis that is embraced by the stem possesses its own cambium, \ 
at the point where it crosses the cambium of the stem. 

The cambium of the axis of the bud, which divides at the 
same rate as the common cambium of the stem, annually pro- 
duces two portions of tissue — namely, a larger one on the inside 

* [Shoots which develop in this way on the trunk are known as Epicormic 
shoots. Such shoots are very common on old Elms, &c. — Ed.] 

f [" Adventitious," because they arise in places where they would not 
normally be expected. — Ed.] 

X [Embryonic tissue, which adds new tissues to those already existing in the 
suppressed bud. — Ed.] 



whose length corresponds to the breadth of the wood-ring, and 
a smaller one on the outside equal in length to the thickness of 
the new bast. A cambium region persists between these two 
portions till the dormant e}'e dies, when the bud-axis, which 
is disposed at right angles to the main stem, ceases to grow 
and is overgrown and enve- 
loped by the advancing wood- 

Numerous bud-axes tra- 
verse the wood of dicotyledo- 
nous trees, exactly as is the 
case with medullar)- rays. 
Should these be stimulated 
to form shoots (Fig. 136, r), 
the latter produce their own 
growth of wood, and both 
they and their medulla form 
an acute angle with the main 
axis of the stem. 

In the case of some trees, 
more particularly the beech, 
a certain proportion of the 
dormant eyes develop in a 
peculiar manner after the ces- 
sation of intermediary growth. 
Concentric growth in thick- 
ness of that portion of the 
wood of the bud-axis which 
is situated in the cortex and 
bast gives rise to the familiar 
wood-balls, or " spheroblasts " 
(Fig. 137), which project from 
the surface of the stem and 

Fig. 136. — Longitudinal section of a 
beech-stem, twelve years old. At a 
two dormant axillary buds are shown 
whose vascular bundles, h, stand at 
right angles to the main axis. A third 
dormant eye, c, had burst forth to 
form a shoot two years previously. 
A dwarf shoot, d, has been formed 
by the unfolding of a bud when the 
main shoot was a year old ; c, a shoot 
that has been dead for four years. 
Natural size. 

frequenth' exceed the size of 

rifle bullets. As they have no connection with the wood of 

the stem, they may be detached by a slight pressure.* 

In the case of our conifers, almost all axillary buds are in the 

* [These " Spheroblasts " are very common on the old Beeches in Windsor 
Park. Burnham Beeches, and elsewhere. — Ed.] 



habit of developing into dwarf shoots,* and consequently dormant 
eyes are very scarce on these trees. In the case of old pines 
only one or two buds remain dormant in each whorl, and in rare 
cases a dormant bud may be perceived to burst forth at the base 
of a shoot where the dwarf shoots (foliar spurs) are absent. 
Should a pine be so injured, by the repeated attacks of cater- 
pillars, that not only all the foliar spurs with their dormant 
buds but also the youngest shoots with their whorls of buds 
wither, the only buds that the tree retains are the dormant 
whorl-buds of the older shoots. These elongate to form the 
so-called " Rosette Shoots," which however 
are unable to preserve the life of the tree. 
The rosette shoots either bear simple lance- 
olate leaves alone, or along with these a few 
foliar spurs. 

In the case of the larch only about 10 per 
cent, of the leaves of the one-year-old shoots 
have buds in their axils, all of which develop 
to normal or dwarf shoots (leaf-fascicles). A 
lost leader can be replaced only by the 
vigorous development of one of these dwarf 

The spruce and silver fir are also but 
sparingly supplied with axillary buds, some 
of which, however, remain dormant until 
special circumstances stimulate them to shoot 
out. These dormant buds are frequently to 
be found in a whorl at the base of the annual shoot. 

The conditions under which dormant eyes may be stimulated 
to form vigorous shoots vary, but all agree in this, that the buds 
receive an accession of nutriment. As examples of stimulating 
conditions I may mention pruning, coppicing, light-thinning, 
defoliation by insects, late frost, &c. 

Adventitious buds are, generally speaking, comparatively 
scarce. Their first inception is not in the axil of a leaf but at other 
points of the stem, roots, or leaves, where they originate in after 
years, and are therefore supplementary to the axillary buds. 

* [e.^^. The pairs of leaves 0:1 a Scotch or Austrian Pine arise each on such 
a dwarf shoot or "foliar spur." — Ed.] 

Fig. 137. — Globular 
shoot ("sphero- 
blast ") of a beech 
which has been 
formed from a 
dormant eye after 
the latter had be- 
come disconnect- 
ed from its vas- 
cular bundles. 


It but rarely happens that adventitious buds originate above- 
ground on uninjured portions of a plant, whereas endogenously 
developed buds occur regularly on the roots of many species of 
trees (root-suckers). On the other hand, their occurrence on the 
callus or investing layer of a wound is a frequent phenomenon 
(Fig. 151). There they originate close beneath the surface in 
the meristematic parenchymatous tissue, where they form their 
ring of vascular bundles, which internally are in intimate union 
with the wood of the callus. 

Adventitious roots, which may occur endogenously both on 
the uninjured cortex and on wound-tissues, have a precisely 
similar origin. 


Of the endless variety of wounds, we need select for discussion 
only a few of the more generally interesting. 


Barking (peeling) by red deer is usually confined to conifers, 
though dicotyledonous trees, for example the beech, are also 
similarly attacked less frequently. Fallow deer, on the other 
hand, abrade most if not all of our forest trees, though certain 
trees, e.^: the ash, are specially liable to attack. Roe deer, 
hares, and rabbits also bark trees under certain circumstances. 
Roe deer cause a special form of injury by rubbing off the bark 
of young trees with their newly formed horns. 

During winter, game bark trees for want of food, the starchy 
cortex of smooth-stemmed trees being nibbled to satisfy hunger. 
In summer, when trees are easily peeled, the more characteristic 
feature of the injury consists in the separation of large flaps of 
cortex, and this is frequently done to a considerable height. 
Views differ as to the motive of peeling during summer. It ap- 
pears to me most probable that the game regard the rich store of 
sugar in the cortex as a toothsome morsel. Some believe that 
the animals find an important aid to digestion in the tannin of the 
cortex. Game are also said to peel trees for the sake of the lime 
contained in the bark. Excellent results attended the feeding 

1 R. Hartig, Zersetziingerschciniini^en, pp. 67 ct seq. 



of red deer in the forest district of Ramsau with a substance 
containing bone meal, as well as with a special powder (Hofeld's) 
consisting chiefly of phosphate of lime and oak-galls. It was 
reported that the trees were not afterwards barked. Others 
again believe that summer peeling is merely the continuance 
of a mode of obtaining food which necessity taught the animals 
during winter, and that game thus get into the habit of barking 
during summer even when other food is present in abundance. 

On account of their periderm remaining smooth for a long time 
up to the height of four or five feet, bark being formed only 
comparatively late in life, the spruce and silver fir are longest 
exposed to the danger of barking. In the case of these trees, 
therefore, it often happens that barking is repeated after an interval 
of several years (Fig. 139), and stems are not unfrequently to 
be met with which show evidences of having been barked at 
various ages as often as five times. 

As the Scotch pine and larch, especially the former, produce 
rough bark early in life, they are exposed to the danger of 
barking only for a short period. It is only that portion of the 
stem of the Scotch pine which is from three to five years old 
that is barked. The portions that are younger are protected by 
the leaves, and those that are older by the thick bark. 

The damage which results from barking varies with the spe- 
cies of tree, time of year, and dimensions of the wound. The 
resinous pine suffers but little, unless the stem is completely 
barked round. The exposed w^ood dries and becomes so 
abundantly impregnated with turpentine and resin that further 
decomposition is prevented, and evaporation of water from the 
internal layers is retarded. The wounds, however, close with 
great difficulty, because the coalescence of the callus-cushions 
is interfered with by the early formation of rough bark 

(Fig. 138). 

The spruce, on the other hand, suffers much more from 
barking, partly because it is not usually attacked till a later 
period of growth, when much larger wounds are formed, and 
partly — and more particularly — because the wounded surface is 
not impregnated with resin to the same extent as in the case of 
the pine. Less damage is done by barking during winter than 
during summer, not onl}' because in the former case the wounds 



are usuall}- smaller, but also because the wounds have the 
opportunit}' to become impregnated with resin before the 
season when a higher temperature favours the formation of 
wound-rot, or the germination of the spores of parasitic fungi. 

Should parasites gain an entrance, decomposition spreads 
rapidly in all directions, and results in the destruction of the 
tree.. In other cases the wound-rot merely induces the inner 
layers of wood to become brown, without, however, attacking the 
wood that is formed in succeeding years. Should the wound 
remain long open, wound-rot may assume very serious propor- 

FiG. 138. — Transverse section 
of a pine-stem showing a 
wound caused by the peeling 
of red deer over which a 
caUus has formed, but which, 
after twenty-four years, is 
not yet quite closed. One 
third natural size. 

Fig. 139. — Transverse section of a spruce-stem 
showing three wounds due to the barking of 
game. One half natural size. 

tions. As a rule it ascends in the stem only to the height of a 
few yards, so that, when this is the form of " Red-rot," the 
timber is sound after the removal of a few short lengths. As 
the spot where the bark has been removed offers the least resist- 
ance, it is evident that should the tree be loaded with snow it 
will break most easily at that point. 


The wood-mouse {Mtis sylvaticus) and the field-mouse {Arvicola 
ai'valis'^) especially injure young dicotyledonous trees by gnaw- 

1 A. arvalis is not a native of Britain, but other species of the same 
genus do considerable damage to trees in this country. — Tr-ans. 

R 2 


ing the cortex during winter. Young beech woods especially 
frequently suffer very severely. If one allows the injured plants 
to remain standing, most of them will develop in spring appa- 
rently in a perfectly normal manner, because the sap is conducted 
up through the wood as before. In the course of the summer 
the exposed wood generally dries up, the outer layers being the 
first to be affected, and wound-rot also makes its appearance. 
Should the cortex have been removed right round the stem 
above the collar, the plant loses the power of conducting water 
at the injured part, and withers. If one delays cutting over the 
plant till this has occurred, it seldom happens that any stool- 
shoots are produced. If, on the other hand, one examines the 
young plantation before the leaves appear, and cuts over all 
injured plants close to the ground, vigorous shoots will be pro- 
duced, at the expense of the store of reserve materials present 
in the roots, and in a short time the young wood will be almost 
as promising as before. The more vigorous plants may remain 
alive for several years, and adventitious roots may even be formed 
above the wound, as is represented in Fig. 140. 


Abrasions of the bark which are caused during the process 
of removing timber from the wood, especially on declivities, 
are amongst the commonest form of wounds to which shallow 
roots and the lower parts of stems are subjected. During the 
dragging of timber, and especially when water is in active 
movement in the tissues, large portions of cortex are detached 
from the base of growing trees. Where cattle are grazed or 
folded, and where roads occur in a wood, the shallow roots are 
subjected to all sorts of injuries, and from these, in the case of 
the spruce, the wound-rot ascends in the stem, attaining to a 
height proportionate to the amount of moisture that enters the 
wound from the soil. On this account wounds that are covered 
by moss or humus are much more dangerous than those which 
are perfectly exposed. 

The majority of the brown patches of red-rot that are observ- 
able on the cut surface of the stool of the spruce, and which 



disappear when one or two short lengths of the tree are 
removed, ma\' be traced to such wounds on the root or collar 

(F"ig. 141 \ Should the mycelium of 
Agaricus melleus gain an entrance into 
such root-wounds, decomposition pro- 
ceeds much more rapidly, and the 
lower part of the stem may become 
perfectly rotten. 

When wood-ants {^Formica Jiercu- 
Icaiia, or F. ligniperda) take possession 
of these wounds, they frequently form 

Fig. 140. — A beech which has 
been severely barked by mice 
above the collar. On the left 
side a stripe of cortex has 
been left. Numerous adven- 
titious roots are seen breaking 
through the uninjured cortex 
above the wound. Natural 

Fig. 141. — The stool of a spruce that had formed 
two stems. One of the stems, a, had been 
removed in the thinnings, and from it decom- 
position subsequently spread downwards into 
the sound stem, b. At c c wounds have been 
formed in the cortex during the dragging of 
timber, and at e wound-rot has spread upwards 
from a damaged root into the stem. One tenth 
natural size. 

galleries which extend far up into the sound part of the stem, 
and rapid decomposition succeeds their excavations. 

Intentionally or unintentionally, man is accountable for the 
most varied forms of bark-wounds. Take, for example, the 


carving of figures or letters. Should these be formed in the 
cortex the wound will be of the same shape as the figure, 
and the latter may be recognized for many decades, even 
after callus has been formed, owing to the difference in the 
appearance of the old and new cortex. If, on the other hand, 
the cortex is first removed from a considerable surface of wood, 
and the figures are carved in the wood itself, they disappear 
when the wound closes. All that can afterwards be perceived 
is the boundary between the old cortex and the place from which 
the cortex had been removed.* 

When it is intended to remove a ring of dead bark from the 
Scotch pine for the purpose of la}'ing on a band of tar, the living 
bast, and even the wood, are frequently unintentionally cut into 
as well. Even after the tar has been laid on, turpentine and resin 
continue to exude from the wound and form a white covering on 
the black tar. This has given rise to the erroneous impression 
that the tar partially dissolves the cortical tissues and causes 
wounds in the bast. 

Precisely similar wounds result from the removal of bark from 
old pines for the purpose of obtaining fuel for laundries, as 
occasionally happens in the neighbourhood of towns. When 
climbing irons are used for scaling trees wounds are also 
extensively formed, and especially so during the harvesting 
of cones, and the cutting down of spruce-branches for litter. 


During the felling of timber in a close wood, it often happens 
that a falling tree, or one of its branches, strikes an adjoining 
tree, stripping off and crushing the cortex. During pruning the 
top rung of the ladder crushes the cortex of the branch against 
which it is laid. In dealing with insect ravages it was formerly 
a common practice to shake the trees by beating them violently 
with the back of an axe so as to frighten the caterpillars and 
make them drop off. In consequence of crushing due to these 

* [Such cut letters, &c., are often found deep down in the wood many years 
later, the successive annual rings formed by the occluding callus having 
covered them completely over. The burying of wire, nails, chains, &c., deep 
in the wood is due to similar occlusion by a callus which gradually forms 
wood over the edges of these objects. — Ed.] 


causes the cortex dies, and growth ceases at the injured spot. 
But more than that, the dead cortex remains for a long time in 
union with that which is Hving and uninjured, and no formation 
of callus can take place, because growth is stimulated along the 
edge of the wound only when the bark-pressure is reduced. 
The formation of wound-rot is encouraged by water collecting 
behind the dead cortex, which becomes locally fissured owing to 
shrinkage consequent on drying, and finally rots away, but only 
after the lapse of many years. 


Turpentine and resin are procured from conifers in various 
ways. In the case of the silver fir, it is only the turpentine 
that is gathered. This collects in vesicles in the cortex, which 
sometimes attain to the size of a pigeon's egg (Strasburg 

In the case of the larch, large holes are bored into the stem, 
and these being afterwards plugged up collect the " Venetian 
Turpentine " which flows down from the vertical resin-ducts of the 
wood. In the case of the black Austrian pine, the cortex is 
removed from the stem in fairly broad stripes, the turpentine 
that exudes freely from the canals of the medullary rays being 
collected in a receptacle that is cut in the stem below the 
wound, while the resin is scraped from the wound after it has 
solidified. On account of the exposed wood soon becoming 
impregnated with resin, and the canals of the medullary rays 
becoming choked up with the same substance, it is necessary 
from time to time to remove fresh portions of cortex at higher 
points on the stem. 

In the case of the spruce, vertical strips of cortex, one to two 
inches in breadth and extending from the base of the stem to a 
height of about six feet, are detached from the tree. When the 
tree is small the resin is taken from one side only, but as it gets 
thicker four sides may be utilized (Fig. 142). When the flow of 
resin ceases, the callus that has been formed along both sides of 
the wound since the last time of stripping is removed, and thus 
a new set of resin-ducts is opened, from which resin continues 
to flow. 



In the course of }'ears the exposed wood dries up, and decay 
begins to make its appearance, being greatly favoured b}' the 
larvse of Sirex, which bore from the surface of the wound deep 
into the wood, and thus enable rain-water to reach the interior of 
the tree. Decomposition frequentl}' spreads from the wound 
high up the tree, and does so much damage that in woods where 
resin is collected the yield of timber ma}' be reduced from seventy 

Fig. 142. — Transverse section of the stem of a spruce which has been tapped for 
resin on four sides for ten to fifteen years. The only wood that is capable of 
conducting water is those portions of alburnum, a, which are marked off by lines 
between the four gutters. The wood, /', beneath the two upper gutters is much 
decomposed, whereas the wood, c, beneath the other gutters has remained sound. 
Numerous galleries, £, formed by Sirex are seen proceeding from the upper 
gutters. One fifth natural size. 

to twenty or thirty per cent, of the gross output. It has 
not been proved that trees that are tapped suffer in growth, 
nor is it to be expected that such will be the case, seeing that 
trees cannot utilize turpentine for growth. Tapping, however 
greatly reduces the value of timber, because the quality to 
a large extent depends on the amount of resin which it 



These are often caused b}- game and mice, though they maj- 
also be due to human agenc}', as, for instance, in a mixed wood, 
where it is desirable to protect a valuable species against its 
more vigorous neighbours. Their effects upon the tree are not 
always alike. It is known that, if even a narrow band of cortex 
be removed completely round the stem, the cambium below 
the girdled portion ceases to be nourished, and there, as a 
consequence, growth in thickness comes to a stand-still. As the 
tree even where ringed retains its power of conducting the 
ascending sap, it remains alive as a rule for some years. What 
the conditions are that limit the duration of life of the portion 
above the ring-wound has not }'et been full}' made out.^ In June 
1 87 1 I selected fifteen equal-sized Scotch pines 120 years old 
which were standing close together, and from these I completely 
removed the cortex to the height of some six feet. While 
certain of the trees died in 1872, several were still perfectl}- 
healthy in 1877. As this shows that it is not the desiccation ot 
the exposed portion of the stem from the surface inwards that is 
the sole cause of death, investigation should be directed to the 
question whether the cessation of growth beneath the ring- 
wound ma\' not prejudicialh' affect the absorption of water b\- 
the roots. 

Those cases where ringed trees remain alive for a long period 
may possibly be explained b}' root-engrafting, the roots of the 
girdled stem being thus nourished b}' neighbouring trees. 


Although the pruning of trees is a subject that has often 
been treated in forestry literature, still the views regarding its 
admissibility are so diverse that a somewhat full discussion of 
the operation may not be out of place here. 

The natural pruning of trees is accomplished by shade, which 
causes the branches to become functionless, and ultimatel}' to 

^ This is not the place to discuss bicoUateral fibro-vascular bundles, 
where the plastic materials may descend in the bast organs near the pith^ 
- R. Hartig, ZerseiziingscrscJicinitngcn, pp. 68 ct scq. 


die. The dying twigs and branches are more or less quickly 
decomposed by saprophytic fungi. 

The rate of decomposition and the period when the branches 
will drop off are most of all regulated by the condition of the 
wood. Branches of dicotyledons which consist only of alburnum 
drop off much sooner than branches which contain duramen. 
On account of the shaded branches of young Scotch pines 
consisting of soft broad-ringed wood, these trees clean them- 
selves much sooner than the spruce and silver fir, the wood 
of whose branches is tough, firm, and durable. The thicker, 
more resinous, and narrower-ringed branches on the upper part of 
the stem of the Scotch pine, on the other hand, retain their 
position for a long time, and are more or less embraced or over- 
grown at their bases by the growth of the stem. This embracing 
of dead branches is the general rule in the case of the silver fir 
and spruce, and as they have no organic connection with the 
adjoining wood-layers they drop out of boards as loose knots 
when the wood shrinks in drying. 

The embracing of dead branches would be a much commoner 
occurrence, were it not for the fact that the base does not die, 
and in the case of the thicker branches it often remains alive for 
a distance of about two inches (Fig. 143). The base of the 
branch, being nourished from the stem, remains alive, and is 
capable of growing in thickness. When, after some }'ears, the 
increase in thickness of the bole of the tree has become equal to 
the length of the living basal part of the branch, the dead part 
of the branch will have become so much decayed as to drop 
off under the action of wind, snow, &c. (Fig. 144). After the 
wound has healed over only a small dark brown blotch remains 
in the interior of the tree to indicate the limits of the enclosed 

It is in the manner just described that the tree protects itself 
against the dead stumps of branches being overgrown. It is 
only the larger branches that frequently do not drop off until a 
portion of the dead base has been embraced by the stem. In 
the case of conifers this portion is saturated with resin, and in 
the case of dicotyledons it is more or less decomposed. After- 
wards, when the branch has become completely rotten and has 
dropped off, a hole remains behind which is only partially filled 


by the occluding callus, and which of course greatly reduces the 
value of the tree for technical purposes (Fig. 145). 

Thus it is always a good plan, in the case of every variety of 
tree, to remove as early as possible all the larger dead branches 
that have succumbed to the natural shading processes. I do not 

Fig. 143. — An oak-branch which 
has succumbed to the natural 
process of shading, its base, /', 
however, still receiving nourish- 
ment from the main stem. 

FiCt. 144. — The snag of an oak-branch which 
has dropped off after being killed by the 
natural process of shading. The basal por- 
tion, h, of the branch which remained alive, 
and originally projected from the stem, has 
been grown over. After a callus has formed, 
the dark brown zone, c, between the living, 
/', and the much -decomposed wood, a, re- 
mains unchanged in the interior of the tree, 
as is shown in the case of a small branch at 
d. The axis of the stem of a latent bud is 
shown at e. 

propose to enter into the technique of the operation, merely 
remarking that it is evident that the expense should be incurred 
only in the case of such trees as promise to yield high-class 
timber. With this limitation there is no doubt that as forestry 
advances the pruning of dead branches will become general. 



The contention that such pruning is too costly is justified only 
when it can be proved that the difference in value between a log 
free from knots and one where they are abundant is not equal to 
the cost of pruning phis interest on the outlay. 

When we come to consider the removal of green branches — 
that is to say, branches or twigs that are living and provided with 
leaves— we find that, except in a few exceptional cases which 
will be presently discussed, a loss of growth attends the oper- 
ation. This is the case no matter whether the separation 

from the stem be effected 
b}' the hand of man or 
b}' such natural agents 
as storms, accumulations 
of snow, &c. If one re- 
duces the number of the 
organs of assimilation 
(the foliage leaves), the 
products of assimilation 
generally suffer to a like 
extent. As I have proved 
conclusively,^ it is only 
in the case of trees that 
are growing in a per- 
fect!}^ open situation, 
whose stems are branch- 
ed to the ground, and 
which have a very large 
mass of foliage, that 
limited pruning may be performed without diminishing the 
amount of growth. In the case of such trees there is a greater 
extent of foliage than is necessary to effect the metabolic 
processes in the plant-food that is taken in by the roots. Of 
course the amount of growth depends essentially upon the 
quantity of such food. Under such circumstances a reduction 
in the extent of the foliage merely results in more active 
assimilation in the leaves that remain. 

In the great majority of cases the practical operation of 
pruning is followed by more or less considerable reduction in 
^ Das Holz der Rothbiichc. Berlin, Springer, 1888. 

Fig. 145. — The dead and rotten stump of an oak- 
branch over which an occluding caHus has 
formed. Two thirds natural size. 


growth. This becomes evident in the lower region of the stem, 
where in fact, if pruning is carried far enough, growth ma}- cease 
altogether, as I have also proved to be the case with trees that 
are very much overcrowded. 

One must always bear in mind that as pruning generally 
interferes with growth there must be important reasons for 
performing the operation if the loss of growth is to be com- 
pensated for. Amongst these may be mentioned, on the one 
hand, the improvement of the form of the stem and the 
production of a clean bole, and, on the other, the admission of 
light to underwood. 

If, for the purpose of obtaining smooth stems, pruning is 
carried further than the mere removal of a few branches, one 
must remember that in such a case there is not only reduction 
of growth, but that there are also indirect dangers consequent on 
such pruning. The first of these dangers is connected with the 
retardation of the healing of the wounds. It is evident that 
the formation of callus over a branch-wound depends to a large 
extent on the supply of plastic substances with which the 
cambium along the edge of the wound or the callus-cushion is 
provided. Very severe pruning will seriously interfere with the 
formation of callus, and consequently with the occlusion of 
wounds. This leads us to consider whether the pruning of the 
stem to the desired height should not be accomplished in two 
operations, separated by an interval of several }'ears. If one first 
of all removes the branches from the lower half of the portion of 
the stem that it is desired to clear, the diminution of the products 
of assimilation does not interfere to such an extent with the 
formation of callus, and the wounds may be covered over in 
a few years. The more vigorous development of the crown 
compensates to a certain extent for the branches removed by 
pruning, so that when the operation is repeated the new wounds 
close sooner than would have been the case had the whole 
operation been performed at one time. 

By dividing the operation in this way there is also much less 
chance of an excessively large number of epicormic shoots being 
produced. Such shoots originate partly in the adventitious 
buds of the callus along the edge of the wound, and partly in 
dormant eyes. In the latter case, it is chiefly the buds that are 


situated on the basal portion of the severed branch that is 
embraced by the stem which produce the shoots. 

When a spruce is pruned, numerous shoots spring apparently 
from the cortex of the main stem. These are chiefly due to 
the vigorous development of small weak dwarf shoots, which 
originated at the base of the branches in their first year, and 
which have become occluded during the increasing thickness 
of the stem. I have not been able to prove that true adventitious 
buds are formed in the case of this tree. 

If in pruning green branches one leaves the stump of a 
branch (snag) without any foliage, the same state of things 
occurs as when branches are suppressed naturally. The 
snag dies, except for an inch or two at the base, and the 
formation of callus is either rendered impossible or is so much 
interfered with and delayed that the dead stump has time to 
become completely rotten. If the bark is removed from the 
snag, the conditions are rendered more favourable for the forma- 
tion of callus, and a covering will more easily grow over the 
snag from the base than is possible when the dead and dry 
cortex remains in situ on the stump. In Fig. 146 I have 
represented the progress of the formation of callus on a thick 
snag, where for clearness the bark has been mostly removed. 
The bark of the dead snag presses firmly on the wood, and the 
formation of the new growth («, U), which already covers more 
than half the stump, has been rendered possible only by its 
pushing in like a wedge and separating the dead cortex from the 
dead wood, so that the thin and primarily non-vascular edge of 
the living tissues has been enabled to grow into the space that 
has thus been formed. The familiar curled growths on the 
stumps of branches are formed when the new tissues advance 
unequally, as is most frequently the case when they are growing 
over an irregularly fractured surface (Fig. 146, x x, in the upper 

As a dead snag interferes with occlusion, the general rule in 
pruning is to cut as close as possible and to make the cut 
parallel to the stem. If this is attended to, a callus is formed in 
the way already described, its formation proceeding most rapidly 
from the lateral edges of the wound. For obvious reasons the 
bark is there most easily raised, much more easily, in fact, than 



along the upper and lower edges. The upper edge, however, 
is greatly favoured as compared with the lower edge, because 
the plastic substances during their passage down the stem are 
conveyed directly to the former, whereas the latter lies out of the 
stream as it were, and is but sparingly supplied with nutriment 
(Fig. 147). 

There is, however, a much more important reason for the 

Fig. 146. — A fractured oak-branch. The wound is being gradually occluded by a 
callus which is slowly advancing beneath and pushing off the thick cortex. At 
a the new gi'owth shows curls, while at b it pushes its thin non-vascular edge 
forward regularly over the dead wood. The dead wood is represented at c. 
One fourth natural size. 

slow formation of callus on the lower part of a wound. In 
that region the cortex is, as a rule, loosened from the wood 
during the operation of pruning. At the time when the 
cambium is active, it is quite impossible to prevent the cortex- 
being loosened, the friction of the saw being sufficient to 
account for it. But the main cause is to be traced to the fact 
that, in order to prevent the cortex being torn off, a cut is first 
of all made underneath, and during the sinking of the branch 
the lower edge of the wound is subjected to severe pressure. 



The cortex of the lower edge of the wound forms a pivot 
round which the sinking branch turns, and, although the effects 
may not be immediately visible, still the crushing and tearing 
at that point kills the cambium for an inch or two back from 
the edge of the wound. Of course, in such a case, the new 
growth — namely, the callus — is not formed at the edge of the 
wound, but at a considerable distance from it, where it is covered 
by the cortex (Fig. 148). The result is that the cortex, which was 
originally in intimate contact with the wood, becomes detached, 

Fig. 147. — A branch-wound on 
an oak which has been half 
occluded by a callus. 

Fig. 148. — The lower edge 
of a branch-wound one 
year after being formed. 
The cortex, a, that has 
been crushed during the 
sinking of the branch, 
dies as far as /', at which 
point the formation of 
callus, c, begins, and the 
cortex is gradually separ- 
ated from the wood. 
Natural size. 

SO that a cavity is formed beneath the wound between the wood 
and the dead tissues. This cavity acts like a gutter to catch the 
rain-water that flows over the surface of the wound, as well as 
all the organisms that it may contain. This forms a specially 
suitable place for the germination of the spores of parasitic 
fungi, and it is from here that water containing the soluble 
products of decomposition finds its way by means of the 
medullary rays into the interior of the wood. This cavity is 
a gutter in every sense of the term, and at the same time the 
point of attack for fungi. Even although the surface of the 
wound may have been coated with tar immediately after 
pruning, this spot remains unprotected, and indeed it is only 


formed after the cortex has been separated from the wood by 
the advancing callus. It is in fact the Achilles heel of the 
branch-wound. In pruning, the main object must be to prevent 
its formation, but this is possible only if pruning be confined 
to autumn and winter, when growth is at a stand-still, and when 
the cortex is least liable to be detached from the wood. If 
one also takes the precaution to support the branch during 
sawing, and at the moment of separation to push it clear of the 
wound, danger is reduced to the minimum. 

The rate at which a wound is occluded depends entirely upon 
the vigour of the tree and the size of the wound. A callus 
forms on young trees, with their relatively broad annual rings, 
faster than upon old trees, and the faster too the higher on 
the stem the wound is situated, because with few exceptions 
the breadth of the rings increases as we ascend. It is equally 
apparent that occlusion will be accomplished sooner where the 
situation is good than where bad. In the case of dicoty- 
ledonous trees, especially the oak, to which my investigations 
have hitherto been confined, branches of a greater diameter than 
4 — 5 inches should not be removed. 

The effects of pruning as regards the health of the tree 
depend chiefly upon the period of the year in which the opera- 
tion is performed. So far as my observations go, it is always 
highly dangerous to prune the spruce during summer, as 
rapidly advancing wound-rot is an almost invariable con- 
sequence. It may be mentioned, however, that in all the cases 
which I examined, the cortex had been injured during the 
process of pruning. This may be avoided by pruning during 
autumn or winter, and as the cut surface becomes immedi- 
ately covered with a resinous exudation the wound is almost 
certainly safe from rot. It is only in the case of the older 
branches, where the heart-wood emits no turpentine, that parasitic 
infection is liable to occur. It thus appears to me that conifers 
may be pruned in autumn and winter, if the wounds caused by 
the removal of the larger branches be coated with tar, but 
since in the case of these trees the branches are generally small 
this will seldom be necessary. 

When the wounded surface of a dicotyledonous tree has not 
been tarred, one first observes a brown colour penetrating to a 




depth of an inch or two, and this is succeeded in a few years by 
wound-rot, which, however, ceases to make progress when the 
wound closes (Fig. 149). If pruning has been done in summer, 
it will be found that beneath the wound brownness appears 
in the youngest annual ring, and often spreads down the stem 
for four or five yards. If one omits to apply a coat of tar, the 
danger of infection by parasitic fungi is naturally increased. 
These, however, penetrate even into tarred wounds should they 

Fig. 149. — Oak pruned in July. 
Rot has spread from and be- 
neath the untarred surface of 
the wound far into the stem. 
One third natural size. 

Fig. 150. — The occluded branch- 
wound of an oak which has been 
infected by Hydniim diversidens. 
One half natural size. 

be formed in spring or summer, because at that time the germ- 
tubes are able to enter beneath the lower edge of the wound 
(Fig. 150). 

Tarring produces satisfactory results only when pruning has 
been done in late autumn or in winter, because it is only then 
that the tar is absorbed by the surface of the wound. It would 
appear that the absorption of the tar is due partly to the 
diminished amount of water in the wood during autumn, and 


parti}' to the consequent negative pressure of the air in the 

When pruning is undertaken in spring or summer,, the tar 
altogether fails to enter the wood, and the thin superficial layer 
does not prevent the cut surface from drying later, and forming 
fissures into which water and fungi may gain an entrance. And 
then, again, the separation of the crushed cortex from the 
lower edge of the wound frustrates the object of tarring. 

From what has been said, it follows that dicotyledonous trees 
may be best pruned in the months of October, November, and 
December — perhaps also in January and February — and that a 
good coat of coal-tar should at once be applied to the wounds. 

Hitherto pruning has usually been undertaken in summer, and 
this explains wh}' the operation has caused such enormous 
damage to trees, especially the oak. It is, however, desirable 
from every point of view that the subject be further investi- 
gated with scientific accuracy. Several species of trees should 
be taken in hand, because the trials which I conducted were 
confined to the oak, and even in their case sufficient time has 
not yet elapsed to make it possible to furnish conclusive 
answers to all the questions that have just been raised.^ 


The shortening of the branches of plants from three to ten 
or twelve feet in height differs from pruning proper only as 
regards the size of the branches. Most of what has been said 
in connection with pruning may be applied here. It therefore 
follows that all shortening of branches is an evil which can only 
be excused when important objects are to be gained. The 
dressing of the younger classes of plants is most admissible at 
the time of transplanting, when the number of the roots has been 
considerably reduced. In the early part of the season when 
foliage is scarce, and when transpiration of water proceeds but 
slowly, the quantity of roots may suffice ; whereas in summer 
the diminished mass of roots may be unable to provide sufficient 

1 It is very desirable that observations be Gontinued on some 240 pruning 
experiments that I carried out in 1S75 in the woods attached to the Forest 
School of Ebersvvalde. 

' S 2 


nourishment for the undiminished crown, which consequently 
withers. The danger is avoided, and the plant gets over the 
loss in a short time, if equilibrium between the roots and the 
foliage is restored at the very first by shortening the longer 

A second reason for shortening the branches is the im- 
provement of the shape of the plants, whether in the nursery or 
in the wood. I do not intend in this place to enter upon the 
technique of the subject, but will merely say that so far as the 
growth of the plant is concerned the usual time — namely, summer 
— is the least suitable. If we dress a plant in spring or autumn 
we remove, in the main, only the branches, the reserve 
materials being left in the storehouses of the stem. But if 
summer be selected for the operation, the reserve materials of 
the stem, being partially utilized in the production of shoots 
and leaves, are lost. If one waits till autumn, the leaves of 
the branches to be removed will have assimilated materials for 
the following year, and these will have been partly deposited in 
the main stem. It appears desirable to institute investigations 
in this direction, and the question whether wounds are least 
attacked by parasitic fungi, such as Nectria, during summer or 
during autumn and spring should also receive careful attention. 
This question has special force with respect to Acer, Tilia, and 
Aesc2iliis, seeing that these genera suffer most from Nectria 
cinnabarina, and in their case even small wounds should be 
protected by grafting-wax. 

The practice of leaving snags destitute of buds on the main 
stem is justly condemned, for the reason that if growth is rapid 
they are partly embraced or completely enveloped when dead and 
withered. On the other hand, it is a mistake to suppose that 
decay spreads in the wood from such snags in after years, for I 
have never been able to observe such a state of things even in 
oaks that had been pollarded or coppiced in youth. 

As the wounds are small and are usually soon occluded by a 
callus, the application of tar is scarcely necessary, except in the 
case of the above-named trees, which are specially liable to suffer 
from Nectria cinnabarina. The technical properties of timber 
are not interfered with by the small brown wounds in the body 
of the stem, for it must be borne in mind that numerous wounds 


of a similar character are also formed when branches drop from 
trees naturall^^ 

As has been already stated, it sometimes happens that 
parasitic fungi, especially species of Nectria, enter through 
branch-wounds and produce cancerous diseases, which afterwards 
spread in the stem. 


When the spruce is grown in open lines in the nursery, it 
tends to develop a double leader when about three or four years 
old, so that instead of a single stem we find two. If one of the 
two stems is not removed till the first thinning, the base dies and 
decays exactly like the snag of a branch (Fig. 141), and becomes 
enveloped more or less by the other stem. The wound-rot spreads 
easily from the stump to the other stem, in which it may ascend 
to the height of four feet. 

In order to avoid this injury, one of the shoots should be 
removed in early life, as is easily done by means of a knife with 
a long handle and a bent blade. In rare cases the technical 
properties of the timber are reduced by a double leader again 
forming in later life. Such an occurrence, however, happens 
but seldom, and probably only when the tree occupies a very 
open situation. 

Less damage is done by removing, during the first thinning, a 
stem that has grown into another at the collar, as sometimes 
happens in a very dense wood. Such cases of natural grafting 
occur most frequently in woods that have been formed by 
planting the young trees in bunches. Seeing that the stems are 
separated by their cortex up to the twentieth or thirtieth year, 
when the first thinning takes place, the coalescence is usually 
only apparent, and the removal of one stem scarcely injures the 


When trees are cut over close to the ground, various 
phenomena of regeneration which vary with species and age 
make their appearance. Amongst conifers the Scotch pine 
produces stool-shoots from dormant eyes only when very young. 


In the case of that tree the axillan- buds of the primar)- leaves 
preserve their vitaht}' until the formation of bark begins, usuall)- 
about the fifth year, when the)' perish, and with them the power 
of producing stool-shoots is lost. 

Those American pines which have three leaves in the sheath, 
for instance P. rigida, retain the power of producing stool-shoots 
till a late age. This is owing to dwarf shoots being developed 
partly in the whorls, and partly on the main axis midwa}' 
bet\\'een the whorls. These dwarf shoots grow each }-ear to an 
extent corresponding to the growth in thickness of the stem, and 
produce but few leaf-fascicles. It is these that give rise to an 
abundant growth of stool-shoots. On account of the absence of 
dormant eyes that are capable of producing shoots, the regene- 
rative power of the stools of conifers is a very limited one, if 
we except those cases that have been quoted. The formation of 
adventitious buds in the callus of wounds is also ver}- exceptional, 
and it is only in the case of the silver fir that I have occasionally 
observed new buds and shoots produced from the callus of the 
stool. On the other hand, it frequentl}' happens that the stools 
of conifers — more especiall}- those of the silver fir, spruce, and 
larch, very rarely those of the Scotch pine — live for several 
decades, and form callus more or less energetically along the 
edge of the cut surface, so that in certain cases the whole of the 
transverse section ma)- be occluded. It is probable that the 
formation of callus on the stool is generally due to the natural 
grafting of the roots of the tree that has been felled (the 
nourished stem) with those of an adjoining tree (the nourishing 
stem). There is, however, no getting over the case quoted by 
Th. Hartig, where a larch-stool showed a growth of callus 
notwithstanding the fact that the tree had stood in a large gap 
in a wood, so that the possibilit)' of nutriment being transferred 
from a neighbouring tree was absolutely precluded. This case 
can only be explained by supposing that in the course of years 
the reserve materials stored up in the roots and stool were 
dissolved and applied to 'the nourishment of the cambium. 

If the cortex and cambium have not been destroyed for some 
distance back, by the drying up and decay of the wood, the 
stools of dicotyledons develop a callus and numerous buds 
during the year succeeding that in which the tree was felled. 



These adventitious buds frequently produce vigorous stool-shoots 
(Fig. 151), which, however, fail to become self-rooted, and suffer 
from the advancing decomposition of the parent stool. The 
stool-shoots that are formed from dormant e}'es are much more 
serviceable, and also more abundant. As it is very desirable 
that these should become self-rooted, so that the new plants 
ma}- be unaffected by the health of the parent stool, it is an 
advantage to have them as low down on the stool as possible. 
For this reason coppice poles are cut as low as possible, and 
in order to destroy all shoots that have 
formed too high up on the stools, and thus 
encourage the formation of deeper shoots, it 
is a common practice in oak coppice to char 
the stools by burning any ground vegetation. 

As the dormant eyes preserve their 
vitality for only a limited period, no shoots 
need be expected from old stools. The stools 
of the older class of birches produce abun- 
dant shoots, which however usually succumb 
after a year or two. The reason for this 
is that the extremely hard bark does not 
yield to the growth in thickness of the 
shoots whose base it envelops. The result 
is that when, on account of the base of 
the shoots being nipped by the bark, the 
supply of water fails to keep pace with the 

accelerated transpiration, the shoots formed early in the' }'ear 
succumb about midsummer. 

When young dicotyledonous trees that have become stunted 
in growth are cut over close to the ground, the young shoots 
often grow so satisfactorily and persistently that the plan is 
frequently practised with good results as a cultural measure. 
Although this matter has not yet been made the subject of 
scientific investigation, it seems probable that after the tree has 
been cut over the reserve materials present in the roots and 
stool are utilized in stimulating root-growth, so that when the 
roots have penetrated to a deeper, fresher, and richer layer of the 
soil the plant continues to grow satisfactorily. Stunted oaks 
that are situated on ground that is covered by weeds or heather 

Fig. 151. — Shoots that 
have formed from 
adventitious buds on 
the one-year-old cal- 
lus of a beech-stool. 
Natural size. 


are often induced to throw out strong shoots, and to show 
vigorous and persistent growth, by setting fire to the whole 


These are partly due to animals, e.g. mice, but are mostly 
caused during cultural operations, and are always prejudicial to 
the plants. The greatest care must therefore be exercised to 
preserve the roots during lifting, transport, and planting. 

Pruning the roots is always an evil, and is admissible only in 
two cases. The first occurs when roots are crushed, nipped, or 
broken off during the process of lifting. A clean cut imme- 
diately above the damaged part encourages the formation of a 
callus from which adventitious roots are produced, and also 
prevents or reduces the chances of decay in the roots. The 
second case where shortening the roots is admissible occurs 
where it would be too expensive to preserve the whole root- 
system during the operations of lifting and planting. It may be 
mentioned that many plants suffer less from their roots being 
shortened than from their being doubled back during planting. 
In order to induce the formation of a dense mass of roots by the 
production of numerous roots in the neighbourhood of the collar, 
repeated shortening of the roots may be necessary where the 
attainment of extra large plants is the object in view. 

The practice, which is unfortunately still so common, of 
aimlessly cutting back the roots is in the highest degree 

Other forms of root-injury are occasioned by removing litter 
from woods, tearing up roots, the attack of cockchafer grubs, 
mice, &c. 


The growth and future success of parts entirely destitute 
of roots, e.g. slips, pole-cuttings, &c., depends essentially upon 
the greatest possible restriction of evaporation from the plants 
until they have produced an abundant supply of roots. For 
this reason one at first suppresses the development of leaves 
by almost entirely burying the cutting, so that only the highest 
bud is able to produce a shoot ; or, in other cases, and more 


especially in Horticulture, the rootless cuttings are placed in 
a chamber where the air is saturated with moisture. 

Cuttings of the Caspian willow, that appear to have rooted 
perfectly, frequently die off on sandy soil in the height of 
summer, or in the autumn of the first year. The reason for this 
is that in the earK' part of the season adventitious I'oots appear 
both upon the cortex and the callus of the cutting, and when 
the upper layers of the loose sandy soil dry up the greater 
portion of the roots on the cortex, most of which are dis- 
posed horizontally, die off. When this is the case, it often 
happens in the height of summer that the roots which have 
originated in the callus of the wound, and which always 
penetrate the soil obliquely, are unable to supply sufficient 
water to satisfy the wants of the leafy shoots, which con- 
sequently wither. On this account the soil of osier-beds should 
be worked to as great a depth as practicable, so as to encourage 
the roots to go deep. 


A technical discussion of the various operations connected 
with the transference of a living shoot or bud from one 
plant to another would be entirely out of place here. It is 
sufficient that we should shortly consider the internal changes ^ 
that are associated with the process. If we except grafting by 
approach, where two adjoining plants are so united to each 
other at one or more places that similar wounds in the cortex 
of both plants are brought into and retained in intimate con- 
tact till complete coalescence has taken place, we find that all 
grafting operations agree as regards the main principles. A 
portion of a plant provided with buds but destitute df roots, the 
so-called scion, or only a portion of cortex furnished with a bud 
(the shield and eye), is united to a rooted plant, called the wild 
plant or stock, in such a way that when coalescence takes place 
water and food-materials will be transferred from the stock to 
the scion, as well as plastic materials from the scion to the 

The operation succeeds, as a rule, only when, on the one 

^ Goppert, I line re Ziistdnde der Biiiinie nacJi iiusseni Verletzinigcn. 
Breslau, 1873. 


hand, the cambium of the stock is active, so that immechate 
coalescence may take place between the callus-tissues produced 
by its cambium and by the cambium region of the scion, and 
when, on the other hand, the scion or bud is at the same time 
inactive. The coalescence, in fact, demands a certain time. 
Should the scion become active before coalescence has been 
effected, or should its buds even be swollen at the time of the 
operation, it withers in consequence of transpiration from the 
young leaves before it can obtain a sufficient supply of water 
from the stock. On this account the scions are prepared as early 
as February, and are preserved in such a manner that by 
repressing the tendency to growth as far as possible they will still 
be inactive at the time when the stock has burst into leaf. As 
is well known, budding is usually undertaken in summer, after 
the new axillary buds have been formed, the buds being united to 
the stock at a time when cell-division is still active in its cambium. 

The scion and stock are united in such a way that their 
cambium layers are brought into as intimate contact as pos- 
sible, care being also taken that no considerable interspaces 
are left between the cut surfaces of the wood. According to 
Goppert's investigations, coalescence is due to two distinct 
processes, for not only does union occur between the cambial 
layers, or the callus-tissues that are produced from them, but also 
between the cut surfaces of the wood. The cells of the paren- 
chyma, both of the medullary rays and of the wood, are stimu- 
lated to divide, and so form a connecting or intermediary tissue, 
which completely fills up the space between the two cut surfaces. 

If the operation has succeeded and the scion has grown, the 
latter is in future supplied with the raw food-materials that are 
absorbed from the soil by the roots of the stock. On the other 
hand, the plastic materials that are elaborated in the scion 
nourish the cambium both of the scion and the stock. Of 
course the new elements that are produced by the cambium 
cells of the scion are the characteristic elements of the scion, and 
similarly with regard to the elements produced by the cambium 
of the stock. The plastic materials produced in the scion afford 
assimilable nourishment to both scion and stock, just as cow's 
milk may serve as nourishment not onl}- to a calf but also to a 
child. But the latter does not on that account assume the pecu- 



liarities of a cow, nor docs the stock assume the pccuUaritics of 
the scion, although it is nourished b)- the metabolic products of 
the latter. If the cambium cells of the stock naturally divide 
more actively than those of the scion, the former will increase in 
thickness more rapidly than the latter, and vice versa. The line 
which marks externally the point of union between the fast- 
growing and slow-growing portions of the stem has been called 
by Goppert the "External line of demarcation," and is often 
recognizable by distinctions in the cortex and bark. Internally 
there is of course also 
a corresponding line of 
demarcation, along which 
the wood of the stock 
and scion unite, and 
which may often be re- 
cognized by a difference 
in the colour of the wood 
(Fig. 152). 

Many cases are known 
where it must be ad- 
mitted that the scion 
exerts an influence on 
the stock. It has been 
observed, for instance, 
that when a scion with 
variegated leaves has 
been used, variegated 
leaves have sometimes 
been produced on the 

shoots that have afterwards formed on the green-leafed stock. 
Such a case forces us to the conclusion that the plastic materials 
produced in the variegated leaves of the scion possess peculiar 
properties which act upon the cambium cells of the stock in 
such a way as to induce variegation in the leaves of the new 
shoots. It is not my intention now to discuss the still more 
potent influences which the scion has been known to exert in 
certain cases on the stock, merel}- remarking that hybrid forms 
have been obtained b}' grafting different varieties of potatoes on 
each other. 

Fig. 152. — Transverse section through the region 
where Sorhits Aria has been grafted on 
S. Aiiaiparia. The boundary line, a a, be- 
tween the slow-growing S. Aria and the fast- 
growing S. Auciiparia is known as the in- 
ternal line of demarcation. One fourth natural 



The effects on dicotyledonous trees of defoliation by insects 
depend on the season when it occurs. When the }^oung shoots 
as well as the leaves are destroyed in spring, fresh shoots are soon 
produced by the dormant eyes of the older branches, or by the 
buds that may have escaped at the base of the young shoots. 
Should defoliation occur in June or July, the trees reclothe 
themselves during August with leaves which spring from the 
buds of the defoliated shoot itself When defoliation takes 
place still later, fresh leaves are either not produced at all or 
only very sparingly. In the following year it usually happens 
that the effects of the defoliation are entirely obliterated. The 
wood-ring formed during the year of defoliation is narrow, and 
the growth of the succeeding year is usually also below the 
normal. The effects on the larch are similar to those met with 
in dicotyledons. In mixed woods of beeches and conifers, 
the former often suffer very severely from bark-scorching after 
the conifers have been stripped of their leaves. 

As compared with dicotyledons, evergreen conifers usually 
suffer very severely from defoliation, but in their case also much 
depends on the season of the year when the damage is done. 
Should this occur in spring, before the new shoots have been 
formed, or in autumn, after the wood-ring has been nearly 
or entirely completed, the life of the tree is not endangered. 
The new shoots, being prevented from deriving any nourishment 
from the leaves of the older branches, do not indeed develop so 
vigorously as those of trees that have not been defoliated, still a 
sufficient quantity of foliage is produced to enable such a tree to 
regain its normal condition in a few years. Fatal results, on the 
other hand, attend total defoliation at, or shortly after, the time 
when the new shoots are formed — that is to say, in May and 
June. On account of their tender condition, the young shoots 
are either totally destroyed or a portion persists long enough for 
the buds to develop. In any case there is a reserve supply of 
dormant buds, which the defoliation may stimulate to further 

^ R. W.zx\\^^ Das ErJo'iDiken iiiid Absterbeii der Fichte iiach der E7itnade- 
lung durch die Nonne. Forst. naturiuiss. ZeitscJirift. Nos. i, 2, 3, 7, jo. 


development, and some of which ma}- even produce short shoots. 
The trees are, however, unable to reclothe themselves with a 
permanent supply of leaves, for the following reasons. During 
the year in which defoliation takes place, all, or nearly all, the 
reserve supplies of plant-food that are stored up in the tree are 
made use of by the cambium in the formation of a wood-ring. 
No growth takes place in the following year. The reserve 
supplies in the young shoots also are quickly consumed by the 
cambium, so that the buds are prevented from forming shoots. 
In the course of the autumn and winter, especially during 
long-continued winter drought, all, or nearly all, the twigs and 
branches of the crown die for want of water. The bole of 
the tree maintains its vitality till the middle of summer — that is 
to say, for a full year after defoliation. During that period, 
however, no growth takes place. Then the inner cortex begins 
to become brown, and the bole dies. This is largely due to 
the high temperature of the tree, which is induced by the lack of 
shade in a wood that has been entirely defoliated by the ravages 
of insects. Especially is this the case with spruces, whose bark 
is but ill-adapted for mitigating the action of direct insolation. 

Should the defoliation not have been complete, the wood 
may slowly recover. This is most likely to occur when a 
mild wet winter follows the season in which the ravages were 
committed. New shoots are formed on those branches which 
have retained a large proportion of their leaves, and which have 
therefore been in a position to produce reserve materials 
even while the insects were at work, and to store them up 
for use during the succeeding season. The thin-barked spruce, 
however, is apt to succumb to bark-scorching, consequent on 
direct insolation. Trees which are to a certain extent protected 
against the action of the sun by the foliage of their neighbours 
may gradually recover, even after having lost a great deal 
of their foliage, but not after complete defoliation. 


Since science has recognized that the occurrence of all 
infectious diseases is perfectly independent of the chemical 
composition of the soil, that section of plant-pathology which 
deals with diseases induced by peculiarities of soil has been 
greatly restricted in extent. 


The supply of water and food-materials in the soil has a great 
influence on the rate of growth of a plant, although it is only 
in rare instances that it produces disease, in the restricted sense 
of the term explained at page 5. 

One such form of disease is the condition where the tree is 
said to become " stag-headed " or *' top-dry," * and which is usually 
to be traced to considerable diminution of the supplies of water 
or food-materials in the soil, and this prevents sufficient nourish- 
ment being continued to plants that have grown up under more 
favourable conditions. 

In beech woods this disease is specially liable to occur, in 
consequence of the removal of litter, and often appears as early 
as the pole-wood stage. The reduction in soil-fertility first 
makes itself noticeable in a general diminution of the rate of 
growth, though frequently also in the withering of the upper 
portion of the crowns, while the lower portion remains green. 

In alder woods top-drought follows excessive draining. 
When oaks that have grown up in a dense wood of beeches, 

*[i.e., the topmost branches become completely leafless, and die off, and 
remain as dry sticks, like antlers projecting above the foliage. — Ed.] 


and that have but poorl}- developed crowns in consequence, arc 
isolated by the removal of the beeches, they clothe their stems 
abundantly with epicormic branches. For some years these as 
well as the crowns thrive perfectly satisfactorily. In the process 
of time, however — and especially on the lighter classes of soil 
which are subject to rapid drought or are liable to produce weeds 
— ^a portion of the topmost branches of the crowns die, and the 
oaks become stag-headed. If the ground is protected in time, by 
under-planting, the top branches either do not die or the disease 
soon fails to make any progress, and the stag-headed condition 
ma}' entirely disappear owing to the dry branches dropping off. 

It is difficult to demonstrate the causes of these phenomena 
experimentally, but the following explanation may be accepted 
as sufficiently accounting for the disease. Directly the oaks are 
isolated the amount of soluble food-materials in the soil is 
augmented, owing to the accelerated decomposition of the humus 
that covers the ground, and, at the same time, the leaves of the 
crown, being more exposed to direct sunlight, are enabled to 
assimilate more rapidly. These two causes combine to produce 
considerable increase of the plastic substances, and consequently 
an increase in growth, and the dormant axillar}' buds are also 
enabled to develop into shoots. 

The first impulse to activit}' is probabl}- communicated to the 
dormant eyes by the increase in the products of metabolism, 
while their further development into shoots is rendered possible 
by the intensified action of the light. When the crowns and 
branches have grown vigorously for a few }'ears, the stock of 
humus becomes exhausted, while the soil dries up in summer to 
considerable depths, owing to the upper layers being deprived of 
their protective covering. The result is that the processes by 
which plant-food is rendered available are interfered with, and the 
stock of soluble food-materials in the soil is reduced. Such a 
state of things is commonly expressed by saying that the ground 
has " become wild." 

The }'ears when plant-food is abundant are followed by a 
period of famine. Owing to the reduced supply of water and 
nutriment, the upper part of the crowns is starved, the lower 
branches appropriating the whole of the water and plant-food. 

Provided the crowns have not been too severely crippled, they 


may recover under the influence of an increased supply of food- 
materials, consequent on the improv^ement of the soil by under- 
planting. Trees that were possessed of well-developed crowns 
before the wood was light-thinned produce but few, if any, 
epicormic branches, nor do they become stag-headed. The 
reason for this is that, owing to vigorous development, the 
crowns are able to make use of the excess of nutriment that 
is produced during the years immediately following the light- 
thinning. No epicormic shoots are produced, so that there are 
none to interfere with the nourishment of the crowns during the 
years of famine. No doubt the general health of the crowns 
suffers, but at all events their upper branches do not die. 

It follows from what has been said that if top-drought is to be 
avoided there must be no temporary reduction of soil-fertility. 
The discovery of the means by which the soil may be protected 
and its fertility conserved falls within the province of sylviculture. 

In the case of agricultural plants we are familiar with a 
number of pathological phenomena which are primarily due to 
the effects of drought on the soil. Here I will only mention 
the " going off " of cereals — namely, the withering of the straw 
before the grain has formed — and the premature ripening of grain, 
where the plants wither after the seeds have formed, but before 
all the nutritive materials have been stored up in the grain. 

Under exceptional circumstances it may also happen that the 
growth of plants is interfered with by excess of nutriment. I 
would, however, again utter a word of warning against hastily 
ascribing sickly appearances to the soil, in the absence of 
scientific evidence. A sudden increase in the supply of plant-food, 
and the consequent important augmentation of the plastic sub- 
stances, may, under certain circumstances, cause the outer tissues 
to rupture, and this occurs when their extension has been unable to 
keep pace with the growth of the internal tissues. It occasionally 
happens, when some cultural operation has suddenly induced 
considerable increase of growth in trees, that the bark, especially 
on the main stem, is ruptured on all sides owing to the powerful 
internal pressure that is set up. When hornbeams ^ that were 
mixed with beeches in a wood were suddenly isolated in the 
seed-felling, their annual sectional growth at breast-height 
1 Untersitchungen aiis dent Forstbot. Inst., \q\. 1 1 1, pp. 141 — 144. 



increased in a few years from O'l 86 sq. in. to 2-124 sq. in., 
and even more. This caused such high tension in the outer 
periderm that longitudinal rupturing was finally induced at 
numerous points. Owing 
to subsequent shrinkage the 
fissures extended to the wood 
(Fig. 153, (?:), and it some- 
times happened that the 
whole of the cortical tissues 
became detached from the 

wood along the cambium region for some distance on each side 
of the fissure (Fig. 153,1!^). The consequence was that the whole 
of the cortex warped like a board that has been dried on one 
side. Most of the numerous wounds healed very quickly in 

Fig. 153. — Diagrammatic representation of 
two ways in which the cortex may rupture 
when the rate of growth is suddenly 

Fig. 154. — Transverse section of the stem of a hornbeam whose cortex had been 
ruptured in 1876 owing to sudden acceleration in the rate of growth, a, a fissure 
in the cortex which does not extend to the wood ; b, an occluded fissure ; (-, a 
fissure which has not yet been completely occluded. The figures correspond to 
the annual rings, these being very narrow in the years 1861 — 1871. One half 
natural size. 

about a year, though some not till later (Fig. 1 54), but for a 
long time the cortex of such hornbeams exhibited an unusual 
appearance (Fig. 155)- 

I have frequently observed the cortex of oaks ^ to be similarly 

1 Op. elf., vol. i. pp. 145—150. 



ruptured, when trees that have grown for a long time in a wood, 
overcrowded owing to neglect in thinning, have been suddenly 
isolated, or when trees that have been reared in restricted light 
have been suddenly exposed by the removal of the standards. 

The augmented supply af 
food-materials in the soil, 
and the intensified action 
of the light, resulted in 
such an acceleration of 
growth that fissures of 
various sizes were formed 
all over the stem. Fig. 
156 represents the trans- 
verse section of such an 
oak a hundred years 
old, and exhibits the 
interesting manner in 
\\hich new tissues are 
produced as a result of 
the formation of the fis- 
sures. These wounds are 
injurious, not only be- 
cause the resulting cica- 
trization and formation 
of callus interferes with 
the splitting of the wood, 
but also because they 
offer a means of ingress 
to parasitic wood--de- 

FiG. 155.— Hornbeam whose cortex has been Stroying fungi. They 

ruptured, a, a crack which does not extend almo'^1- alwavs he 

to the wood ; fi, a fissure reaching to the wood, ^^Y almost al\\ ays DC 

but which has been occluded by the formation avoided by Strongly thin- 

of callus (see Fig. 154, d); c, a crack which . , . . 

extends to the wood only in the upper portion. ^rig tne plantation SOmC 

One half natural size. years before it is in- 

tended to " lighten " it. 

It being taken for granted that roots rot and the whole plant 
dies when excess of stagnant water in the soil prevents the 
entrance of air to the roots, and, further, that the same cause 
induces the formation of injurious humic acids, increases the 



danger of frost in the case of man>' plants, and conduces to 
seedlings being thrown out by frost, &c., the subject need 
not be further discussed. 


Fig. 156.— Transverse section of an oak which in consequence of much accelerated 
growth has ruptured in two places, x and j', two years before being felled At 
the three places marked a b the cambium has occluded the surface of the wood 
with new tissues, which are possessed of an independent cortex, d d. The loose 
flaps of cortex have TiX e e formed new wood on their inner surface. This has 
formed a kind of callus-cushion at c, which constitutes the edge of the wound. 
The wood-nng formed underneath the cortex in 1876, the year in which the 
cortex was ruptured, is a sort of double ring, and consists of two parts,/ and ^^ 
both of which contain a porous zone and a zone where vessels are comparativeW 
scarce. The porous zone of the inner of these two pr.rts, namely/ was formed 
m the spring before rupturing had taken place. 


The metabolic processes in the roots demand an abundant 
supply of oxygen. The roots die owing to asphyxia if 
they are excluded from a constant supply of this element. 
Oxygen is necessary not only for growth but also for the 
formation and solution of reserve materials, processes which 
are specially active in roots. The air in the soil is impoverished 
to an extent corresponding to the amount of oxygen thus 
abstracted. Under normal conditions the loss is abundantly 
compensated for, partly by the variations of temperature in the 

^ R. Hartig, Zersetziaigserschcinungen, pp. 75 ^/ seq. 

T 2 


superficial layers of the soil, partly by the processes of diffusion, 
and partly by the entrance of water containing dissolved oxygen. 
The greater the daily and annual variations of temperature in the 
upper layers of the soil, and the greater the depth at which they 
operate, so much the more thorough is the interchange of gases, 
or, as it is sometimes called, the " respiration " of the soil. As 
is well known, the temperature of the soil depends, in a great 
measure, on its specific heat. The lower the specific heat, so 
much the more quickly is the soil heated or cooled. Water and 
humus possess a high specific heat, and the more of these sub- 
stances a soil contains the greater will be the quantity of heat 
required in order to raise its temperature. A forest soil that is 
unprotected by umbrage, that is easily dried owing to exposure, 
and that has lost the greater part of its humus, is much more 
easily warmed than a soil that is protected by a dense wood, 
is constantly moist, and contains abundance of humus. 

It is further evident that a forest soil which is exposed to 
direct insolation is much more easily warmed, though it also cools 
much more easily owing to radiation of heat, than one where 
the crowns of the trees and a covering of leaves and humus 
afford a double protection. 

So far as the diffusion of air in the soil is concerned, we know 
that it only occurs to a considerable extent in porous soil 
which is not over-wet. In the case of dense, firm wet soil the 
mixing of gases proceeds with extreme slowness. It may 
happen, under certain circumstances, that the interchange of 
gases in the soil is so limited as to induce asphyxia and deca}' 
in the roots of plants. I have applied the term "Root-Rot" to 
cases where the roots die by asphyxia, in contradistinction to 
infectious root-diseases. 


This disease is specially destructive in the young Scotch pine 
woods of the north of Germany. It seldom appears before the 
twentieth year, usually not till the thirtieth, and is characterized 
by the trees appearing unhealthy for a short time and then 
falling over while still perfectly green, after snow or a strong 
gale has supplied an external impulse. The tap-root will be 
^ Op. cit., pp. 74 ct scq. 


found to be wet and rotten almost back to the stool, while on the 
other hand all or most of the shallow lateral roots remain per- 
fectly sound. Only in rare cases is the withering of the tree due 
to saturation of the stool with resin, consequent on the decay of 
the tap-root. Root-rot is to be distinguished from the ravages 
of Tranietes radicipcrda — both being frequently met together 
in pine woods — by the tap-root rotting and the lateral roots 
remaining sound, whereas in the case of the parasite the tree 
is killed, though not thrown, owing to the lateral roots being 

The disease is also to be met with in spruce woods growing 
on decidedly shallow soils that contain stagnant water. Under 
such circumstances, however, it is less destructive, because the 
shallower root-system of the spruce makes the tree more 
independent of the decay of the few roots that penetrate deep 
into the soil. 

In the case of pine woods, root-rot appears only on soils 
where, at a short distance from the surface (usually about \\ ft.), a 
stratum is encountered which offers no obstacle to the entrance 
of the tap-root when the trees are young, but which is of such a 
texture as to prevent the free circulation of air after the wood 
has become close. This stratum, which usually consists of 
argillaceous loam or of very fine-grained quartz (alluvial loam), 
is so difficult to work with the spade as to necessitate the use of 
the pick. As such conditions of soil are also unsuitable for 
agricultural purposes, we very frequently find such strata where 
farming has been replaced by forestry. For this reason the 
subsequent disease of the pines has been erroneously ascribed 
to the previous tillage operations. At first young pine woods 
thrive admirably on such soils. The tap-roots penetrate to the 
deeper layers of the soil, to which at first the circulation of air 
also extends. It is only when the branches begin to interlace 
and to form a dense umbrageous canopy which protects the 
soil summer and winter, and when a thick layer of leaves and 
humus forms on the ground, that the circulation of air in the 
soil is interfered with. Insolation becomes impossible, and both 
heating and cooling are rendered alike difficult. As the soil 
remains constantly moist, while the air is largely excluded from 
soil that is argillaceous and very impervious, or consists of dense 


sand, the processes of diffusion proceed but slowly. Although it 
may not be for some decades, this interference with the air- 
circulation may ultimately induce asphyxiation of the deeper 
roots, by preventing their obtaining a sufficient supply of 

Root- rot never occurs in dicotyledonous trees, and only with 
extreme rarity in pines that are mixed with dicotyledons. 
Possibly this may be explained by the fact that during half of 
the year the soil is subjected to the minimum of umbrageous 
shelter, and consequently there is more air-circulation than in a 
wood composed entirely of conifers. 

This brings us to the immediate consideration of the best 
means of prevention. These must always be directed towards 
securing better aeration of the soil. The circumstances of any 
particular case must determine which of the following courses 
is to be taken : cultivating mixed woods of dicotyledons 
and conifers, or, should this be impracticable, the pine may 
be replaced by the shallow-rooted spruce ; the removal of 
excessively large accumulations of leaves in hollows ; or the 
abstraction from the soil of stagnant water by drainage. 

The death of the deeper roots on trees that have been 
too deeply planted may to a certain extent be described as a 
variety of root-rot. The heavier the soil, so much the more 
dangerous is it to plant deeply. It is best that such a tree 
should succumb at once, but in most cases it lingers through 
several decades without being able to produce new roots to 
replace those that have rotted off. Only a few trees, such as 
willows, poplars, and especially shrubs, develop a plentiful supply 
of adventitious roots immediately beneath the surface of the 
ground, by means of which a new root system is formed, as in 
the case of cuttings which are absolutely destitute of roots. 

Similar conditions are induced when the roots of older trees 
are covered with a thick layer of earth, as often occurs during 
the operations attending road-making, mining, &c. In such cases 
less damage is done if the air can get at the roots from the side, 
as usually happens when trees grow on sloping ground, but if 
the entrance of air to the roots is rendered a matter of great 
difficulty the trees either die off entirely, or at all events their 
growth is seriously impaired. I found close beneath the surface 


of the earth-heap that adventitious roots were being abundantl}' 
produced from the uninjured cortex of smooth-barked trees, such 
as the beech and hornbeam, even when the stems were eight 
inches in diameter. 

Where it has been deemed desirable to preserve valuable trees, 
excellent results are said to have been got by ringing the stems 
a short distance beneath the surface of the ground, or at least 
by removing the bark in patches as far in as the wood. From 
the callus that formed at these places numerous roots were 
produced, which by ramifying close beneath the new surface of 
the heaped-up soil preserved the life of the tree. 

It is scarcely necessary to mention that failure in the natural 
regeneration of beech woods is often to be traced to the 
insufficient aeration of soil that is covered with a thick layer 
of humus. Small seeds especially, that are buried too deepl}\ 
frequently fail to produce plants on account of the supply of air 
being insufficient to replace the carbonic acid gas that is pro- 
duced during germination. The familiar fact that unsatisfactory 
results are almost always got when the germination of alder- 
and birch-seeds is tested in a room, although these seeds ger- 
minate splendidly when they are sown outside, is probably due 
to the circumstance that it is only outside that the air in the 
neighbourhood of the seeds is constantly being changed, owing 
to the daily variations in the temperature of the soil. In the 
room the temperature is uniform and the air is comparatively 
still, so that the carbonic acid gas which is given off during 
germination cannot be removed quickly enough from the 
neighbourhood of the seed. Death occurs in heaps of germinat- 
ing seeds for similar reasons. 

Analogous to the root-rot that has already been described 
is also the decay of the roots of plants that are culti- 
vated in glazed pots, which render the free circulation of air 


In the narrower sense of the term poisons are taken to mean 
only such substances as are directly injurious to plant-cells and 
effect their destruction. Such substances may be naturally pre- 
sent in the soil, but are more often imported into it. As a rule. 


the meaning is extended to include innocuous soluble substances 
— -which may even be valuable constituents of plant-food — when 
the solutions in which they are present in the soil are in a too 
concentrated form. The endosmotic process by which the roots 
take in water can go on only if the cell-sap of the roots is 
so much more concentrated than the solutions in the soil that 
it can absorb water >rom the environment. On this account 
any strong solution of food-materials in the soil will prove 
injurious, and may even attract water from the roots. The 
result is that the plants wither. . Such a state of things may 
be frequently observed when very soluble mineral manures are 
applied in excessively large quantities. Other soluble salts which 
are innocuous in themselves may also cause plants to wither. 

When spring tides have inundated woods situated behind 
dunes, the water, being unable to return, has slowly percolated 
into the soil, and the chloride of sodium which sea-water 
contains has frequently proved extremely injurious.^ The pine, 
alder, oak, and beech succumbed altogether and were found to 
suffer most, while the birch was least affected. In July 1874, 
along with Herr Schiitze, the chemist at Eberswalde, I insti- 
tuted investigations on the action of common salt, using solu- 
tions of the strength of the water of the Baltic (27 per cent.) 
and of the North Sea (3'47 per cent.) The Scotch pine, spruce, 
false acacia, and beech were selected for experiment, beds of 
seedlings and transplanted trees being sprayed with the salt 
water, each square yard receiving 2-57 gallons at a time. One- 
and three-year-old spruces succumbed both to the weak and 
the strong solutions, while six-year-old plants were only killed 
by the stronger solution, though they became partially brown 
under the action of the other. When spruces some six feet 
high each received fully three gallons of the stronger solution, 
some were killed, while others showed only a temporary brown- 
ness and ultimately recovered. False acacias one year old were 
also killed by the weaker solution, while, strange to say, in the 
case of thirty-year-old beeches it was only the points of the 
leaves that died, some time after the solution had been applied. 

^ Schiitze, " Untersuchung von Boden und Holz aus Bestiinden, welche 
durch Sturmfluthen der Ostsee beschadigt sind," Zeitschrift fur Forst- 
und Jagdwesen, 1876, p. 380. 


In this experiment the Scotch pine proved least sensitive, a 
result which was possibly due to its deeper roots. 

The injurious effects of urine oh plants are generally well 
known, and may be sufficiently explained from its saline 

Many acids and leys act as true poisons, and are sometimes 
conveyed to the soil in large quantities in the impure water that 
flows from factories. As experience proves, they are highly 
injurious, but this is not the place to discuss the many poisons 
that may occur in such contaminated water. 

A certain amount of interest also attaches to the injurious 
influences exerted on vegetation by continuous exhalations of 
carbonic acid gas from the soil. At the baths of Cudowa in 
Silesia many springs of water containing carbonic acid are 
distributed throughout the park. At such places one finds 
only grass, shrubs being unable to grow. This is probably due 
to the soil being so permeated by free carbonic acid that the 
respiratory processes of the roots are rendered impossible. Grass, 
however, is enabled to grow because the circulation of air close 
beneath the surface of the ground is sufficient to maintain the 
roots alive. 

It has been proved that the roots of trees are injured by 
coal gas when it escapes from pipes into the soil in large 
quantity. The unhealthy condition or death of trees that line 
the streets of towns is, however, not altogether to be attributed to 
this form of injury. The cause is rather to be found in the close 
paving of the streets and footpaths, which precludes the entrance 
of water and even air, so that the tree-roots suffer both from 
want of moisture and of air. 

It may be shortly mentioned here that coal gas also interferes 
considerably with the cultivation of flowers in rooms. This is the 
case even when but little gas is burned, for small quantities are 
always escaping from the pipes. Camellias, azaleas, and ivy are 
very sensitive to gas, the least sensitive plants being palms and 

* [In many cases, at least, these injuries are due to the sulphurous anhy- 
dride of which traces are frequently present. — Ed.] 




The action of frost on plants, whether fatal or otherwise, can 
be understood only when one has gained a clear idea of the 
sources of heat of which plants can avail themselves. 

The metabolic processes which make the more highly 
developed animals independent to a greater or less degree of the 
influences of external heat constitute a factor in the vegetable 
kingdom which may be neglected, in comparison with the effects 
exerted on plants by the heat of the surrounding media. In 
the case of the older classes of trees, especially those which are 
covered by thick bark, the temperature of the lower and inner 
portions of the tree is chiefly determined by that of the soil. 
The temperature of the surrounding air has, however, most 
influence on branches and twigs. 

At the time of active growth, and in fact whenever trans- 
piration of water is proceeding energetically, the temperature 
of the interior of a plant is brought into conformity with 
that which prevails in the soil by means of the water that 
is absorbed by the roots. This has been placed beyond 
the shadow of a doubt by the following experiment. Two 
trees alike in all respects and equally exposed to the sun 
were selected, of which one was deprived of its branches. It 
was then found that the temperature of the tree that had been 
left intact was 1 8° F. lower than that of the tree which had been 
pruned. When the former was also pruned, and the ascent of 


water consequcntl}' stopped, the temperature at once rose 18° F. 
When the soil is frozen so that no water can enter by the 
roots, the tree receives heat from the soil only by the process 
of direct conduction. This, however, is always of sufficient im- 
portance to explain why the temperature of the interior of 
a tree, even during prolonged cold, rises as we descend ; and 
also why a deep soil, in which the roots descend to long dis- 
tances, has a more favourable thermic effect on trees than a 
shallow soil. This also explains why a natural or artificial 
covering on the soil is so useful in enabling fruit and ornamental 
trees to resist the winter's cold. The reason also why certain 
trees that are easily frosted when young are apparently less 
sensitive to cold in later life — or become " hardened," as it 
is called — is to be traced to the greater amount of heat which 
the roots receive when they have penetrated to greater depths. 

The extraordinary rapidity with which shrubs and trees 
become green in spring after a heavy shower of warm rain is 
also due to the rise in temperature of the soil. Finally, the 
early appearance of leaves on the smaller classes of trees in a 
wood, as compared with the larger trees, is due to the fact that 
the soil-strata in which the roots of the former are chiefly 
distributed experience a rise in temperature at a time when the 
cold of winter still prevails in the deeper strata, and it is from 
the latter that the stronger and more vigorously developed roots 
derive their heat. 

It is the temperature of the surrounding air that chiefly 
determines the temperature of twigs and branches, as well 
as of all the more delicate parts of plants generally. Heat 
penetrates with extreme slowness into the interior of those 
portions of a stem which are covered with a very thick periderm 
or a layer of bark. It is only when insolation is uninterrupted 
that the side of a tree which is exposed to the sun's rays may 
become heated to such a pitch as to induce such pathological 
phenomena as "Bark-scorching" and "Sun-cracks." As op- 
posed to the heat which plants receive, we have the loss of heat 
which they experience. Owing to the evaporation of water, heat 
is directly abstracted from the tissues where this process is active- 
The process of assimilation is also connected with loss of heat. 

The rate of cooling is, however, most influenced by radia- 


tion of heat. This proceeds most energetically in the more 
divided up parts of plants, where the surface is large in 
proportion to the mass of the organ. The depression of tem- 
perature consequent on radiation of heat not only explains the 
phenomena of hoar-frost, dew, &c., but is also in most cases 
accountable for late frosts which not unfrequently occur during 
still clear weather, even when the temperature of the air is 
above the freezing-point. From what has been said it is 
sufficiently evident that the readings got from thermometers 
inserted in holes of different trees are the result of the joint 
action of various heat-producing and cold-inducing factors. The 
determination of the internal temperature of trees at the Forestal 
Meteorological Research Stations has absolutely no scientific 
value, and represents a waste of time on the part of the observer 
that is quite unjustifiable. 

When the temperature of any portion of a plant sinks below 
the minimum necessary for the production and continuance of 
the chemical processes of metabolism — that is to say, for the 
calling into action of the vital forces — a period of rest ensues, 
which continues until the necessary thermal conditions are again 
restored in the tissues. Should the temperature sink considerably 
below 32° P., the plant is frosted ; in other words, a portion of the 
water of imbibition in the cell-walls and a portion of the water 
of the cell-sap separates in the form of ice crystals, while a 
more concentrated solution with a lower freezing-point remains 
behind in the liquid form. 

In the wood of a tree, where for the most part intercellular 
spaces are absent, the water of the cell-walls can only separate 
out to form ice crystals in the lumina of the cells, while the 
walls themselves become drier but do not freeze. As the lumina 
of the wood-cells contain abundance of air besides water, there 
is ample space to admit of the expansion which the water 
undergoes in changing into ice. The lower the temperature 
sinks, so much the more water leaves the walls, and so much 
the drier do they become. This explains why trees shrink in 
exactly the same way during intense cold as felled timber does 
on drying. The volume of the cell-walls is reduced proportion- 
ally to the water that is withdrawn, and the stem ruptures 
longitudinally and displays frost-cracks or frost-fissures. These 


are most abundant on the north-east side of trees, because intense 
cold usually occurs with a north-east wind. As a rule, frost- 
cracks are formed only when a great reduction of temperature 
occurs suddenly, and when the interior of the tree is therefore 
relatively warm, so that excessive shrinkage is confined to the 
outer layers of the wood. 

It is a familiar fact that, when such frost-cracks have closed up 
with the restoration of a higher temperature, they become oc- 
cluded by the callus that forms along their edges. The reduced 
pressure of the bark 
causes the forma- 
tion of new tissues 
along both sides of 
the crack, and these 
project from the 
surface as a " frost- 
rib." On account 
of the thin callus- 
layers being easily 
ruptured, it requires 
but a few degrees 
of frost in succeed- 
ing years to re- 
open the crack. 
Repeated opening 
and closing of the 
wound sometimes 
induces the forma- 
tion of strikingly 

prominent frost-ribs. Should several mild winters occur in 
succession, a frost-crack may close up entireh', as is seen in 

Fig- 157- 

In the interior of old oaks I have sometimes noticed numerous 
radial and peripheral cracks which did not extend to the outside 
of the stem, nor had they reached the surface even at the time 
when they were formed. At present it is uncertain whether 
these cracks are also to be attributed to the action of frost, and 
no satisfactory explanation has been given as to the circum- 
stances under which they originated. 

Fig. 157. — An oak-stem showing a frost-crack which has 
been produced in the winter before the wood-ring, 
a, was formed. Originally the crack extended from 
n to d. For nine years in succession the crack has 
been annually reopened, so that the frost- rib, a to b, 
has been formed, and this has ruptured laterally 
at c. During the last five years the crack has re- 
mained closed. One half natural size. 


When the tissues of the leaves and cortex, and in fact when 
any parenchymatous tissues are frosted, pure water is withdrawn 
into the adjoining intercellular spaces, but the cells themselves 
do not generally freeze. The result is that the cells lose their 
turgidity, and at the same time begin to droop. This explains 
the familiar phenomenon of lilies, hyacinths, &c., which have been 
caught by late frost, being prostrated on the ground, until the 
ice melts and the cells reabsorb the water into their interior 
and again become turgid, when the plants resume an erect 

Cells which contain a concentrated solution part with water 
only under the influence of very intense cold, and I have often 
found that the cortex and bast of trees showed no signs of ice 
when the wood was hard frozen. 

As a rule, when living plant-tissues that contain much water are 
frosted — and this applies especially to young leaves and shoots 
that are affected by late frost — large masses of ice are formed in 
certain regions, and notably underneath the epidermis of leaves 
and shoots, and in the medulla. The tissues, however, remain 
entirely free of ice, merely shrinking in proportion to the 
quantity of water that is lost. These masses of ice consist of 
parallel prismatic crystals, which are arranged at right angles 
to the tissues from which the water has been abstracted. The 
cortical parenchyma of the shoot usually contains numerous 
intercellular spaces, especially along the line that marks the 
limits of the collenchymatous tissues of the outer cortex. 
Owing to the formation of a sheet of ice in this region, a 
separation of the cortical tissues may take place, which however 
may occasion but little damage to the plant. I have noticed 
after a late frost that the epidermis on the under side of the 
leaves of the sycamore was pushed out into numerous vesicular 
swellings, but it was only after the lapse of several weeks that 
this forcible separation exercised any prejudicial influence on the 
health of the leaves. 

On account of its numerous large intercellular spaces, it is 
evident that the spongy parenchyma of the under part of the 
leaf offers specially favourable conditions for the formation of ice. 
In the case of the false acacia and other trees that are still green 
when the first frost occurs in autumn, a sheet of ice forms in the 


layer of cells that has previously been formed across the petiole 
of the leaf. Simultaneously with the formation of the ice the 
connection of the leaf with the tree is severed, the result being 
that on the following morning there is a general fall of leaves. 

When a thaw occurs in the frosted parts of a plant, the tissues 
usually regain the condition which characterized them before the 
frost appeared. As the water is set free by the melting of the 
ice it is slowly absorbed by the cell-walls and the cell-contents. 
In many cases, however, it is found that the parts have been 
killed. Instead of the chemical processes that are revived under 
the action of a recurrence of heat inducing normal metabolism, 
they initiate chemical decomposition. The views are divided 
as to the time when frost proves fatal. While Goppert con- 
cludes that death occurs during the continuance of the frost, 
Sachs is of the opinion that the tissues die only after they 
have thawed, and that a fatal issue depends very much on 
the manner and rate of thawing. The two views may to 
a certain extent be reconciled, for it is possible that during 
winter death occurs during the continuance of the frost, whereas 
in the case of a late spring frost it appears at the moment 
of thawing. 

The death of a plant under the action of frost during winter 
bears a close resemblance to the effects of drought on the 
tissues. No matter whether the deficiency of water in the 
tissues is due to the action of frost, or to evaporation being in 
excess of the absorption of water by the roots, the cells must die 
if the deficiency exceeds a certain limit. A change is induced 
in the molecular constitution of the protoplasm, the main 
feature of which is that the protoplasm is rendered incapable 
of retaining any considerable quantity of water. This change is 
probably connected with the dissociation of molecular groups 
in the protoplasm in consequence of the abstraction of water. 
In a living condition the micellse * of the protoplasm are 
surrounded by water, the water and the micellae being held 
together by that kind of molecular attraction whose action in an 
organic substance is spoken of as the force of imbibition. It 

*[The hypothetical structural units of an organized body have been 
termed, among other names, micellse : each micella is supposed to have its 
own molecular structure also, much as a crystal has. — Ed.] 


may be assumed, although it cannot be demonstrated, that the 
arrangement or grouping of the ultimate particles of the 
protoplasm suffers a change during excessive abstraction of 
water, and that, when the supply of water is again restored, they 
are unable to regain their original position. Should the critical 
limit of drought not be overstepped, the cell passes from the 
condition of plasmolysis * into that of turgescence ; but, on the 
other hand, a cell withers and is unable to regain its normal 
vital condition if the limit of drought has been exceeded. The 
same holds true when the loss of water is induced by frost. A 
cell is able to bear a certain amount of cold with impunity, the 
molecular derangement that causes the death of the plant — that 
is to say, the changes in the normal properties of the protoplasm 
— occurring only when the loss of water due to the action of frost 
or drought has exceeded a definite limit. 

In order to illustrate the molecular derangement of the 
protoplasm, reference may be made to the familiar changes 
that occur in starch-paste under the action of frost. When 
that substance freezes it parts with more or less of its water, 
and the comparatively dry residue suffers a molecular change 
which prevents its reabsorbing as much water as it originally 
possessed. When the thaw occurs, the clear water remains 
outside the disorganized ■ paste, which consequently loses its 
glutinous character. 

In the condition of vegetative inactivity our perennial plants 
are capable of withstanding our coldest winters without perishing 
from frost. In other words, our winters are never so cold that 
our forest trees succumb to a molecular disorganization of the 
protoplasm of the cells. On the other hand, trees that have 
been introduced from warmer countries — and these include most 
of our fruit trees — perish from frost during unusually severe 
winters. The winter 1879-80 furnished a lamentable instance 
of this fact. Exotic plants exhibit every degree of hardiness, 
down to the point which is reached even in our mildest winters, 
and which precludes the possibility of their passing the winter 
out of doors. Apart from specific peculiarities, we also find 

* [Plasmolysis is a condition of collapse of the living contents of the cell, 
so that water escapes : the cell cannot grow until it is again distended with 
water (turgescent). — Ed.] 


individual differences, and it is this fact that makes it possible 
for us to acclimatize plants. As the capacity to resist frost 
varies amongst individuals of the same species, just like any 
other physiological or morphological pcculiarit}% it becomes 
possible to acclimatize a tender plant by propagating hardy 
varieties. It is also probable that hardier varieties are produced 
in the struggle for existence that takes place along the line 
which limits the natural geographical distribution of a plant, 
where the increasing severity of the climate bars the way to 
a further advance. From this it follows that in attempting to 
introduce a certain species it must be advantageous to procure 
the seeds from such frontier regions. 

Indigenous shrubs and forest trees suffer from winter frost 
only under very exceptional circumstances. The roots of young 
trees, more especially oaks up to four years old, may be killed if 
severe and long-continued frost finds the lighter classes of soil 
unprotected by snow or any other covering. The periderm on 
roots is thinner than that on stems, and consequently the former 
are less protected and moreover growth is active for a longer 
period in roots, where it frequently continues till the middle of 
winter, so that when frost occurs the tissues arc not in the inert 
condition which assists them to resist cold. Such plants burst 
their buds in spring, but wither up whenever transpiration 
from the delicate young shoots has exhausted the stock of 

Shoots that have not completed their growth, especially the 
Lammas shoots of the oak, suffer from winter frost. This is a 
matter, however, that belongs to the seconci division of our 
subject, which treats of the phenomena induced by frost in 
plants that are affected while in a state of vegetative activity. 

Even our indigenous trees, more especially evergreen di- 
cotyledons and conifers, may succumb during winter owing to 
their supplies of water being abstracted not by cold but by 
transpiration.^ The absorption of water by the roots ceases 
when the ground is frozen to a depth that is reached by the 
roots of young plants. No harm is done if the trees are 
protected above-ground against evaporation, by snow or any 
other covering. They die, however, if they are exposed for 
^ R. Hartig, Untersuchtmgen, I. p. 133. 



months to the action of air and sun, as was the case, for instance, 
in the winter of 1879-80. In this case drought alone was 
accountable for death. Even in the course of the winter 1879-80 
the leaves of middle-aged spruces and silver firs became brown 
and died where the foliage was exposed to the direct rays 
of the sun, and where constant air-currents encouraged trans- 
piration, as, for instance, on the southern edges of woods, on 
railway embankments, or on spruce hedges, &c. It was said 
that in Alpine regions which were much exposed to the south 
wind even old woods of silver firs succumbed entirely to the 
influence of the frost. In my opinion these phenomena can 
only be explained by the circumstance that repeated thawing and 
accelerated transpiration are induced in the leaves by the direct 
action of the sun during the bright wintry weather that usually 
prevails in these parts, or by the warm south winds, as the 
case may be, and that the leaves wither because they are unable 
to obtain any water from the stems which have been frozen 
under the influence of long-continued and severe cold. Man}- 
of the phenomena accompan}-ing the defoliation of pines, as well 
as the death of the branches of old pines, may also be explained 
in this wa}'. The injurious effects of repeated thawing and 
freezing, long-continued frost, or strong drying winds are to be 
explained by the scarcity of water that results from the inter- 
rupted or at least reduced passsage of water. 

The limits of forest growth in northern latitudes and in 
mountainous regions are determined not so much by the low 
temperature as by the action of drought on those parts of the 
tree that project from the snow during the long period of 
vegetative inactivity. On this account, too, we find that the 
limits of tree-growth are reached at a considerably lower 
elevation on south and west slopes, where the action of the sun, 
augmented as it is by reflection from the snow, is stronger than 
on north and east slopes. 

We are still awaiting a satisfactory explanation of the 
familiar fact that trees, especially exotic conifers, are more easily 
killed by frost in a wet situation than in a dry one, and that in 
general the more succulent parts of plants are more liable to suc- 
cumb to frost than those portions which are comparatively dry. 

When trees have suffered from frost during winter the 


injurious effects manifest themselves in a variet)' of wa}'s which 
have not hitherto been sufficiently investigated. After ver\' 
severe and long-continued winter cold, the cortex, bast, and 
cambium, and the wood-parenchyma as well, die and become 
brown. The trees either fail to produce leaves in the following 
season, or if they do bear leaves, flowers, and even fruit the\' 
wither up entirely in the course of the summer or autumn. As 
the wood does not lose its power of conducting water all at once, 
trees that are injured by frost may be able to produce leaves. 
The power, however, disappears as decomposition spreads from 
the parenchymatous cells to the conducting organs, or as the 
wood dries up from without inwards. Sometimes the cortex 
and bast are only killed in patches, and when this is the case a 
callus may gradually form over the damaged parts. 

It sometimes happens, especially in the case of exotic conifers, 
occasionally also in dicotyledons, that the cortex, bast, cambium, 
and frequently also the youngest annual wood-rings exhibit 
immunity from frost ; the wood-parenchyma, especially that in 
the neighbourhood of the medulla, being alone destroyed. In 
such a case conifers usually die suddenly from drought in the 
beginning of Ma}- ; whereas dicotyledons, whose cambium 
becomes active during the bursting of the buds, frequently 
remain alive. This result is due to the fact that the cambium 
having remained unaffected, forms a new wood-ring before the 
old frosted wood has lost its power of conducting sap ; or else 
the youngest annual rings escape the frost and suffice for the 
transference of the sap. Although the shoots and leaves are but 
poorly nourished for some years after the occurrence of the frost, 
such trees ultimately recover. Under such circumstances it 
often proves an excellent plan to prune severely, so as to bring 
evaporation into proportion with the diminished quantity of 
water that finds a passage through the wood. In very dr}' 
years, however, many trees ultimately succumb to the after- 
effects of the frost. 

When frost affects plants during the season of growth — and 
this is the case with late and early frosts — a fatal issue no longer 
depends on the hardiness of the plant, but upon the manner of 
thawing. When in a state of vegetative inactivity, our indigenous 
trees can withstand the most severe cold of winter with 

U 2 


impunity, whereas if the leaves have appeared they suffer from 
a few degrees of frost. In this case the view is undoubtedly 
correct that death from frost only occurs with the thaw. When 
plant-tissue is frozen during active growth, it exhibits the 
conditions that have already been described. Should the plant 
thaw very gradually, the water is absorbed by the walls and 
contents of the cells at the same rate as it is formed from the 
ice-crystals by the gradual accession of heat, so that when the 
cells have attained the temperature at which chemical processes 
are possible the normal conditions of imbibition have also 
been again restored, and the metabolic processes which were 
temporarily suspended are resumed under the influence of the 
higher temperature. The case is different, however, when the 
frosted parts of plants are rapidly thawed, as occurs, for instance, 
when they are brought into a warm room, or are touched by the 
warm hand, or are suddenly warmed by the sun. The rapid 
accession of heat induces the ice in the intercellular spaces to 
thaw rapidly, and the ice-water, being but slowly absorbed by 
the cell-walls and protoplasm, flows into the intercellular spaces, 
and drives out the air, with the result that leaves which are 
suddenly thawed become translucent. The normal conditions of 
imbibition have not been restored when the chemical processes 
start afresh under the influence of the rise in temperature. 
Instead of these processes assuming the normal features of 
metabolism, they lead to chemical decomposition in the com- 
paratively dry and withered tissues ; in other words, they induce 
death from frost. It is therefore emphatically to be recom- 
mended that plants affected by late frost should be protected 
against a too rapid thaw. 

It often happens, even in the case of our indigenous trees, 
e.g: the oak, that after a cold wet summer the vigorous Lammas 
shoots have not ceased growing when the first early frost 
appears. Exotic trees, whose vital processes demand more heat 
for their normal maintenance than our climate has to offer, find 
themselves every year in an unprepared condition on the advent 
of winter. The youngest organs of the annual shoots have not 
completed their development (and especially is this the case 
when growth in height continues till the latter part of summer, 
as happens with Ailanthus, Sic), the youngest elements of the 



wood-ring arc still in an cmbr\-onic condition and with their 
walls unlignified, and the plastic substances have not yet been 
converted into reser^•e materials. Such trees display the same 
sensitiveness to winter frost that our indigenous trees do to 
late spring frost. After a rapid thaw the interrupted chemical 
processes induce decomposition. 

Numerous pathological phenomena in plants have been 
crroneousi}- attributed to frost, and in particular so-called tree- 
canker has frequent!}' been ascribed to this cause.* Most of the 
forms of canker are infectious diseases, and it is only in a few 
extrcmel}' frost}' localities that I have had 
the opportunity of noticing cancerous spots 
which were undoubted!}' due to frost. These 
were met with on a great variet}" of dicot}'- 
ledonous trees, and in order to distinguish 
this form of disease from the work of can- 
ker-inducing fungi I have designated it 
" Frost-canker." ^ 

Frost-canker always occurs at the base 
of a lateral branch that has been killed 
b}' severe late frost. The first symptoms 
are found in the callus which surrounds 
the base of the dead branch. Should the 
locality (frost-hollow) be \-isited b}' late 
frosts during a series of }-ears, the callus, 
which has not had time to protect itself 
b}- a dense firm periderm, is killed in the 
month of May by frosts which occur after 

the tissues have resumed the state of vegetative activity. The 
tissues often die to the distance of half an inch or more from 
the base of the branch (Fig. 158). Subsequenth' a new callus 
forms under the dead and rapidly decomposing cortex. Should 
the plant be unaffected by late frosts for several successive years, 
these canker-spots ma}' heal up complete!}'. But, on the other 
hand, should such frosts recur, the canker-spot increases in size 
with each unfavourable year. The fact that frost-canker makes 

' R. Hartig, Untef^siichi/ngen, I. p. 135, Table VII. 

* [See Sorauer, PJianzenkrankheiten^ B.I. 1SS6, for the arguments in 
favour of this view. — Ed.] 

Fig. 158. — The branch 
of a beech showing 
frost-canker in the 
vicinity of the base of 
a shoot that has been 
killed by frost. The 
wood is brown intern- 
ally. Natural size. 


progress only in a frosty year distinguishes it from fungoid 
canker, which spreads every year. It is further to be noted that 
the late frost also kills the wood at the exposed region as far in 
as the medulla. The products that result from the decomposi- 
tion of the contents of the dead cells distribute themselves more 
or less both up and down the stem, whereas in the case of 
fungoid canker the exposed wood usually becomes brown only 
on the surface. 

In the case of many trees, especially exotic dicotyledons, the 
small fissures in the cortex which are induced by cold prove the 
primary cause of canker. 



In science and in practice two entirely different phenomena 
are referred to under the first of these terms. The more 
frequent phenomenon, which I shall specially designate bark- 
scorching, is caused during the months of July or August by the 
action of unusually strong sunshine on the bark of smooth- 
stemmed trees which have been suddenly exposed after growing 
up in a close wood. 

The trees that suffer most from bark-scorching are the beech, 
hornbeam, spruce, Weymouth pine, and silver fir.* The 
commonest causes of exposure are the formation of roads, 
railways, or rides, or the retention of certain trees for the 
production of seeds, or as standards. 

The injury to the bark by drying up and exfoliation occurs 
almost always on the south-west side, the reason being that this 
is the side on which the sun's rays impinge at the time of the 
maximum daily temperature. 

The extensive clear-felling of spruce woods that had been 
entirely or largely defoliated in Upper Bavaria by the nun 
moth afforded an opportunity for some careful observations on 
the temperature of isolated trees. On August i8, the warmest 
day of 1892, when the thermometer registered 96"8° F. in the 

* [These injuries occur not unfrequently even in our climate. It should be 
noted that the word "bark" is here vised in a somewhat loose sense : true 
bark is dead, and it is the living tissues below which suffer. — Ed.] 


shade, and I04'9°F. on a felled area that was not exposed to 
the wind, it was found that on the south-west side of eighty- 
year-old spruces fully exposed to the sun the temperature was 
131° F. between the wood and the bark. Four weeks later the 
whole of the south-west side of most of the trees had died. 
The high temperature may possibly be explained by the fact 
that the trees had small crowns, and that consequently but little 
water found its way up the younger wood-rings. B}- com- 
paring the temperature of the cambium of beeches, spruces, and 
pines of the same age and thickness, the influence of the cortex 
and bark in modif)'ing the temperature of trees fully exposed 
to the sun was determined. On September 30, at 10 A.M., when 
the temperature of the air was 6g-8° F., the temperature on the 
south-east side of the thin-barked beech was 98'6° F., of the thin- 
barked spruce 82'4° F., and of the thick-barked pine 68° F. This 
would appear to indicate that in trees with thin periderm or 
bark the branches on isolated individuals come well down the 
stem, so as to afford protection against the sun, a state of things 
that one does not find to the same extent in trees with thick 

On standards in a young wood bark-scorching first appears, 
and is most severe, near the surface of the ground. There are 
two reasons for this. First, the rays reflected from the ground 
increase the temperature ; and, secondly, the air-currents that 
assist so materiall}' in cooling those parts of a tree which are 
exposed to the sun are interfered with by the young trees. 

Even the parenchymatous tissues of the injured parts of the 
stem succumb to drought, and the alternate desiccation and 
saturation with external moisture induces rapid decomposi- 
tion, which of course speedily affects the internal portions of 
the stem. Should parasitic tree-fungi effect an entrance, the 
tree may be rapidly killed, but otherwise the decomposition 
retains the simple character of wound-rot. 

I investigated and described a disease which I found in a 
wood of Weymouth pines about forty years old.^ This disease 
both agrees with and differs from bark-scorching, and may be 
designated " bark-drought." The extraordinary drought of 1876 
had reduced the supplies of water in the trees of a wood growing 
1 Uiitcrsiichiingcn^ III. pp. 145 — 149. 


on dry ground intermixed with a silicious moor-pan, to such an 
extent that the cortical and other Hving tissues beneath the 
bark exposed to the drying winds became completely withered. 
This occurred on the south and west sides of the trees, and 
especially at a height of from three to six feet, although 
portions both above and below these heights were also affected. 
The Weymouth pine is found naturally in marshy situations, 
and, adapting itself to the natural habitat of the tree, its 
cortex is but poorly protected by periderm and bark. It 
is thus easy to understand that on a dry soil and in a hot 
dry year the wood is unable to furnish the cambium and 
cortical tissues with sufficient moisture. It follows therefore 
that this species of tree should not be cultivated on excessively 
dry ground, especially where water cannot be expected to ascend 
from the subsoil. 

Of quite another character is the pathological phenomenon 
appropriately called " sun-crack," which is sometimes met with in 
late winter or spring in the beech, hornbeam, Acer, and oak.^ In 
spring, fissures varying in length form in the cortex, which 
separates from the wood for an inch or more on both sides of 
the wound. In the case of the beech, with its thin cortex, the 
rind * not only becomes detached but also dies. Owing to the 
vigorous formation of callus, such a sun-crack frequently heals 
up after a few years, whereas in the case of bark-scorching it is 
very seldom that healing occurs. Fig. 159 represents, in one half 
the natural size, the cross section of the upper part of an oak 
taken from the south side of the stem. The tree, which was 
about 170 years old, and was taken from a light pole-wood of 
beeches on a fairly steep north slope, showed that numerous 
sun-cracks had been formed all over the stem, at various periods 
of its existence. 

The cold ground, which in spring was hardly affected by the 
sun even at midday, must have kept down the temperature 
of the wood of the oak to a low point, even when the stem 
was intensely heated by the sun's rays. It is probable that 
the cortex had become so warm at certain places under the 

^ Lhitersuchimgen, I. p. 141. 

* [The word " rind " is here used in a general sense to denote all the tissues 
outside the cambium. — Ed.] 


influence of the sun's rays that it expanded violentl)', and so 
became detached from the wood. The question, however, has 
not yet been settled by experiment, and unfortunately it is 
scarcely possible in this wa\- to determine the factors that 
combine to produce sun-cracks. 

As a further result of dryness of the air and of excessivel)- 
strong sun, the premature withering and fall of leaves may here 
be mentioned. In 1876 Ihad the opportunity of observing this 
in an intensified form in all the beech woods on south and \\-est 

Fig. 159. — Transverse section of an oak-stem showing numerous sun-cracks. ( )ne 

half natural size. 

slopes in the northern Harz. The beech pole-woods were 
almost entireh' defoliated in the end of August — that is to 
sa}', nearl}' two months before the normal time of the fall of 
the leaf. As this state of things was manifest even on fairly 
fresh ground, it must be attributed to an abnormal rate of 
transpiration from the leaves during the hot di')' summer, to 
compensate for which water could not be convej-ed quickl}- 
enough from the ground. 

When plants have been kept in a humid atmosphere, as, for 
instance, in a forcing-house, a conservatory, or under the shade 
of a close wood, the shoots, but especiall}' the leaves that are 


produced under such circumstances, are peculiar in possessing 
an epidermis which is comparatively non-tuberous.* On this 
account it is ill-adapted to prevent the excessive transpiration 
which is encouraged by air-currents and a dry atmosphere, and 
such plants wither or lose a portion of their leaves prematurely. 

A sudden accession of light in too large quantity has also a 
prejudicial influence on the health of plants, and especially 
on the leaves of trees, whether dicotyledons or conifers. 
Under normal conditions the chlorophyll-corpuscles protect 
themselves against the action of too bright light, which would 
destroy their green colouring matter, by so arranging themselves 
in the cells of the leaf that their narrow edge only is exposed to 
the intense illumination. The leaves of plants that have been 
reared in shade become yellow or brown when suddenly exposed 
to the action of direct sunlight. In such a case, however, it is 
always difficult to determine how much of the damage is to be 
ascribed to the accelerated transpiration induced by the intense 
sunlight, and to the consequent withering of the cells. 

On the other hand, it is a familiar fact that pathological 
phenomena may also be induced by deficiency of light. A plant 
that has grown up in unrestricted light possesses a certain stock 
of the products of metabolism which have not, so far, been 
utilized in the construction of cells. These may take the form 
of reserve materials which have been stored up in the plant, or 
of active plastic substances which are distributed throughout the 
leaves and organs of the stem. By means of these substances a 
plant is able to grow for a certain time even without light, until, 
in fact, the substances have been utilized and the supply has 
been exhausted. Shoots and leaves that have been produced 
in the dark are, however, abnormally constructed. They are 
spindly, and " drawn," and display the phenomenon of so-called 
etiolation. As chlorophyll can normally be produced only under 
the action of light, and as the supply of nutritive substances is 
insufficient, the shoots and leaves are yellowish and not properly 
developed. Seeing that light cannot exert its retarding influence, 
the shoots become abnormally elongated. Such drawn shoots, 

* [There are other anatomical differences also in the cellular tissues of 
such shade leaves, as Stahl has shown, which are calculated to make them 
less resistant. — Ed. 


being unprovided with a properly developed epidermis, wither or 
easil}' succumb to other influences when the plants are again 
fully exposed to the light, and are incapable of developing into 
normal shoots. 

The laying of cereals is the result of the shading of the lower 
internodes in consequence of thick seeding or heavy manuring. 
The restriction of light that results from drilling seed thickly 
stimulates spruces, pines, and other plants to make increased 
height-growth, but this is secured at the expense of the lateral 
shoots and the health of the plants. 


Reference may here be made in a few words to such 
mechanical injuries as are due to atmospheric precipitations 
and violent gales, and especially as these often lead to other 

Flowers and leaves are damaged by heavy hail, which ma}- 
also severely injure the cortex of trees, especially when the rind 
is smooth. At the places where the hailstones strike, the rind 
is crushed, or, it may be, knocked off altogether. Although as a 
rule a callus very soon forms over such wounds, still it not 
unfrequently happens that the injured portion of the stem dies. 
In young spruce woods in the neighbourhood of Munich I found 
that the leading shoots which were affected b)- hailstones died — 
a result doubtless due to the excessive evaporation from the 
wood, which in many cases was stripped of its cortex on one side 
of the shoot to the distance of about an inch. 

It very frequently happens that the wounds caused by 
hailstones form an entrance for parasitic fungi. The spores of 
Ncctria ditissinia are specially apt to germinate on such places, 
and to produce canker in the beech (Fig. 39, page 93). Larches, 
too, are often similarly infected by Peziza Willkovnnii. 

There is not much to be said about the damage that is induced 
by snow-crushing. For obvious reasons, this occurs almost 
exclusively in woods of evergreen conifers, where it takes the 
form either of the breaking off of the tops and branches, or of 
the fracturing of young poles. It may be worth noting, however, 
that wounds are very often formed at the base of branches 
which are bent down by a load of snow. .Should the ground be 


covered with snow, and should the apex of such a branch become 
frozen into the upper layers, it may readily happen that during 
the gradual melting and shrinking of the snow the branch is 
forcibl}' detached from the stem altogether. Such wounds 
frequenth' form the means of entrance for the above-named 
parasitic fungi. 

Gales ma}' fracture stems or tear trees up by the roots, but 
such injuries fall rather within the limits of a treatise on 
s}'lviculture or forest management than of pathology. 


Attention ma)- here be directed to the fact that the destructive 
effects of the passage of fire over the ground of a wood depend 
not onl}- upon the intensity and duration of the conflagration, 
but also upon the species and age of the trees, or, in other words, 
upon the amount of protection afforded by the cortex and bark. 
As is known, the lower portions of the bark of old pines may be 
perfccth' black and charred without the cambium being killed. 
This is due to the low conductivity of the bark for heat.* If ro 
brownness is to be observed in the younger layers of the bast, 
it is evident that the fire can have done no damage. On the 
other hand, trees with thin bark are very sensitive to fire, and by 
making a few incisions in the cortex one may determine whether 
it has been killed or not. Although trees whose lower cortex is 
damaged ma}' produce fresh leaves, one must not be deceived 
b}' such a state of things. Trees that are no thicker than one's 
arm become green in spring when the lower cortex is charred 
or withered right round. A similar state of things occurs with 
beech-saplings that have been barked by mice, but in both cases 
the trees ultimately wither up entirel}^ During the growing 
season the starch that is stored up in the stem at a lower level 
than the dead cortex is utilized by the cambium — which is no 
longer nourished from above — in the formation of the wood-ring 
so that when the trees die in the course of the summer the 
stools, being destitute of reserve supplies, are unable to produce 

* [In cases where the cambium is scorched for some distance, but not 
entirely, round the stem, the remnant of living cambium may slowly creep 
round and form callus over the injured side : years afterwards, on felling , 
such parts of the stem present " ring-shakes."' — Ed.] 


fresh shoots. Shoots arc produced much better from the stools 
of trees that have been entirely consumed, or that have been cut 
over close to the ground directh' after the injury occurred. In 
such a case the whole of the plastic materials stored up in the 
subterranean parts of the tree are at the disposal of the new- 
shoots. If the injured stem is sufficiently young to hold out the 
prospect of stool-reproduction at all, it can only do harm to 
delay cutting it over. 



In the neighbourhood of extensive blast furnaces or similar 
centres of industry, where large quantities of coal are consumed, 
it has alwa}-s been noticed that vegetation suffers from the 
smoke. To such an extent is this the case that in industrial 
towns like Essen scarcely any vegetation exists. In the direction 
of the prevailing winds very serious damage is not unfre- 
quently caused even for a distance of two miles from the fur- 
naces. The views at one time held that the damage was due to 
metallic poisons (arsenic, zinc, lead) present in such smoke, or 
to the soot deposited on the leaves, hav'e proved to be incorrect- 
The investigations of Stockhardt - and Schroder ^ have shown 
that the damage is due entirely to the sulphurous acid present 
in the smoke. It has been determined by experiment that the 
sulphurous acid being absorbed by the surface of the leaves 
induces local death and brownness in the tissues. The tissues 
prove most resistant in the neighbourhood of the larger ribs. 
Although the leaves of conifers absorb less sulphurous acid than 
those of dicotyledonous trees, still, on account of their being 
longer exposed to the prejudicial influences, they generally 
suffer more than the foliage of deciduous trees.* If one examines 

^ Hasenclever, Ucbcr die Bcschiidigung dcr Vegetation dureh satire Case. 
Berlin, 1879. 

- Stockhardt, Tharander forstl. JaJirbiicli^ 1871, p. 218. 

■'■ ScYixodeY, La?idwirthsc/iaftL Versuehsstationen, 1872 and 1873. 

* [I have investigated many such cases, and find the Larch sutlers greatly- 
The cases are complex, and it is by no means clear that the action of the 
acid-gases is merely local on the leaves ; there is evidence to show that the 
damage is largely due to the gases passing through the stomata and into the 
lacunae of the living leaves. — Ed.] 


spruces that are still living, though full)' exposed in the 
neighbourhood of blast furnaces, it will be found that it is only 
on the }'oungest shoots that the leaves are still green. The 
farther one moves from the seat of the mischief, so do the 
annual crops of leaves that still maintain their position on the 
spruce-shoots increase. It is thus evident that the duration of 
the leaves depends in large measure on the intensity of the 
action of the smoke. Amongst dicotyledonous trees the beech 
is the most sensitive, after which come the oak and the 
sycamore, while the elm, ash, and mountain ash, and, amongst 
conifers, the black pine are some of the most resistant. In 
towns where it is only in winter that large quantities of coal 
are used as fuel, the conifers alone suffer. In summer the air 
is almost free from sulphurous acid, and it is only on the 
approach of cold weather that the deleterious influences begin 
to make themselves manifest. At this time the deciduous trees 
have shed their leaves, so that it is only the conifers that are 
affected. The sulphurous and sulphuric acids that collect in 
large quantities in snow that has covered foliage for some time 
prove injurious to the trees. 

The ease with which sulphurous acid is oxidized to hydrated 
sulphuric acid not only explains how this plant-poison is 
constantly being removed from the atmosphere, but also 
indicates how we may remove sulphurous acid from the smoke 
of blast furnaces and factories generally. To some extent this 
has already been put into practice. By leading the sulphur 
gases through moistened hydrated lime 90 per cent, is rendered 
innocuous. Another plan is to conduct the gas through a long 
pipe in which a stream of water flows in the opposite direction. 
By this process a conversion into hydrated sulphuric acid is 

According to recent observations the chlorine and soda fumes 
that are produced in certain factories also prove injurious to 


Up to the present the way in which lightning affects the 
health of trees remains unexplained. 

When lightning strikes a wood, its effects may be confined to 


a single tree, or the}' ma}' be noticeable on a whole group of 
trees. As regards the former case, we find that all trees are 
subject to be struck b}' lightning, but that some are more liable 
to suffer than others. Oaks and the Lombard}' poplar would 
appear to be struck most frequently, though the Scotch pine is 
also very often affected ; whereas the beech enjoys comparative 
immunit}' from such injur}'. Even in trees of the same species 
the form of damage varies exceedingly. As a rule the injury is 
confined to the separation from the wood of a strip of cortex 
about an inch in breadth. This lightning score, w^hich begins 
in the crown, is frequentl}' interrupted over considerable portions 
of the stem. It may leave one side of the tree and appear on 
another, again to return to the original side at a different level. 
In stems with straight fibres it runs straight, but in trees 
showing spiral growth it follows a similar course. At the 
bottom of the tree it disappears between two roots close to the 
surface of the ground ; or it runs for some distance along the 
under side of a strong lateral root, and then suddenly disappears. 
B}' this treatment the health of the tree is in no wise affected. 
The narrow strip of wood is either wholly uninjured or else 
reveals a small crack down the centre. Externally it shows but 
little brownness, and in a few years it becomes entirely covered 
over b}' a callus. 

In other cases trees (pines) that are struck b}' lightning 
reveal externally the same form of injury, but in a few days 
the entire cortex — with the exception of that on the collar, 
the roots, and the upper part of the crown — dies and becomes 
brown. Such trees generall}' wither up after an interval varying 
from a few months to a }'ear or so, although the}- may remain 
alive for four or five years, to die at the end of that period. In 
some cases the electric current barks the tree and leaves the 
stem almost naked, or it splits the stem longitudinally into 
several parts, dismembering it almost entirely, and scattering 
large splinters to a distance of one hundred yards. In certain 
cases all that is left in the ground is a short stump. 

It is onl}'' when the tree is perfectly dry, or possesses dr}' 
branches or at least dr}' rotten wood, that the lightning sets it 
on fire. Combustion does not follow in a fresh living tree. 

So far no explanation is forthcoming to account for the death 


of the trees over considerable areas that have been affected by 
hghtning, a state of things that I have several times observed 
both in young and old pine woods. ^ In such cases it was 
remarkable that death, instead of affecting the whole area 
simultaneously, spread centrifugally and radially from a given 
point, and frequently continued to carry off the trees for five 
years or more. An investigation of the trees showed that only 
one or a few examples revealed traces of lightning, but that 
between the crown and the collar of such trees, and many others 
in their vicinity, the cortex was dead. In an old pine wood the 
dead bark hung loose from the boles, while the crowns retained 
perfectly green foliage. In a younger wood about thirty years 
of age I found three stems showing traces of lightning along 
the margin of the devastated area which had been steadily 
extending for five years previously. The first of these had died 
within the past year, the second still possessed a green crown 
although its cortex and bast had died between the heights of 
one and a half and eight feet, while the third, in spite of the 
lightning having detached a broad strip of cortex, was perfectly 
healthy in all parts. I confess that in face of these observations 
I am unable to offer an explanation of the action of the 
lightning. The fact that trees struck by lightning sometimes 
remain alive for five years is to be explained in the same way 
as the frequent survival for several decades of pines that have 
been girdled. The water and plant-food move upwards in 
the wood, and the crown, utilizing the products of metabolism, 
remains healthy, and forms new organs. Death occurs only 
when the exposed wood of the bole has graduall}- dried up 
to such an extent that water is unable to pass upwards in 
sufficient quantity. That a tree scored by lightning may 
remain perfectly healthy, while a neighbouring tree not so marked 
may die, may possibly be explained by supposing that in the 
former case the electric current was confined within narrow 
limits, whereas in the latter it was distributed over the whole 
surface, or throughout the entire cortex, of the stem. 

^ R. Hartig, Zcitschrift fii)- Forsf- i/nd Jagdwcsc?z, 1876, pp. 330 cl scq. 


The number placed after the name of a disease refers to the page in 
the text-book where a description will be found. 


1. The seedlings droop and die : PhytophtJiora oi/mivora, 58. 

2. Young plants in the nursery become yellow or die, the stems 
contracting suddenly close to the surface of the ground : Pestalozzia 
Hartigii^ 136. 

3. Young plants are enveloped by a brown fungus : Thelephora 
iactniata, 35. 

4. The leaves bear numerous columnar Kcidia on their under 
surface : Melampsora Goeppertiana, 161. 

5. The leaves display long sporogenous layers on their unde'' 
surface. These rupture and emit yellow spores : Ccro/na Abietis 
pectifiatce, 184. 

6. The leaves, whicn are deformed, are pale yellow, and bear 
aecidia. The branches form witches' brooms : yEciditun elatitmin^ 179. 

7. The leaves are yellowish brown, while, on the under side, the 
mid-rib bears a black longitudinal ridge : Hysterimn nervisequium, 108. 

8. The leaves are yellow, and remain attached to the branch by 
being enveloped in white mycelial filaments : Trichosphcei-ia parasifico, 

9. The branch or stem shows a spheroidal swelling : Aicidiuni 
elatmum, 179. 

10. The branches bear mistletoe, or the stem shows perforations : 
Viscuvi, 25. 



11. The cortex dies right round the branch or stem, and bears 
black tubercles : Phoma abietina, 138. 

12. The stem bears irregular or bracket-shaped sporophores which 
show very fine pores : Polyporus Hartigii, 1 94. 

13. The stem bears sporophores with large pores: Irametes Fitii, 

14. The stem bears tawny yellow cap-shaped sporophores, which 
.spring from rhizomorphs : Agaricus melleus, 207. 

15. The roots bear white sporophores : Trametes radidpej-da, 186. 

16. The roots are attacked by rhizomorphs : Agaricus melleus, 207. 


1. The seedlings show black blotches on the leaves or stem, and 
may decay : Cercospora acerina, 135 ; Phytophthora omnivora, 58. 

2. The leaves show white blotches : Erysiphe bicornis, 70 ; E. 
Tulasfiei^ 70. 

3. The leaves show black blotches : Rhytisma acerinum, 105. 

4. In autumn the leaves show persistent green blotches, which 
ultimately bear black spots : R. piaictatitui, 1 06. 

5. The branches wither, while a transverse section of the wood 
shows dark green blotches : Nectria cinnabarina, 96. 

6. The branch or stem dies, while the cortex bears cinnabar-coloured 
fungus bodies : JV. cinnabarina^ 96. 

7. The stems of young plants contract suddenly above the roots : 
Pestalozzia Harligii? 136. 

8. The branches show canker-spots : Frost-canker, 293. 

9. The branches bear mistletoe : Viscuvi, 25. 

Acer platanoides 

The branches die in spring, and show oblong fungus-bodies : 
Septogloeuni Hartigiamim^ 141. 


The branches die, and bear cinnabar-coloured fungus-bodies on the 
cortex : Nectria cinnabarina, 96. 


1. The leaves show yellow vesicular swellings : Exoascus flavus, 133. 

2. The leaves show greyish white downy corrugations : E. epiphyllus, 


3. The cones show pocket-like outgrowths : E. alfiitorqims, 133. 

4. The branches show canker-spots : Nectria ditissima, 91, 

5. The wood shows red-rot : Polyporus sulphureus, 200. 

6. The roots show fleshy outgrowths : Schinzia Ahii, 39. 


Alnus glutinosa 

The leaves show vesicular corrugations : E. ainiforquus, 133. 

Alnus incana 

The branches bear witches' brooms : E. borealis, 133. 

Alnus viridis 

The branches wither, and black tubercles appear on the dead 
cortex : Valsa oxy stoma, 151. 


The leaves show golden yellow blotches : Piurinia gramhiis, 155. 


1. The leaves show small yellow fungus-bodies : Melampsora 
hetnlina, 171. 

2. The leaves show vesicular swellings : Exoascus carnea, E. Betuke^ 

3. The branches bear witches' brooms : Exoascus turgidus, 133. 

4. The stem bears large bracket-shaped sporophores : Polyporus 
befuli7ius, 206. 

5. The stem bears brown crust-like sporophores : Polyporus Icevigatus, 


The roots bear fleshy outgrowths : PlasmodiopJiora Brassicce, 39. 


1. The leaves bear small golden yellow fungus-bodies : Melampsora 
Carpini, 171. 

2. The branches bear witches' brooms : Exoascus Carphii, 135. 

3. Branches or stem show canker-spots : JSlectria dilissima, 91 
Frost-canker, 293. 


The branches show prominent swellings, and bear a " mistletoe " : 
Lorant/ms, 30. 


r. The leaves show small brown blotches : SphcErella, 88, 

2, The leaves show white dusty blotches : Erysiphe guttata, 70. 

3. The branches show canker-spots : Nectria ditissima, 91. 


1. The leaves bear golden yellow swellings, which produce secidia : 
Gyinnospora7igium clavariceforme, 158. 

2. The leaves show white dusty blotches : Erysiphe guttata, 70. 

3. The branches bear witches' brooms: Exoascus bullatus, 133. 

X 2 



The leaves show small yellow fungus-bodies : Crojiarthon asckpia- 
deujH, 175. 


1. The seedlings show dark patches on the leaves and stem, and 
decay or wither : Phytophthora onifiivora, 58. 

2. The stem of young plants contracts suddenly close to the surface 
of the ground, and the tree withers : Pestalozzia Hartigii, 136. 

3. Young plants in the nursery are enveloped by a brown fungus : 
Thelephora laciniaia, 35. 

4. The leaves show white blotches : Erysiphe guttata., 70. 

5. The leaves show brown blotches : Sphcerella Fagi, 88. 

6. The cortex shows canker-spots : Nectria difissi??ia, 9 1 ; Frost- 
canker, 293. 

7. The cortex is covered with a white woolly substance : Chermes 
fagi, 96. 

8. The cortex shows pustular swellings : ilnd. 

9. The cortex of branches is ruptured longitudinally : Lachmis 
exsiccator, 96. 

10. The cortex of the stem withers on the south side : Bark- 
scorching or Sun-crack, 294. 

11. The stem bears large bracket-like sporophores : Polyporus 
fonientarms, 206. 

12. The wood shows a verdigris-green colour: Peziza (srugijiosa, 


The cortex shows canker-spots : Nectria ditissi/na, 91. 


Gentiana asclepiadea shows yellow fungus-bodies : Cronarthim 
asclepiadeuDi, 175. 

The branches bear mistletoe : Viscian, 25. 


1. Culm and leaves show fungus-bodies, which are first yellow and 
later brown : Pucci?iia graminis, 155. 

2. The spikelets are covered with a sweetish secretion or produce 
black fungus-bodies : Claviceps purpurea, 98. 

3. The spikelets produce dark brown powder : Ustilago Carbo, 68. 


The bulb becomes soft and slimy, and emits a repulsive s^iiell : 
Bacterium, 37. 



1. The branches bear mistletoe : Viscu/n, 25. 

2. The stem bears sulphur-yellow sporophores. The wood shows 
red-rot : Polyporus sulphureiis, 200. 

Juniperus communis 

1. Leaves and branches enveloped in dark brown mycelia : Herpo- 
trichia fiigni, 76. 

2. Branches show swellings which, in spring, produce abundant 
yellow or brownish spores : Gymnospora?igiu)n cofiicum, 1573 G.davarice- 
forme, 158; G.tremelloides^ 159. 

3. Roots bear white sporophores : Tramefes radiciperda, 186. 

Juniperus Oxyeedrus 

Branches bear a " mistletoe " : Arceuthobium Oxycedri, 30. 

Juniperus Sabinse 

Branches show swellings which, in spring, produce abundant yellow 
spores: Gyunwsporangium Sabince^ 158, 


The cortex and branches die : Cucurbitaria Labtirni, 87. 


1. The seedlings droop and wither : Phytophthora omnivora, 58. 

2. The young trees die, and reveal mycelia on their roots : Rhizina 
undulata, 123. 

3. The leaves show yellow fungus-bodies : Melampsora Tremul(B, 

4. The leaves become brown, and show black fungus bodies : 
Hysterium laricitiian, 117. 

5. The cortex shows canker-spots : Peziza Willkominii^ 117. 

6. The cortex, on its inner side, shows white mycelial sheets : 
Agaricus me ileus, 207. 

7. The cortex bears brown crust-like sporophores : Trametes Pint, 

8. The cortex bears large sulphur-yellow sporophores : Polyporus 
sup/mreus, 200. 

9. The cortex bears cap-like tawny yellow sporophores : Agaricus 
melieus, 207. 

10. The roots are dead, and show rhizomorphs : ibid. 

11. The wood shows red-rot : Poiyporus ScJiweijiitzii, 198. 

12. The wood is decayed, and marked by white blotches : Trauietes 
Pifii, 191. 

13. The wood shows red-rot and luxuriant white fungus-growths : 
Poiyphorus suiphureus, 200. 



The leaves are marked by brown blotches, and show small yellow 
fungus-bodies: Chrysomyxa Ledi. 179. 

Medicago, see Trifolium 


1. The seedlings droop soon after appearing : Phytophthora o/?inivora, 
58 ; Nedria cuairbiiuia, 89. 

2. Plants in the nursery become yellow and die, the stem being con- 
tracted close above the surface of the ground : Pestalozzia Hartigii, 136. 

3. Young plants, or the branches of older trees, are enveloped in 
dark brown mycelia : Herpotrichia nigfa, 76. 

4. Young plants are enveloped in the sporophore of a fungus : 
Thelephora laciniata, 35. 

5. Young and old trees die, their roots showing mycelia : RJiizimx 
undulata, 123. 

6. The leaves and branches wither in winter and spring : Frost, 

7. The leaves bear golden yellow vesicles : ChrysoJiiyxa Rhododendri, 
177 ; C. Ledi, 179. 

8. The leaves become yellow, and show golden yellow longitudinal 
ridges on their under surface : Chrysoinyxa Abietis, 175 

9. All the leaves of a young shoot are abnormally short, and rupture 
on their four sides : yEcidium coriiscans, 183. 

10. The leaves become red, and later yellowish brown. They either 
show longitudinal black ridges, or fall prematurely : Hysterium macro- 
sporutn, 109. 

11. The branches die in May or June : Septoria parasitica, 143. 

12. The scales of the cones show numerous round brown swellings 
on their upper surface : Aicidiiim strobiIi7ium, 182. 

13. The scales of the cone show two large tecidia on their lower 
surface: ^cidijim amorum Piccce, 183. 

14. The cortex shows dead patches beset with groups of red fungus- 
bodies : Nedria Cucurbitula, 89. 

15. The cortex in the lower part of the stem shows resinous exuda- 
tion: Trametes radidperda, 186. 

16. The cortex shows white mycelial sheets on its inner surface : 
Agaricus melleus, 207. 

17. The cortex shows evidences of injury by sun : Bark-scorching, 

18. The root is dead, and bears small yellowish white fungus-bodies, 
or large white sporophores : Trametes radidperda, 186. 


1 9. The root shows red-rot and white mycelia : Folyporus vaporarius^ 

20. The root is dead, and shows black mycelial strands which form 
white enlargements between the cortex and wood : Agaricus melkus, 

21. Branch-wounds bear brown sporophores : Tra metes Phii^ 191 r 
Polyporus Hartigii, 194. 

22. Wounds bear large white sporophores : Polyporus borealis, 196. 

23. The wood shows white-rot : Polyporus Hartigii, i()s^. 

24. The wood shows white-rot. The pure white patches have usually 
a black spot in the centre : Trametes radiciperda^ 186, 

25. The wood shows white-rot, and contains numerous cavities : 
Trametes Pi?ii, 191. 

26. The wood shows white-rot, and crumbles down into very small 
cubes : Polyporus borealis, 196. 

27. The wood shows red-rot : Polyporus vaporarius, 198. 

28. The wood shows dark brown blotches or cavities : Wound-rot, 
236, 243. 

29. The wood shows green-rot: Peziza aruginosa, 224. 

Pinus Cembra 

The roots show numerous Mycorhizce, 71. 

Pinus montana 

The branches with their leaves are enveloped and killed by dark 
brown mycelia : Herpotrichia nigra, 77. See also the diseases, i, 6, 8^ 
10, 13, 16, 17, under Pinus sylvestris. 

Pinus Strobus 

1. The leaves die, and display black fungus-bodies : Hysterium 
brachysporum, 117. 

2. The cortex shows resinous exudation and golden yellow vesicles : 
Peridermiuni Strobi, 175. See also diseases i, 4, 10, 13, 14, 16, 17, 
under Pinus sylvestris. 

Finns sylvestris 

1. The seedlings droop and die: Phytophtho7-a omnivora, 58: 
Nectria Cucurbitula, 89. 

2. Seedlings and older plants show brown blotches, which afterwards 
bear small black tubercles : Hysterium Pinastri, no. 

3. Seedlings and older plants are entirely yellow and finally brown 
or the discoloration spreads gradually back from the apex of the shoots 
"Bhght," III. 


4. Young plants in their lower parts are enveloped in the brown 
sporophores of a fungus : Thekphora laciniata, 35. 

5. The leaves become suddenly brown in summer : " Frost-bhght," 
I II. 

6. The leaves show^ golden yellow^ vesicles : Coleosporiiim Se?iedo?us, 
C. Eiiphrasice, C. Ttissilagi?iis, 172, 

7. The young shoots in the end of May show golden yellow spots 
■on the cortex. These afterwards rupture, and the shoots either die or 
become contorted : Melampsora Tremulce, 164. 

8. The cortex shows golden yellow A^esicles filled with spores : 
Teriderinhim Pmi, 172 ; CroJiartmm asclepiadeian, 175. 

9. The cortex gradually dies, and shows resinous exudation : 

10. The cortex dies, and shows large white sheets of mycelium on its 
inner surface : Agariais melleus^ 207. 

11. Branch-wounds bear brown bracket-Uke sporophores: Trametes 
Fini, 191. 

12. Wounds bear large reddish brown cushion-like sporophores: 
Poly poms Schweinitzii, 198. 

13. The cortex close to the ground bears cap-like yellow sporophores : 
Agariais vielleus, 207. 

14. The cortex close to the ground bears white cushion-like sporo- 
phores : Trametes radidperda, 186. 

15. The cortex or wood close to the ground bears white porous 
crust-like sporophores : Polyporus vaporarius, 198. 

16. The roots are dead, and bear yellowish white cushion-like sporo- 
phores : Trametes radiciperda^ 186. 

17. The roots are dead, and show resinous exudation. Between the 
wood and cortex white mycelial sheets and black mycelial strands are 
found : Agaricus 7JieUeus, 207. 

18. The roots are dead, and show white floccose mycelial strands : 
Polyporus vaporarius., 198. 

19. The roots show mycelial growths : Elaphomyces, 71. 

20. The leading shoot or the branches die above a black mark from 
which resin flows: Crojiartium asdepiadeum, 175; Peridermiuju Pi?ii, 

21. The wood shows white-rot, with numerous small round or oval 
holes: Trametes Piiii^ \Q)\. 

22. The wood shows red-rot, without much smell. Floccose mycelial 
growths and strands are found : Polyporus vaporarius., 198. 

23. The wood shows red-rot, and emits a very strong smell of turpen- 
tine. Thin white mycelial incrustations are found in the cracks : 
Polyporus Sdiweinitzii, 198. 


24. The wood shows holes, the branches bear mistletoe : Visciim, 25. 

25. The wood (alburnum) shows a dark blue colour : Ceratostoma 
pilifej'um^ 224. 

26. Old and young trees die, the roots showing mycelia : Rhizina 
jmdiilata., 123. 

The leaves and young shoots die, or the former become brown along 
the ribs : Glaosporium ner-viseqiimm^ 140. 


1. The leaves show small yellow blotches, which afterwards become 
dark brown : Melampsora, 164. 

2. The leaves show yellow vesicular swellings : Exoascus aureus, 


3. The branches bear mistletoe : Viscum, 25. 

4. The flowers exhibit golden yellow much-enlarged ovaries : 
Exoascus mireiis, 135. 

Populus pyramidalis 
The branch and twigs die : Didyinosphceria popiilijia, 104. 

Prunus Cerasus 

1. The leaves are crumpled, and frequently also blood-red in colour : 
Exoascus IVtesnen, 132. 

2. The leaves become prematurely yellow and die, and remain 
attached to the tree during winter : Gnomonia erythrostoma, 88. 

3. The branches form witches' brooms : Exoascus Wiesneri, 132. 

4. The cortex bears brown sporophores : Polyporus igniarius, 201. 

Prunus domestica 

1. The flowers show yellowish red fleshy blotches : Poly stigma 
rubrum, 97. 

2. The fruit forms "pockets" : Exoascus Pruui, 131. 

3. The branches form witches' brooms : Exoascus defortiians, 132. 

4. The branches show black tuberous swellings : Plowrightia mor- 
bosa, 102. 

Prunus instititia 
The branches form witches' brooms : Exoascus InstititicE, 133. 

Prunus Padus 

1. The fruit forms "pockets": Exoascus Pruni, 131. 

2. The cortex shows canker-spots: Nectria ditissima, 91. 

Prunus spinosa 

1. The leaves show yellowish red fleshy blotches: Polystigvia 
rubrum, 97. 

2, The fruit forms "pockets " : Exoascus Prufii, 131, 



I The young shoots die, and become brown : Botrytis Donglasii^ 130. 
2. The branches bear a "mistletoe," and show witches' brooms: 
Arceuihobhim Dotiglasii^ 30. 

Pyrus communis 

1. The leaves show yellow swellings, which produce ^cidia : 
Gymnosporanghim SabincB, 158. 

2. The leaves show vesicular swellings : Exoascus bullafus, 129. 

3. The stem bears brown cushion-like or bracket-shaped sporo- 
phores : Poly poms ignariits, 201. 

4. The branches bear mistletoe : Visciun, 25. 

Pyrus Malus 

1. The leaves bear yellow swellings, which produce ascidia : Gyimio- 
spora?2gh(m tremelloides, 159. 

2. The branches show canker-spots: Nectria ditissi?iia, 91; Frost- 
canker, 293. 

3. The stem bears brown cushion-like or bracket-shaped sporo- 
phores : Poly poms ig7iarius^ 201. 

4. The branches bear mistletoe : Viscum, 25. 


1. One- and two-year-old plants wither, and show mycelial strands 
and black tubercles on their roots : Rosellinia querchia^ 78. 

2. The leaves show vesicular swellings : Exoascus ccerulescetis, 135. 

3. The leaves show round brown blotches : Sphcerella, 88. 

4. The cortex shows canker-spots: Nectria ditissi?)ia, 91; Frost- 
canker, 293. 

5. The cortex of young oaks dies over large areas of the stem, and, 
should the trees have survived, a callus forms along the margin of the 
wound : Aglaospora Taleola, 99. 

6. The wood is dry, and shows red-rot : Polypoms sulp/inreus, 200 ; 
FistuHna /lepatica, 206 ; Dc^dalea quercbia^ 206. 

7. The wood shows white-rot: Polypoms ignarius, 201; Hydman 
diversidens, 202. 

8. The wood shows red-rot with white stripes : Stereuvi hirsuiian, 

9. The wood shows red-rot with white blotches and cavities : 
Thelepho7-a Perdix, 203. 

10. The wood shows irregular oblong patches of red-, white-, and 
yellow- rot : Polypoms dryadeiis, 201. 

11. The branches bear a deciduous "mistletoe" and prominent 
swellings : Loraiithus europceiis, 30. 



The leaves and shoots show golden yellow swellings : Puccinia 
coro7iata^ 156. 


1. The leaves bear large galls : Exobasidhim Vaccinii, 185. 

2. The leaves show brown blotches : Chrysomyxa Rhododendri^ 177. 


The leaves show yellow swellings : Melampsora Hartigii, 170. 


The wood shows red-rot. Sulphur-yellow sporophores appear upon 
the cortex : Polyporiis sulp/iureus, 200. 


1. The leaves show small yellow fungus-bodies, which become 
brown in autumn: Melampsora sa/ichia, 170. 

2. The leaves show large black thickened blotches : Rhytisma 
salicmian, 107. 

3. The leaves show white dusty blotches : Erysiphe adiinca, 70. 

4. The wood shows red-rot, and sulphur-yellow sporophores appear 
upon the surface of the stem : Polyporiis sulpliureiis, 200. 


Leaves and stem show reddish yellow fungus-bodies : Cokospormm 
Sefiedonis, 172. 


1. Leaves and stem show black blotches : Phytophthora mfestans, 64. 

2. The tubers show disease : P. infestans^ 64 ; Bacterium, 38. 

Sorbus Aria 

The leaves show fungus-bodies, which produce secidia : Gymnospor- 
angium tretnelloides, 159. 

Sorbus aucuparia 

1. The leaves show large golden yellow blotches, which produce 
secidia: Gymnosporangium coniaan, 157. 

2. The leaves show small yellow fungus-bodies : Melampsora Sorln, 


3. The cortex shows dead patches, which bear small fungus-bodies : 

Ciicurbitaria Sorbi, 88. 

4. The branches bear mistletoe: Viscum, 25. 

Sorbus torminalis 

The leaves show yellow blotches, which bear cecidia : Gymjiosporan- 
gium coniaan, 157. 



1, The twigs and branches die, and produce cinnabar-coloured 
fungus-bodies : Nedria ciiutabarma, 96. 

2. The cortex shows canker-spots : Nedria ditissima, 91. 

Trifolium and Lucerne 

1. Roots attacked by violet Rhizodonice, 82. 

2. Close to the root-collar white mycelia and black resting-mycelia 
may be detected : Peziza ciboriodes, 130. 


The leaves and shoots enveloped in white mycelia, the former 
dying: Trichosphceria parasitica, 72. 


The leaves show vesicular blotches : Exoasctis U/mi, 135. 

Vaccinium Myrtillus 

1. The young shoots die and the berries shrivel up : Sderotinia 
baccarum, 130. 

2. The leaves show small brown blotches : MeIa?npsora Vacdnii, 

Vaccinium Vitis-idaea 

1. The stem becomes much elongated, and attains the thickness of 
a goose-quill : Melampsora Goeppertiaiia, 161. 

2. Leaves, flowers, and stem are swollen, and dusted with white 
spores: Exobasidium Vacd?iii, 185. 

3. Leaves, fruit, and young shoots become brown : Sderotinia 
Vacdnii, 130. 


1. Leaves, stem, and berries show mildew : Oidiuvi Tuckeri, 70. 

2. The leaves show yellow blotches above and white blotches 
below : Peronospora viticola, 65. 

3. The berries wither : Physalospora Bidwellii, 103. 

4. The berries are pale in colour, and their stalks decay : Coiiiothy- 
riii/ii diplodiella, 103. 

5. The roots are killed by Rhizoctonice and Rhizomorphs : Demato- 
phora 7iecatrix, 82. 

6. All parts of the plant show brown or black blotches : Gloeosporiuvi 
ainpelophagum, 104. 

Leaves, flowers, and stem show black vesicles filled with spores : 
Ustilago Maydis, 68. 



Abies pectinata, see Fir, Silver 
Abnormal predisposition to disease, 

9 . . . 

Acclimatization of exotics, 2S8 

Acer, Dematophora necatrix on, 82 

Erysiphe bicornis on, 70 

Nectria cinnabarina on, 96 
ditissima on, 92 

Rhytisma acerinum on, 105 
punctata on, 106 

campestre, Septoglceum Hartigi- 
anum on, 141 

platanoides, Mildew on, 70 

Seedlings, Disease of, 58 
Acorns, Influence of size of, on growth 

^cidium abietinum, 49, 177 

asperifolii, 156 

Berberidis, 155 

Clematitis, 166 

columnare, 49, 161, 184 

conorum Piceee, 183 

coruscans, 183 

elatinum, 51, 179, 195 

pencillatum, 1 59 

Rhamni, i 56 

strobilinum, 182 
Agaricus adiposus, 180 

melleus, 47, 48, 50, 83, 186, 187, 
207, 245 
Rhizomorphs of, 43 
Age in relation to disease, 7,8, 11 
Aglaospora Taleola, 99 
Air in soil. Circulation of, 275 

wood, 53 
Alder, see also Alnus 

Nectria ditissima on, 92 

New parasite of, 147 

Polyporus on, 200 

Stigmatea on, 88 

Witches' brooms on, 133 
Alder-roots, Disease of, 39 
Algal fungi, 57 
Almond, see Amygdalus 

Alnus, see also Alder 

Erysiphe guttata on, 70 
glutinosa, Exoascus alnitorquus 

on, 133 
E. epiphyllus on, 133 
flavus on, 133 
incana, Exoascus alnitorquus on, 


E. borealis on, 133 
epiphyllus on, 133 
flavus on, 133 
viridis, Valsa oxystoma on, 151 
Alpine Rose Apples, 186 
Alternation of generations, 154 
Amygdalus, Polystigma fulvum on, 98 
communis, Exoascus deformans 
on, 132 
Annuals, Death of, 6 
Antheridia of Peronosporeas, 58 
Anthracosis of Vine, 104 
Anthusa officinalis, /Ecidium asperi- 
folii on, 156 
Ants, 203, 245 
Aphidas on beech, 96 
Apothecium of Discomycetes, 105 
Apple, see also Pyrus 
Apple-trees, Nectria ditissima on, 92 
Probable bacterial disease of, 
Apricot, Valsa on, 88 
Arceuthobium Douglasii, 30 

Oxycedri, 30 
Arona, Rccstelia cornuta on, 1 58 
Arsenic in smoke, 301 
Arvicola, Barking by, 243 
Asci, 69 
Asclepius Vincetoxicum, Cronartium 

on, 175 
Ascomyces Betulae, 135 
Ccerulescens, 135 
Tosquinetii, 133 
Ascomycetes, 57, 69, 153, 155 
Imperfectly known, 135 
Spore-formation in, 44 



A-sexual generation, 44 
Ash, Erysiphe guttata on, 70 
Nectria ditissima on, 92 
Seedlings, Disease of, 58 
Asparagus, Puccinia on, 1 56 
Aspens and Pine Twist, Connection 

between, 9 
Atmospheric influences, Injuries due 

to, 282 
Autoecious parasites, 154, 176 

BaciUus Olea tubeixulosis, 38 
Bacteria, 37 
Bacteriosis, 37 
Bacterium Hyacinthi, 37 
Barberry, Parasite of the, 25, 155 

in relation to wheat-rust, 9, 49, 

Bark, 226 
" Bark-drought," 295 

Scorching by sun, 268, 269, 283, 
Barking by Cattle, &c., 244 

Game, 241 
Barley, Puccinia striteformis on, 156 

Ustilago Carbo on, 68 
Bary's, de, Work, Value of, 3 
Basidia, Abscission of spores by, 44 
Basidiomycetes, 57, 153 
Beans, Dematophora necatrix on, 83 
Beech, Dematophora necatrix on, 83 

Erysiphe guttata on, 70 

Green-rot of, 42 

Hydnum diversidens on, 202 

Nectria ditissima on, 92 

Peziza aeruginosa on, 224 

Polyporus fomentarius on, 206 

Spha^rella on, 88 

Trametes radiciperda on, 186 

Tuber on, 71 

Wood-balls, or Spheroblasts, on, 


Seedlings, Disease of, 58 
Beef-steak fungus, 206 
Beet, Dematophora necatrix on, 83 

Peziza Sclerotiorum on, 130 
Beetles in wood and bark, 219 
Betula, Erysiphe guttata on, "]"] 

Exoascus on, 133 

Melampsora on, 171 

New parasite of, 147 

Peziza aeruginosa on, 224 

Polyporus on, 206 

Trametes radiciperda on, 186 
Birch, see Betula 
Bird Cherry, see Prunus Padus 

"resin," 175 

Bird's Nest, Yellow, see Monotropa 

Black knot, 103 

Rot of Vine, 103 
Blanc de racines, 82 
Blanquet, 82 
Blight of apple- and pear-trees, 38 

pines, III 
Blister of larch, 117 

pine, 172 
Blotches, Disease, 88 
Borago, yEcidium asperifolii on, 1 56 
Botrytis cinerea, 130 

Douglasii, 130 
Bramble, Parasite of the, 25 

rust, 156 
Brands, 68 
Brood-cell, 44 

Brownness of conifer leaves, 1 1 1 
Buckwheat, see Fagopyrum 
Buds, Adventitious, 238 

Preventitious, 238 
Bunt, 68 
Byssothecium circinnans, 82 

Cabbages, Club-root of, 39 
Casoma Abietis pectinat^e, 184 
Evonymi, 171 
Laricis, 166, 169 
Mercurialis, 166 
pinitorquum, 166 

in regard to moisture, 46 
Ribesii, 171 
Calluna, Dodder on, 34 
Callus, Formation of, 226, 229 

on the stools of conifers, 262 
Calyptospora without effect on old 
tissues, 51 

Goeppertiana, 161 
" Cambium, Wound-," 231 
Canker due to frost, 293 

Nectria ditissima, 92 
soil, 129 
on larch, 117 
pine, 172 
silver fir, 179 
Cap fungi, 1 84 
Carbonic Acid gas. Injurious effects 

of, 281 
Carpinus, see Hornbeam 
Carrot, Peziza Sclerotiorum on, 130 
Castanea vesca, Loranthus europa>us 
on, 30 

Rhizina undulata on, 126 
Caterpillars, Surface, 147 
Ceratostoma piliferum, 224 


Cercospora acerina, 135 

a facultative parasite, 47 
Resting-mycelium of, 43 
Cereals, Puccinia coronata on, 156 

Rust of, 155 
Champignon blanc, 82 
Chermes abietis, 144 
Fagi, 96 

Laricis, 118, 169 
Cherry, Black-knot of, 103 
Gnomon i a on, 88 
Valsa on, 88 
Bird, see Primus Padus 
Wild, see Prunus avium 
Chestnut, Horse, Nectria cinnabarina 
on, 96 

Spanish, see Castanea vesc^ 
Chlorine, Effects of, on vegetation, 

Chrysomyxa, 175 

abietis, 13, 153, 175, 177 
Ledi, 49, 179 
Rhododendri, 9, 49, 177 
Chytridiaceae, 57 

Cinchona bark, Cultivation of, 230 
Clarkia seedlings. Disease of, 58 
Claviceps purpurea, 98 

Sclerotia of, 43 
Clematis vitalba, Ca;oma on, 166 
Climatic predisposition to disease, 

Clover, Dodder on, 34 

Peziza ciborioides on, 130 
Red, Orobanche minor on, 25 
Club-root, 39 
Coal-brand, 68 

Smoke, 300 
Cockchafer grubs, 147, 264 
Coleophora laricella, 117 
Coleosporium Euphrasia^, 172 
Senecionis, 153, 172 
Tussilaginis, 172 
Conidium, 45 

Conifers, Agaricus melleus on, 207 
Hysterimii on, 107 
Mycelium on the roots of, 71 
Polyporus Hartigii on, 194 

vaporarius on, 198 
Rhizina undulata on, 123 
Seedlings, Disease of, 58 
Withering of the leaves of, 112 
Coniothyrium diplodiella, 103 
Coppicing, 261 
Cork, 226 

" Woimd," 227 
Cortex, 225 
Roots. 26 

Corticum amorphum, 117 
Corylus, see Hazel 
Cow-wheat, see Melamp^rum 
Cowberry, see Vaccinium 
Cracks, Frost, 284 

Sun, 283, 294, 297 
Crataegus, Erysiphe guttata on, 70 

Exoascus bullatus on, 133 

RjEstelia lacerata on, 1 58 

Witches' brooms on, 133 
Crickets, 147 
Cronartium, 172 

asclepiadeum, 175 * f ' 

rfbic/la, 175 -• ' * 

Crucifers, white-rust of, 66 
Cryptogams, 35 
Cryptorhynchus lapathi, 151 
Cucurbitaria Laburni, 89 

morbosa, 102 

Sorbi, 88 
Cupressinoxylon, Agaricus on, 207 
Cupuliferje, Mycelium on roots of, 71 
Cuscuta Epilinum, 34 

Epithymum, 34 

eiu'opaea, 34 
Cuscutea;, 33, 34 
Cynips, 186 
Cystopus candidus, 66 
Cvtisus Laburnum, Cucurbitaria on, 

Deedalea quercina, 206 

Damping off of seedlings known to 

early writers, i 
Deadening material, 217 
Death due to external causes, 7 
frost, 2S7 
internal causes, 6 
of animals, 6 

plants, cause of, 6 
Natural and accidental, 6 
Natural, discussed, 6 
Debility of old age as a factor of 

disease, 7, 8, 11 
Deer, Fallow, Barking by, 241 
Red, „ ' 241 

Roe, „ 241 • 

Defoliation by insects, 268 
Demarcation, Line of, between scion 

and stock, 267 
Dematophora necatrix, 43, 82 

an example of mycelial infection, 
Development dependent on variation, 

Didymosphaeria populina, 104 




Discomycetes, 69, 105 

Protection of sporophores of, 
against drought, 45 
Disease-blotches, 88 
Disease, Causes of, 4, 20 

determined by two factors, 8, 9 

Investigation of, 16 

and sickliness distinguished, 5 
Diseases classified, 8 

due to soil, 270 
Dodder, see Cuscuta 
Dogwood, Nectria ditissima on, 92 
Dormant eyes, 239 
Draining, Effects of excessive, on 

alders, 270 
Drawn shoots, 298 
Drought, Effects of, on cereals, 272 

Leaves of conifers injured by, 1 1 1 

of Bark, 294 
Dry-rot, 214 

fungus, 219 

see also Alerulius lacrymans 
Duramen, 33, 44 
Dust-brand, 68 

Echium, ^cidium asperifolii on, 156 

Egg-spores of Peronosporese, 58 

Elsagncce, Tubercles on the roots of, 39 

Elaphomyces granulatus, 71 

Elm, Exoascus on, 135 

Endophyte defined, 42 

Endophytes, Mode of attack of, 50 

Engrafting, 228 

Entomology, Forest, an old study, 2 

Entomophthoreee, 57 

Enzyme, 1 1, 50 

Epicormic branches, 238, 253, 271 

Epidermis, 225 

Epiphyte defined, 42 

Epiphytes, Mode of attack of, 50 

Ergot, see Claviceps purpurea 

Erysiphe, see also Mildew 

Epiphytic character of, 42 

adunca, 70 

bicornis, 70 

guttata, 70 

pannosa, 70 

Tuiasnei, 70 
Erysiphe^, 69, 70 
Etiolation, 298 

Euphrasia, a partial parasite, 24 
Even-aged woods specially liable to 

disease, i 
Evonymus, C^eoma on, 171 
Exoascus, 131 

alnitorquus, 133 

aureus, 135 

Exoascus Betulce, 135 

borealis, 133 

buUatus, 133 

carnea, 135 

Carpini, 135 

CcCrulescens, 135 »• 

deformans, 132 

epiphyllus, 133 

flavus, 133 

Instititias, 133 

Primi, 131 

Sadebeckii, 133 

turgidus, 133 

Ulmi, 135 

Wiesneri. 132 
Exobasidium \^accinii, 185 
Exotics, Acclimatization of, 288 
Experiments in infection, 17, iS, 19, 

Eye-bright, a partial parasite, 24 

Factories, Refuse of, 281 
Facultative parasites, 47 

saprophytes. 47 
Fagopyrum seedlings, Disease of, 58 
Fagus, see Beech 
Felling in summer and winter, 214 
Ferments exuded by parasites, 50 
Figures carved in bark, 246 
Filaments, Truncated, of Erysiphe, 69 
Fir, Douglas, as a protection against 
pine-blight, 116 

Botrytis on, 130 
in relation to drought, 12 
Rhizina undulata on, 125 
Scotch, see Pinus sylvestris 
Silver, ^-Ecidium columnare on, 

elatinum on, 179 
Ca^oma on, 1 84 
Columnar rust of, 161, 184 
Hysterium macrosporum on, 109 

nervisequium on, 108 
Nectria Cucurbitula on, 89 
Pestalozzia Hartigii on, 136 
Phoma abietina on, 138 
Rhizina undulata on, 125 
Trichospha.ria parasitica on, 72 
in connection with adventitious 
buds, 262 
Fire, Effects of. 300 
Fistulina hepatica. 206 
Flower-pots, Failure of plants in glazed. 

Fly wood, 205 
Frank's work, 4 
Fraxinus, see Ash 


Frost, Action of, 282 
Causes of, 284 
Death due to, 287 
Leaves of conifers injured by, 112 
Shedding of leaves owing to, 286 
Transplanting favours injury by, 

I4« 15 
Frost-beds, 10 

canker, 293 
cracks, 284 
"Rib," 285 
Fruit - trees, Dematophora necatrix 

on, 82 
Fungi, 40 

Biology of, 45 
Classification of, 57 
Heat requirements of, 46 
Importance of moisture for, 46 
may attack uninjured plants, 50 
enter only through a wound, 50 
Fungus, Beef-steak, 206 

House, see Merulius lacrymans 
Fusicladium Tremulas, 105 
Fusidium, 141 

candidum, 94 

Gales, Damage due to, 299 
Galium, Dodder on, 34 
Game, Barking by, 241, 249 
Gas, Coal, Effects of, 28 1 
Gastropacha pini, 218 
Genista, Dodder on, 34 
Germ -pore, 159 

tube, 40 
Germination of seed, 279 
Glceosporium ampelophagum, 104 

nervisequium, 140 
Gnomonia erythrostoma, 88 
Gonidia of Peronospore^e, 57 
Gonidium defined, 45 
Grafting, 265 

Wax as a dressing for wounds, 260 
Gramineae, Ergot on, 98 

Rust of, 155 
Grape, see Vine 

Disease, 70 
Grapholitha pactolana, 89 
Grasses, see Graminea; 
Green-rot, 42, 224 
Growth-borer, Pressler's, 18 

in height, how limited, 7 
Gymnoasce£E, 131 
Gymnosporangium, 51, 157 

clavariteforme, 158 

conicum (Juniperum), 157 

Sabin^e (fuscum), 158 

tremelloides, 1 59 

Hail, Effects of, 299 
may predispose to disease, 15, 118, 
Hallier's early work, 4 
Hardening of trees, 283 
Hares, Barking by, 241 

in regard to potato disease, 65 
Hartig's, R., chief works, 4 
Haustoria of fungi, 42 

PeronosporejE, 57 
on the roots of Phanerogams, 
24, 34 
Hawthorn, see Crataegus 
Hazel, Dodder on, 34 

Erysiphe guttata on, 70 

Nectria ditissima on, 92 

Sphasrella on, 88 
Healing in general, 225 
Heat-requirements of fungi, 46 
Height-growth, how limited, 7 
Helotium Willkommii, 117 
Hemp, Parasite of, 25 
Heredity in relation to disease, 15, 16 
Herpotrichia, an example of mycelial 
infection, 47 

nigra, 76 
Hetercecious fungi, 49, 154, 176 
Honey Agaric, 207 

Dew, 98 
Honeysuckle, see Lonicera 
Hop, see Humulus 
Hornbeam, Exoascus on, 135 

Melampsora on, 171 

Nectria ditissima on, 92 
Hornbeams, Fissures in the cortex of, 

Host, Watery condition of, favourable 
to parasites, 13 

Active growth of, unfavourable to 
parasites, il 
House-fungus, see Merulius lacrymans 
Humulus, Dodder on, 34 
Hyacinths, Hyphomyces on, 38 

Bacteriosis of, 37 
Hydnum diversidens, 202, 258 
Mycelial growths of, 42 
Hymenomycetes, 184 

Klycelial growths of, 42 
Hypha;, see also Mycelium 

action on calcium oxalate, 52 

Characteristic action of, 52 

Septate, 40 
Hyphomyces on hyacinths, 38 
Hypoderma, 107 
Hysterium, 107 

Effects of iodine on, 42 

resemblance to Rhytisma, 106 

Y 2 



Hysterium brachysporum, 117 
laricinum, 1 17 
macrosporum, 51, 109, 177 
nervisequium, loS 
Pinastri, 1 10 

Immersion in water, Effects of, on 

timber, 214, 215 
Individual predisposition to disease, 12 
resistance to disease, S 
rate of growth, 22 
Infection, Animal agency in, 48 
experiments, 17 — 20, 120 
by mycelium, 47, 120 
spores, 48, 49 
Injuries due to fire, 300 

Phanerogams, 23 
plants, 22 

precipitations and gales, 299 
Mechanical, 299 
Insects, Effects of defoliation by, 218, 

may predispose to disease,! 5,1 17 
Intercellular growth of endophytes, 42 
Intermediary growth, 26, 238 

tissue, 228, 266 
Intracellular growth of endophytes, 42 
Ivy, Orobanche on, 25 

Juniperus, Gymnosporangium on, 157 
Sabina; on, 158 
Herpotrichia nigra on, 76 
communis, Gymnosporangium 
clavariaeforme on, 158 
conicum on, 157 
tremelloides on, 1 59 
Oxycedrus, Arceuthobium on, 30 

Knots, 250 

Kiihn's contributions to plant-path- 
ology, 3 

Laburnum, see Cytisus Laburnum 
Lachnus exsiccator, 96 
Larch, Cteoma on, 166, 169 

Hysterium on, 117 

Polyporus on, 199, 200 

Rhizina undulata on, 125 

Aphis, 118 

Blister, 117 

Disease, 117 

in relation to moisture, 46 
pure woods, 122 
Larix europjea, see Larch 
Lathrjea, a partial parasite, 24 
Lead in smoke, 301 
Leaders, Double, Removal of, 261 
Leaf-Cast, 113 

Leaves, Natural shedding of, 225 
Premature withering of, 297 
Shedding of, owing to frost, 2S6 
Ledum palustre, Chrysomyxa on, 179 
Leguminosie, Tubercles on roots of, 39 
Letters carved in bark, 246 
Lichens on trees, 35 
Liebig's discoveries erroneously ap- 
plied, 2 
Life, Prolongation of, by slips, 7 
Light, Effects of excessive, 298 

restricted, 294, 298 
" Lightening " woods. Effects of, 274 
Lightning, Effects of, 302 
Lime, Nectria cinnabarina on, 96 
ditissima on, 92 
Sphccrella on, 88 
Limits of this treatise, 5 
Linum usitatissimum. Dodder on, 34 
Liparis monacha, 218 
Litter, Consequences of removing, 

from a wood, 5, 270 
Local predisposition to disease, 9 
Lonicera Periclymenum, Damage done 
by, 23, 24 

Erysiphe guttata on, 70 
Loranthacea;, Partial parasitism of, 25 
Loranthus europasus, 25, 30, 31 
Louse Wort, Partial parasitism of, 23 
Lucerne, Dodder on, 34 

Orobanche rubens on, 25 
Rhizoctonia violacea on, 82 

Maize, Ustilago on, 68 

Mai nero, 82 

Malformation, Difficulty of defining a, 5 

Malformations induced by fungi, 51 

Maple, see Acer 

Maple, Norway, see Acer platanoides 

Meadow grasses, Ustilago Carbo on, 68 

Medullary rays contain living cells, 46 

Melampsora Arias, 171 

betulina, 171 

Caprearum, 171 

Carpini, 171 

epitea, 171 

Goeppertiana, 49, 161, 184 

Hartigii, 170 

mixta, 171 

Padi, 171 

populina, 165 

salicina, 170 

Sorbi, 171 

Tremula:, 164 
Laricis, 169 
pinitorquum, 9, 12, 166 

Vaccinii, 171 



Melampyrum arvense, a partial para- 
site, 24. 
Merulius lacrymans, 19S, 216, 219 

Mycelial growths of, 42 
Mice, Barking by, 243, 249, 300 

injuring roots, 264 
Micella, 287 
Micrococcus, 12S 
Mildew, see also Erysiphe 

Epiphytic character of, 42 

of Vine, 65 
Millet-brand, 68 
Mistletoe, Common. 25—30 

Oak, 30 
Mites, on Black Austrian pine, 140 
Mixed woods in relation to disease, 

Mock plums, 131 
Moisture in relation to fungus-growth, 

Monosticha oxystoma, 151 
Monotropa Hypopitys, Doubtful para- 
sitism of, 25 
Monstrosity, Difficulty of defining a, 5 
Morbo bianco, 82 
Moth, Pine Beauty, 175 

Nun, 218, 294 
Moth-wither, 175 
Mus sylvaticus, Barking by. 243 
Mycelial nest, 1S9 
Mycelium, see also Hyphas 

of fungi, 40 

Cell-sap in the, 40 

Oil or fat in the, 40 

Resting, 43 

Strandlike, 43 
Mycorhiza, 71 
Myxomycetes, 39 

Nectria, 89, 150, 260 

Facultative parasitism of, 47 

cinnabarina, 96 

Cucurbitula, 89 

ditissima, 91, 299 

the possible cause of "blight,'' 
Normal predisposition to disease, 9 
Norway maple, see Acer platanoides 
Nostoc communis, 1 59 

Oak, see also Ouercus 

Aglaospora Taleola on, 99 
Daedalea on, 206 
Dematophora necatrix on, 82 
Erysiphe guttata on, 70 
Exoascus cajrulescens on, 135 

Oak, Green-rot on, 42 

Hydnum diversidens on, ?02 
Nectria ditissima on, 91 
Peziza aeruginosa on, 224 
Polyporus dryadeus on. 201 

fomentarius on, 206 

igniarius on, 201 

sulphureus on, 200 
Rosellinia on, 78 
Sphasrella on, 88 
Stereum hirsutum on, 205 
Thelephora Perdix on, 203 
Tuber on, 71 
Oaks, Predisposition of, to top-drought, 


Rupturing of the cortex of, 273 
Oats, Puccinia coronata on, 156 

Ustilago Carbo on, 68 
Obligative parasites, 47 
Occlusion of wounds, 229 
Oidium Tuckeri, 70 
Olive, Bacterial disease of, 38 
Oogonia of Peronosporeae, 58 
Oospores, Vitality of, 45 

of Peronosporea;, 58 
Orchidacea.-, Parasitism of, 25 
Organisms, Low, in relation to death, 6 
OrobanchacecC, true parasites, 25 
Orobanche, Parasitism of species of, 

Outgrowths on Alder-roots, 39 

Paraphyses, 108 

Parasite, Definition of the term, 46 

Difficulty of defining a, 23 
Parasites, Facultative, 46 

Obligative, 46 

Partial, 24 

Pseudo, amongst cryptogams, 35 
Parenchymatous cells scarce in wood, 

Partridge wood, 203 
Pathology, Vegetable, Development of, 

Pear, see also Pyrus 

Erysiphe guttata on, 70 

Polyporus sulphureus on, 200 

Probable bacterial disease of, 38 

Exoascus bullatus on, 133 

Stigmatea on, 88 
Pedicularis, a partial parasite, 24 
Perennials, Partial death of, eachvear, 

Periderm, 225 
Peridermium Cornui, 175 

elatinum, 179 

oblongisporium, 172 



Peridermium Pini, 172, 175 

in regard to turpentine, 51 
acicola, 172 
corticola, 172 
Strobi, 175 
Peridium of Uredineaa, 153 
Perithecium of Pyrenomycetes, 72 
Peronospora Sempervivi, 58 

viticola, 65 
Peronosporese, 57 
Persica vulgaris, Exoascus deformans 

on, 132 
Pestalozzia Hartigii, 136 
Peziza aeruginosa, 42, 224 
calycina, 117, 139 
ciborioides, 130 
Fuckeliana, 130 
Sclerotiorum, 130 
Willkommii, 117, 299 

in regard to moisture, 46 
Pezizje, 1 17 
Phacidea;, 105 

Phanerogams, Injuries due to, 23 
Phelloderm, 226 
Phellogen, 226 
Phoma abietina, 138 
Phragmidium incrassatum, 1 56 
Rubi Idfea, 156 
subcorticum, 156 
Phycis abietella, 175 
Phycomycetes, 57 

Formation of spores by, 44 
Phyllactinia, 70 
Phylloxera vestatrix, 83 
Physalopsora Bidwellii, 103 
Phytophthora Fagi, 58 
infestans, 36, 64 
omnivora, 58 

a facultative saprophyte, 47 
its action on starch, 52 
its manner of spreading, 49 
vitality of oospores of, 45 
Picea, see also Spruce. 

Menziesii, Septoria parasitica on, 

Sitkaensis, Rhizina undulata on, 
Pileus, 35 

Pine, Dematophora necatrix on, 82 
Elaphomyces granulatus on, 71 
Hysterium pinastri on, no 
Nectria Cucurbitula on, 89 
New parasite of, 147 
Rosette shoots of, 240 
Black Austrian, Disease of, 140 
-Mountain, Herpotrichia nigra on, 

Pine, Scotch, see Pinus sylvestris 
Weymouth, see Pinus Strobus 
Beauty moth, 175 
leaf blight, 1 1 1 
twist, 166 

in regard to aspens, 9 
moisture, 46 
Pinus halepensis. Bacterial disease of, 

Gymnosporangium Sabins on, 

rigida in regard to stool-shoots, 

Strobus, Bark-drought of, 295 
Hysterium brachysporum on, 

Peridermium on, 175 
Polyporus Schweinitzii on, 198 
Restocking diseased areas with, 

Rhizina undulata on, 125 
Withering of the leaves of, 
II t 
sylvestris, see also Pine 

Peridermium Cornui on, 175 
Polyporus Schweinitzii on, 
Plant-food in soil, 270 
Planting, Deep, 278 
Plants, Importance of rejecting weak, 

Injuries due to, 20 
Plasmodiophora Alni, 39 

BrassicK, 39 
Plasmolysis, 288 
Platanus, Glceosporium nervisequium 

on, 140 
Plowrightia morbosa, 102 
Plum, see also Prunus 
Black-knot of, 103 
Plums, ]\Iock, Pocket, or Starved, 

Poa pratensis. Bunt on, 68 
Pocket plums, 131 
Poisons, Plant, 279, 301 
Polyporei, Preventive measures 

against, 206 
' Polyporus betulinus, 206 • 
borealis, 42, 55, 196 
dryadeus, 201 < 
^' fomentarius, 206 
fulvus, 194 ., 
Hartigii, 180, 194 
v> igniarius, 51, 201 >, 
l^evigatus, 206 
mollis, 198 
Schweinitzii, 53, 56, 198 



^ Polyporus sulphureus, 42, 52, 53, 
200 y 

vaporarius, 42, 19S, 199, 213 
Polystignia fulvum, 98 
ochraceum, 98 
rubrum, 97 
Poplar, see also Populus 

Lombardy,Didymosphi'eria 011,104 
Populus, Dodder on, 34 

Erysiphe adunca on, 70 
alba, Exoascus aureus on, 135 
nigra, „ „ 135 

tremula, „ „ 135 

Potato disease, 64 

Potatoes, Dematophora necatrix on, 

Engrafting, 267 
Reserve materials in, 40 
Resistance of, to disease, 12 
Pourridie de la Vigne, 82 
Pourriture, 82 
Predisposition to disease, 8 
Abnormal, 9, i 5 
Climatic, 10 
Individual, 12, 13 
Induced by hail, 15 

conditions of growth, 14 
insects, i 5 
soil, 10 
Local, 9 
Normal, 9 
Seasonal, 11 
Temporary, 10 
to frost, 14 
Pressler's growth-borer, 18 
Preventitious buds, 238 
Preventive measures against parasites, 

Progress, Organic, depends on varia- 
tion, 5 
Promycelium of smut-spores, 66 
Pruning, 249 

of roots, 264 
Prunus, Polystignia rubrum on, 97 
avium, Agaricus on, 207 
Exoascus deformans on, 132 
Polystignia ochraceimi, 98 
Cerasus, E. deformans on, 132 
Chamaecerasus, ., 132 

domestica, Agaricus on, 207 
Exoascus deformans on, 132 
Pruni on, 131 
instititia, Exoascus on, 133 
Padus, „ Pruni on, 131 

Melampsora on, 171 
Nectria ditissima on, 91 
Polystigma fulvum on, 98 

Prunus spinosa. Dodder on, 34 
Exoascus Pruni on, 131 
Polystigma rubrum on, 98 
Valsa on, 88 

Pseudo-parasites amongst cryptogams, 

Pseudotsuga Douglasii, see Fir, 

Puccinia Asparagi, i 56 

coronata, 156 

graminis, 155 

straminis, 1 56 

striieformis, 156 
"Pugging," 217 
Pure woods specially liable to disease, 

I, II 
Pycnidium of Cucurbitaria, 88 
Pyrenomycetes, 69, 72 
Pyrus, see also Apple and Pear 

R^estelia cancellata on, 158 
Pythium de Baryanum, 66 

(^uercus, see also Oak 

Cerris, Loranthus on, 31 
Quickens, see Triticum repens 
()uinine, 230 

Rabbits, Barking by, 241 
Red-rot of birch, 206 

spruce known to early writers, 
Trametes radiciperda, a cause 
of, 186 
"stripe," 215 
Rejuvenescence, Annual, of trees, 7 
Remedial measures against parasites, 

Reserve materials m regard to prun- 
ing, 260 
Resin-ducts, 232 

tlux, 210 

glut, 210 

leader, 175 

top, 175 
Resinous degeneration, Early mention 

of, I 

saturation of wood, -234 
Resistance to disease. Individual, 7 
Resting-myceliuni, 43 
Rhaninus, ^Ecidium on, 156 
Rhinantheje as partial parasites, 

Rhinanthus Crista-galli, a partial para- 
site, 24 
Rhizina undulata, 123 
Rhizoctonia violacea, 82 



Rhizoctoniae, 43, 48, 78 
Rhizoids, 26 
Rhizomorphs, 43, 48 
Rhododendron, Chrysomyxa on. 177 
Rhododendrons and spruce-leaf blister, 


Rhytisma acerinum, 70, 105 

punctatum, 106 

salicinum, 107 
Ribes, Caeoma on, 171 
Robinia, Polyporus sulphureus on, 200 

seedlings, Disease of, 58 
Roesleria hypog^ea, 83 
Rsestelia cancellata, 158 

cornuta, 1 58 

lacerata, 158 
Root-fungus of conifers, 123 

rot of vine, 82 

suckers, 241 
Roots, adventitious, 240, 264, 279 

Eftects of heaping earth on, 278 

Injuries to, 264 

Natural engrafting of, 262 
Rose, ]\Iildew of, 70 

Rust of, ■ 1 56 
Rosellina quercina, 78 

an example of mycelial infection, 

Effect of iodine on, 42 
in relation to water, 1 2 
Mode of attack of, 50 
Sclerotia of, 43 
Rosette shoots on the pine, 240 
Rot, Dry, 214 

Green, 42, 224 

Red, I, 186, 244 

Root, 276 

White, 195 

Wound, 236, 244, 247, 257, 262, 

Rubus cassius, Phragmidium incrassa- 
tum on, 1 56 

fruticosus, Phragmidium incrassa- 

tum on, 1 56 
IdfEus, Phragmidium incrassatum 

on, 156 
Rust, Spruce-leaf blister, 175 
of Asparagus, i 56 

Crucifers, White, 66 

Larch, 169 

Pine, 172 

Poplar, 164 

Rhododendron, 177 

Rose, 156 

Spruce, 175 

^^'heat, 48, 155 

Colour of, 40 

Rust, Protection against drought of, 45 

Fungi, 153 
Rye, Puccinia striaeformis on, 1 56 

Brand, 68 
Sac Fungi, 69 

Saffron, Rhizoctonia violacea on, 82 
Salix, see also Willow 

aurita, Rhytisma salicinum on, 107 

Caprea, ,, ,, 107 

nigricans, „ „ 107 

purpurea, „ ,, 107 

Saprolegiacea;, 57 
Saprophyte, the term defined, 46 
Saprophytes, Facultative, 47 
Schinzia Alni, 39 
Schizomycetes, 37 
Sclerotia, 43, 48 
Sclerotinia, 129,130 

baccarum, 130 

Libertiana, 130 

megalospora, 130 

Oxycocci, 130 

Vaccinii, 130 
Scorching of bark by fire, 300 

sun, 300 
Scrophulariaccce as partial parasites, 24 
Sea-water, Effect of, on trees, 20, 280 
Seed from stunted pines, 16 

Germination of, 44 

Importance of care in selecting, 
13, 22 
Seeding, Effects of thick, 299 
Seedling beech disease. Early mention 
of, 58 

conifers. Effects of drought on, 112 

Parasitic disease of, 147 

pines, Caeoma on, 166 
Sempervivum seedlings. Disease of, 

5S . 
Senecio, Coleosporium on, 172 
Septoglffium Hartigianum, 141 
Septoria parasitica, 143 
Sexual generation, 44 

processes in fungi, 44 
Shake — Bark, Ring, or Heart, 191 
Shedding of pine-leaves, 1 1 1 
Shoots, Stool, 261, 262 
Shortening of branches, 259 
Shrinkage of wood due to frost, 284 
Sickliness and disease distinguished, 5 
Sinkers of Mistletoe, 26 
Sirex, 248 
Slime Fungi, 39 
Slips, 264 

may prolong life, 7 
Sloe, see Prunus spinosa 
Smoke, Damage due to, 300 



Smoke, Special liability to injury from, 

Smut, Formation of spores of, 66 
Popular use of the term, 66 
of Wheat, how spread, 49, 67 

Smuts, Protection of. against drought, 


Snags of branches. 254, 260 

Snow, Damage due to, 299 

Soda-fumes, Effects of, 300 

Soil-canker, 129 

in relation to disease, 10, 270 

Solutions in soil, Effects of strong, 

Soot, Effects of, 300 

Sorauer's work. 4 

Sorbus, ^cidium pencillatum on, i 59 
IMelampsora on, 171 
Rjestelia cornuta on, 15S 
Aucuparia. Cucurbitaria on, 88 

Spear-top, 175 

Sfthacelia segetum, 99 

Sphaceloma ampelinum, 104 

Sph^erella, Leaf-blotches due to, 88 
Fagi, 88 
Fragaria?, 88 
maculiformis, 88 
punctiformis, 88 

Sphaeria dryina, 224 

Sphasrotheca pannosa, 70 

Spheroblasts on the beech, 239 

Sporangia of Peronosporea;, 57 

Spore-mother-cells, 162 

Spores, how formed, 43 
Germination of, 45 
Infection by, 48 

Sporidia, 154 
of smut, 67 

Sporocarp of Basidiomycetes, 153 

Sporophore of Fungi, 43 
PeronosporccC, 57 
Pezizffi, 117 

Spruce, see also Picea 

yEcidium abietinum on, 177 

coruscans on, 183 

Chrysomyxa abietis on, 175 

Ledi on, 179 
Dematophora necatri.x; on, 82 
Green-rot on, 42, 224 
Herpotrichia nigra on, 76 
Hysterium macrosporum on, 109 
Nectria Cucurbitula on, 89 
Parasitic disease of, 147 
Pestalozzia Hartigii on, 136 
Peziza aeruginosa on, 224 
Polyporus borealis on, 196 
Trichospheeria parasitica on, 72 

Spruce, Withering of leaves of, in 

cones, /Ecidium strobilinum on, 

conorum Piceaeon, 183 

leaf blister, 9, 13, 175 
redness, 109 
rust, 175 
Spruces as a protection against pine- 
blight, 116 
Stag-headed condition, 270 
Starved plums, 131 
Stem-brand. 68 
Stereum hirsutum, 205 
Sterigmata, 142 
Sticky-brand, 68 
Stigmatea, Leaf-blister due to, 88 

Alni, 88 

Mespili, 88 
Stink-brand, 68 

Stone-fruit trees. Black-knot of, 103 
Strawberrry, Sphaerella on, 88 
Stumps of branches, 254, 260 
Sucker-tubercles of fungi, 42 
Sulphurous acid in smoke, 301 
Sun, injurious effects on conifers, 112 

cracks, 283, 294, 296 
Swarm-spores of Peronosporea;, 57 
Swellings induced by fungi, 50 
Sycamore, see Acer 
Symbiosis, 71, 181 

Tannin as food for fungi, 50 
Taphrina aurea, 135 

betulina, 133 

Populi, 135 
Tar-girdles, Effect of, on j^ines, 246 
Tarring wounds, 258 
Teleutospore, 154 
Temperature of trees, 283 
Teratology defined, 5 
Thelephora laciniata, 35 

Perdix, 54, 203 
Thinning woods, Effects of, 271 
Thrushes as distributors of mistletoe, 

26, 30 
Thunder brooms, see Witches' brooms 
Thymus, Dodder on, 35 
Tidiness, Importance of, 57 
Tilletia Caries, 68 

laevis, 68 
Timber, Coniferous, Blue colour of, 224 

Structural, Attack of fungi on, 212 
Merulius lacrymans in, 219 
Tinder-fungus, 206 
Tinea sylvestrella, 175 
Toad-stools, 46 
Tobacco, Parasite of, 25 


Tooth Wort, a partial parasite, 24 
Top-drouglit or Top-drying, 5, 14, 20, 

Town trees. Unhealthy condition of, 

Trametes Pini, 191 

Action of ferment of, 53 
Mycelial growths of, 42 
radiciperda, 186, 277 

an example of mycelial infec- 
tion, 48 
associated with root-rot, 21 
may pierce suberose tissues, 50 
may spread by spores, 49 
Transplantation in regard to frost, 14, 

Trenches as a preventive measure, 57 
Trichosphteria, an example of mycelial 
infection, 47 

parasitica, 72 
Triticum repens, Bunt on, 68 

Damage due to, 24 
Tsuga Mertensiana, Rhizina undulata 

on, 125 
Tuber, 71 
Tuberacei, 69, 71 
Tubercularia, 89 
Tulasne's work, 3 
Turgescence, 288 
Turpentine, Strasburg, 247 

Venetian, 247 
Twigs, Shedding of, 225 
Twitch, see Triticum repens 
Tyloses, 228, 235, 236 

Uncinula Aceris, 70 

adunca, ']o 
Uredine^e, 153 

as obligative parasites, 46 
Uredo linearis, 155 
Uredospores, 154 
Urine, Effects of, on plants, 281 
Urocystis Anemonis, 68 

Cepulee, 68 

occulta, 68 

Violas, 68 
Urtica, Dodder on, 34 
Ustilaginete, 57, 66 
Ustilago Carbo, 68 

destruens, 68 

Maydis, 68 

Vaccinium, Exobasidium on, 185 
Melampsora on, 171, 185 
Myrtillis, Sclerotinia baccarum 

on, 130 
Oxycoccus, Sclerotinia on, 130 

uliginosum, Sclerotinia megalo- 
spora on, 130 
Vaccinium Vitis Idaea, Effects of 
Calyptospora on, 51 
Melampsora Goeppertiana on, 
Valsa oxystoma, 151 

Prunastri, 88 
Variation in relation to development, 5 
Possible directions of, 12 
probably initiated in the oosphere, 


Vine, see also Grape 

Botrytis cinerea on, 130 
Coniothyrium diplodiella, 103 
Dematophora necatrix on, 82 
Dodder on, 34 
Glceosporium on, 104 
Physalopsora Bidwellii on, 103 
Mildew, 65 

Viscum album, 25 

Walnut, Polyporus sulphureus on, 200 
Water, Contaminated, Effects of, 281 

Salt, Effects of, 280 

Stagnant, Effects of, 276 

in soil, 270 
Weinstockfaule, 82 
Wet-rot of potato, 64 
Wheat, Bunt of, 68 

Puccinia striasformis on, 156 

Ustilago Carbo on, 68 

rust in relation to barberrv, 9, 

\\ hite piped wood, 205 
rot of birch, 206 
conifers, 21 1 
fir and spruce, 195, 196 
oak and beech, 203, 205 
vine, 104 
Willkomm's pioneer work, 4 
Willow, see also Salix 
Dodder on, 34 
Melampsora on, 171 
Mildew of, 70 

Polyporus sulphureus on, 200 
Caspian, Behaviour of, on sandy 
soil, 265 
Wind injuring leaves of conifers, 1 1 1 
Witches' brooms, 51 
on alder, 133 
birch, 135 
hawthorn, 133 
Prunus, 131 
silver fir, 179 
Wood, Distinction between dead and 
living, 46 


Wood, " Wound," 230, 231 
balls on the beech, 239 
Works, Early, on plant-diseases, 2, 

3, 4 
Wound-rot, 236, 244, 247, 257, 262, 

\\ ounds, 225 

Agencies that produce, 50 

Effects of, on wood, 232 

General treatment of, 237 

Natural, 225 

Ring, 249 

Varieties of, 241 

Wounds, due to crushing, 246 

resin-collecting, 247 
predispose to disease, 1 5 
Wurzelpilz, 82 

Yellow-piped wood, 205 
Rattle, see Rhinanthus 

Zea, see Maize 
Zinc in smoke, 301 
Zoogonidia of Peronosporea?, 58 
Zygomycetes, 57 
Zygospores, 42 



Books on Botany. 


TREES. By Professor R. HARTIG. Translated by Dr. W. 
SoMERViLLE, Profcssor of Agriculture and Forestry at Durham 
College of Science. With a Preface by Prof. H. Marshall 
W^ARD, F.R.S. With numerous Illustrations. Medium 8vo. 


and Foreign. By THOMAS LASLETT. Timber Inspector to 
the Admiralty. New Edition. Revised by Prof. H. Marshall 
Ward, F.R.S. Crown Svo. \_I?i the press. 


CROPS, chiefly such as are caused by Fungi. By WORTH ING- 
TON G. SMITH, F.LS., M.A.I., Member of the Scientific 
Committee, Horticultural Society. With 143 Illustrations, Drawn 
and Engraved by the Author. Fcap. Svo, 4^". 6d. 


Reviews of Works on Botany and Related Subjects, 1 834-1 887. 
Vol. II. Essays, Biographical Sketches, 1841-1886. 8vo, 21s. 


(Sixth Edition). 

Vol. I. Structural Botany : or Organography on the Basis of 
Morphology. To which is added The Principles of Taxonomy and 
Phytography, and a Glossary of Botanical Terms. By ASA GRAY. 
Svo, 10s. 6d. 

Vol. II. Physiological Botany. I. Outlines of the History of 
Ph^nogamous Plants. II. Vegetable Physiology. By G. LINCOLN 
GOODALE, A.M., M.D., Professor of Botany in Harvard University. 
Svo, 10.T. 6d. 


Products, and Uses. By JOHN SMITH, A.L.S., Author of 
" Historia Filicum," " History of Bible Plants," &c,, &c. Medium 
Svo, 14s. 


Books on Botany. 


By Prof. F. O. BOWER, D.Sc, F.R.S., Author of " A Course of 
Practical Instruction in Botany." Globe 8vo, 35. 6d. 

Regius Professor of Botany in the University of Glasgow. Crown 
8vo, ici". dd. 


BOTANY. By G. T. BETTANY, M.A., B.Sc, F.L.S., late 
Lecturer on Botany in Guy's Hospital Medical School, sometitne 
Examiner in Botany Cambridge Univ^ersity Local Examinations. 
Pot Svo, i^. 

BOTANY. By Sir J. D. Hooker, F.R.S. 

With Illustrations. Pot Svo, \s. \Science Privicrs. 


Edition. Globe Svo, \os. 6d. 


By DANIEL OLIVER, F.R.S. The Part on Systematic Botany 
based upon material left in manuscript by the late Prof. 
Henslow. With numerous Illustrations. Third Edition. Fcap. 
Svo, 4J". 6d. 


By DANIEL OLIVER, F.R.S. With numerous Illustrations. 
Globe Svo, 6^. 6d. 

SCIENCE PAPERS, Chiefly Pharmaco- 
logical and Botanical. By DANIEL HAN BURY, F.R.S. Edited 
with Memoir by Joseph Ince, F.L.S., F.C.S. With Portrait. 
Medium Svo, 145. 


Nature Series. 

Crown 8vo. 

KELVIN, P. R. S. In Three Vols. 
Vol. I. THE CONSTITUTION OF MATTER. Second Edition, -js. 6d. 


With Illustrations. 3^'. 6d. 


ON LINKAGES. By A. B. KEMPE, F.R.S. Illustrated. li-. 6./. 

ON LIGHT. The Burnett Lectures. By Sir GEORGE 
GABRIEL STOKES, F.R.S. Three Courses. L Ox the Nature of 
Light. II. On Light as a Means of Investigation. III. On Bene- 
ficial Effects of Lighi'. 7^. 6J. 


Illustrated. 2s. 6(f. 


F.R.S. Illustrated, ^s. 6d. 




LODGE, F.R.S. Illustrated. 6s. 6d. 


HALL. 45. 6d. 


A. TRIBE. 2s. 6d. 


MELDOLA, F.R.S. Illustrated. 6s. 


Nature Series. 

Crown 8vo. 
CHARLES DARWIN. Memorial Notices reprinted 

from A'a/inr. By THOMAS H. HUXLEY, F.R.S. : G. T- ROMANES, 
F.R.S. ; Sir ARCHIBALD GEHs:iE, F.R.S. ; and W. T. DYER, F.R.S. 
2s. 6d. 




INHERITED? An E.xamination of the View held by Spencer and Darwin. 
By W. PLATT BALL. 3^-. 6d. 




Prof. H. M. WARD, F.R.S. Ilhistratcd. 6s. 


GRANT ALLEN. Illustrated. 3.-. bd. 


JOHN LUBBOCK, M. p., F.R.S. Illustrated. 45. 6tZ. 


Illustrated, j^s. 6d. 


INSECTS. By Sir JOHN LUBBOCK, M-.P,, F.R.S. AVith Illustrations. 
Zs. 6d. 


Illustration^. 7^. 6d. 


By Sir D. WILSON, LL.D., F.R.S.E. Illustrated. 4.,-. 6d 


CLIFFORD, F.R.S., Diagrams, z^. 6d.