DISEASES OF
FIELD AND GARDEN CROPS
FIG. 138.— DARWIN'S POTATO.
Solanum Maglia, Sch.
Tor
DISEASES
OF
FIELD AND GARDEN CROPS
Cfjteflg surfj as are caused fig Jungt
BY
WOBTHINGTON G. SMITH, F.L.S., M.A.I.
MEMBER. OF THE SCIENTIFIC COMMITTEE
ROYAL HORTICULTURAL SOCIETY
WITH ONE HUNDRED AND FORTY-THREE ILLUSTRATIONS
DRAWN AND ENGRAVED BY THE AUTHOR
SINE-FORTlOhE
/SINE -B4VORE •„
MACMILLAN AND CO.
1884
Printed by R. & R. CLARK, Edinburgh.
TO
DR. BULL OF HEREFORD
AND
THE OFFICERS AND MEMBERS OF THE
WOOLHOPE CLUB
WITH THE GRATEFUL AND PLEASANT REMEMBRANCES
OF
THE AUTHOR
PEEFACE.
THE following notes on the Diseases of Field and
Garden Crops are reports of a series of addresses
given at the request of the officers of the Institute of
Agriculture at the British Museum, South Kensington.
The addresses were preceded by a course of twenty
lectures upon Vegetable Physiology in relation to
Farm Crops, by Professor G. T. Bettany, M.A., B.Sc.,
F.L.S.; so that it was unnecessary, in speaking of
plant diseases, to revert in detail to the structural
and physiological branches, which had been covered
by Professor Bettany.
The nature, limits, and objects of the addresses
will be seen in the introductory remarks. In the
lecture room actual examples of all the diseases in
different stages of growth were exhibited, and the sub-
jects were illustrated with camera-lucida drawings, as
well as larger coloured drawings made from living
examples. In the lecture room the simpler subjects
were taken first, the students being gradually led on
to the more involved ones. This arrangement has
been adhered to in the present work, where the
addresses have been put into book form, and numer-
ous additions made.
PREFACE.
The engravings are either reductions from the
large drawings used at the lectures, or from others
made from nature since. They are all (with the
exception of one or two which it was necessary should
be copies) original, and from living examples.
We hope the illustrations will be useful ; we
believe they are correct j they show what we have
seen, or think we have seen.
Excellent preparations for the microscope of the
fungi mentioned in these pages may be purchased, at
moderate prices, from the Eev. J. E. Vize, M.A.,
Forden, Welshpool.
Our thanks are due for assistance to our friends
Mr. Chas. B. Plowright, M.R.C.S., King's Lynn;
and to Mr. A. Stephen Wilson of North Kinmundy,
Summerhill, Aberdeen.
Most of the original camera-lucida drawings, with
many of the actual examples, and a large series of
microscopic preparations used at the lectures and for
this work, are now in the department of Botany,
British Museum (Natural History), South Kensington.
W. G. S.
LINDEN HOUSE,
DUNSTABLE,
BEDS.
CONTENTS.
CHAPTER I.
PAGE
INTRODUCTORY 1
CHAPTER II.
CLOVER SICKNESS — CLOVER MILDEW. — Peronospora tri-
foliorum, D.By 6
CHAPTER III.
PERONOSPORA EXIGUA, W.SM. . . . .12
CHAPTER IV.
NEW DISEASE OF POTATOES. — Peziza postuma, Berk, and
Wils 15
CHAPTER V.
FUSISPORIUM DISEASE OF POTATOES. — Fusisporium
Solani, Mart 30
CHAPTER VI.
SMUT OF POTATOES. — Tubercinia scabies, B. ... 35
xii CONTENTS.
CHAPTER VII.
PAGE
SCAB AND CRACKING OF POTATOES 37
CHAPTER VIII.
NEW DISEASE OF ONIONS. — Puccinia mixta, Fl. . . 39
CHAPTER IX.
MILDEW OF ONIONS. — Peronospora Schleideniana, Ung. . 45
CHAPTER X.
MOULD OF ONIONS. — Mucor suUilissimus, B. . . .51
CHAPTER XL
ONION SMUT.— Urocystis cepulce, Far 54
CHAPTER XII.
NEW DISEASE OF GRASS. — Isaria fuciformis, Berk. . 55
CHAPTER XIII.
STRAW BLIGHT 69
CHAPTER XIV.
SURFACE MILDEW OF TURNIPS. — Oidium Balsamii, Mont. 75
CONTENTS.
CHAPTER XV.
PAGE
PUTREFACTIVE MILDEW OF TURNIPS AND CABBAGES. —
Peronospora parasitica, Pers. . . . .80
CHAPTER XVI.
WHITE-RUST DISEASE OF CABBAGES, ETC.—Cystopuscan-
didus, Lev. . 86
CHAPTER XVII.
CLUB- ROOT OF TURNIPS, CABBAGES, MANGELS, AND
ALLIED PLANTS. — Plasmodiophora Brassicce, Wor. . 94
CHAPTER XVIII.
EAR- COCKLE, PURPLES OR PEPPERCORN IN WHEAT,
OATS, AND RYE. — Tylenchus tritici, Bast. . .105
CHAPTER XIX.
CLOVER DODDER. — Cuscuta Trifolii, Bab. . . . 115
CHAPTER XX.
GRASS MILDEW. — Erysiphe graminis, D.C. . . .126
CHAPTER XXI.
CORN MILDEW — SPRING RUST AND MILDEW. — Puccinia
RuUgo-vera, D.C 135
xiv CONTENTS.
CHAPTER XXII.
PAGE
BORAGE BLIGHT. — sEtidium asperifolii, Pers. . . 143
CHAPTER XXIII.
CORN MILDEW — SUMMER KUST AND MILDEW. — Puccinia
graminis, Pers. . . . . . . .147
CHAPTER XXIV.
BARBERRY BLIGHT. — dEddium Berleridis, Pers. . . 159
CHAPTER XXV.
THE POSSIBLE CONNECTION OF THE ^FUNGUS OF CORN
MILDEW AND THE FUNGUS OF BARBERRY BLIGHT . 169
CHAPTER XXVI.
NEW DISEASES OF WHEAT, BARLEY, AND RYE -GRASS
CAUSED BY Fusisporium culmorum, hordei, and Lotii,
W.Sm 208
CHAPTER XXVIL
ERGOT. — Claviceps purpurea, Tul. ..... 214
CHAPTER XXVIII.
WILSON'S VARIETY OF CLAVICEPS ON ERGOT. — Claviceps
purpurea, Tul., var. Wilsoni, W.Sm. . . . 233
CONTENTS.
CHAPTER XXIX.
PAGE
MILDEW OF PARSNIPS. — Peronospora nivea, Ung. .. . 239
CHAPTER XXX.
BUNT OF WHEAT. — Tilletia Caries, Tul. . . . 245
CHAPTER XXXI.
SMUT OF CORN. — Ustilago carlo, Tul 254
CHAPTER XXXII.
TARE OR VETCH, AND PEA MOULD. — Peronospora vicice,
Berk 263
CHAPTER XXXIII.
PEA MILDEW. — Emjsiplie Martii, Lk 266
CHAPTER XXXIV.
LETTUCE MILDEW. — Peronospora ganglioniformis, Berk. 269
CHAPTER XXXV.
POTATO DISEASE, I. — Peronospora infestans, Mont. — Its
Active State , 2'5
CONTENTS.
CHAPTER XXXVI.
PAGE
POTATO DISEASE, II. — Peronospora infestans, Mont. — Its
Passive State . 295
CHAPTER XXXVII.
PARASITIC FUNGI AS FOUND IN A FOSSIL STATE . . 330
CHAPTER XXXVIII.
CONCLUSION . 336
LIST OF ILLUSTBATIONS.
CLOVER MILDEW.
FIG. PAGE
1. Peronospora trifoliorum, D.By., x 400. Conidium of
do., xlOOO 10
2. Peronospora exigua, "W.Sm., x 400. Conidium of
do., xlOOO 12
NEW DISEASE OF POTATOES.
3. Potato steins with sclerotia of Peziza postuma, B. and
Wils. Natural size 17
4. Sclerotium of Peziza postuma, B. and Wils., x2 . 18
5. Microscopic structure of the sclerotium of Peziza
postuma, B. and Wils., x 400 . . . .19
6. Peziza postuma, B. and Wils. Natural size . . 25
7. Section through cap of Peziza postuma, B. and Wils.
Enlarged 10 and 20 diameters .... 26
8. Section through fragment of cap of Peziza postuma,
B. and Wils., x400 27
9. Asci, paraphysis, and sporidia of Peziza postuma, B.
and Wils., x 500 27
FUSISPORIUM DISEASE OF POTATOES.
10. Fusisporium Solani, Mart. , x 400 . . . .32
11. Spores of Fusisporium Solani, Mart., in different
stages of growth, x 1000 33
NEW DISEASE OF ONIONS.
12. Puccinia, mixta, Fl., on flower scape of chives, x5 . 40
13. Section through a sorus of Puccinia mixta, Fl., x 200 41
14. Teleutospores of Puccinia mixta, Fl., x 1000 . . 42
b
xviii LIST OF ILLUSTRATIONS.
MILDEW OF ONIONS.
FIG. PAGE
15. Peronospora Schleideniana, Ung., x 200 ; resting-spores,
x400; conidmm, x 1000 46
16. Peronospora Schleideniana, Ung., part of conidiophore
and conidia, x 400 48
NEW DISEASE OF GRASS.
17. Panicle of sheep's fescue grass invaded by fsaria
fuciformis, Berk., x 2 56
18. Stems of sheep's fescue grass, with Isaria fuciformis,
Berk, x5 57
19. Tip of one of the minor branches of Isaria fuciformis,
Berk., x 100 58
20. Tip of branchlet of Isaria fuciformis, Berk., x 1000 58
21. Living wasp, with fungus growths, x 2 . . .59
22. Torrubia ophioglossoides, Tul. ; parasitic on Elapho-
myces variegatus, Yitt. Natural size, and x 5, 200,
and 1000 61
23. Mnium hornum, Hedw., invaded by mycelium of
Torrubia ophioglossoides, Tul. Natural size . . 64
24. Parasite of Isaria fuciformis, Berk. ; Saprolegnia
philomulces, W.Sm., x 400 68
STRAW BLIGHT.
25. Fragments of diseased wheat stems, x 5 . . .69
26. Fragment of wheat stem with fungus growth,
x200 71
SURFACE MILDEW OF TURNIPS.
27. Oidium Balsamii, Mont. , x 400 .... 77
28. Spore of Oidium Balsamii, Mont., germinating on
fragment of turnip leaf, x 1000 . . . . 78
PUTREFACTIVE MILDEW OF TURNIPS AND CABBAGES.
29. Peronospora parasitica, Pers., x 200 . . . 81
30. Conidmm and part of conidiophore of Peronospora
parasitica, Pers., x 1000 83
31. Oospores or resting-spores of Peronospora parasitica,
Pers., x400 84
LIST OF ILLUSTRATIONS.
WHITE RUST DISEASE OF CABBAGES.
FIG. PAGE
32. Cystopus candidus, Lev. , x 400 ; with conidium and
zoospores x 1000 88
33. Reproductive organs and oospores of Cystopus can-
didus, Lev. 91
CLUB-BOOT OF TURNIPS, CABBAGES, MANGELS, AND
ALLIED PLANTS.
34. Clubbed-root of young turnip, one-half the natural
size 97
35. Section through a small club, x 10 . . . . 98
36. Section through fragment of turnip-root, showing the
plasma of Plasmodiophora Srassicce, Wor. , x 200 . 98
37. Section through fragment of turnip -root showing
spores of Plasmodiophora Srassicce, Wor., x 200 . 100
38. Spores of Plasmodiophora Brassicce, Wor. ; and rest-
' ing-spore of Peronospora parasitica, Pers., x 1000 101
39. Germinating spores of Plasmodiophora Srassicce,
Wor., xlOOO 101
EAR-COCKLE, PURPLES OR PEPPERCORN IN WHEAT,
OATS, AND RYE.
40. Spikelet of wheat, x2 106
41. Horizontal section through spikelet of wheat, x 4 . 106
42. Grain of wheat enclosed in its pale, x5 . . . 108
43. Scale or lodicule of wheat grain, x 50 . . .108
44. Spikelet of wheat, the grains replaced by the galls
of Ear-Cockle, x2 109
45. Galls of Ear- Cockle from a wheat spikelet, x 5 . 110
46. Fragment of wall of gall of Ear-Cockle with Nema-
toid worms, Tylenchus tritici, Bast., in situ, x 40 112
CLOVER DODDER.
47. Cuscuta Trifolii, Bab., growing on clover, x2 . 116
48. Seeds of red clover, yellow trefoil, Dutch clover, and
clover Dodder, x 5 117
49. Seeds of perennial red clover and clover Dodder, seen
in section, arid germinating, x 10 . . .118
LIST OF ILLUSTRATIONS.
FIG. PAGE
50. Clover and Dodder seeds germinating, x 2 . . 119
51. Germinating Dodder seed, fixing on seed leaves of
clover, x 10 120
52. Infant Dodder plant on young clover leaf, x 10 . 120
53. Fragment of clover stem, with Dodder entwined,
showing connection of latter by suckers, x 15 . 122
54. Anatomical connection of clover Dodder with clover
stem, x50 124
GRASS MILDEW.
55. "Wheat stem invaded by Erysiphe graminis, D.C., x2 127
56. Oidium monilioides, Lk. The early condition of
Erysiphe graminis, D.C., x 400. Germinating
conidium, x 1000 127
57. Conceptacle of Erysiphe graminis, D.C., x 100 . 129
58. Horizontal section through conceptacle of Erysiphe
graminis, D.C., x 200 130
59. Vertical section through conceptacle of Erysiphe
graminis, D.C., x 100 131
60. Asci and sporidia of Erysiphe graminis, D.C., x 500.
Germinating sporidium, x 1000 .... 132
61. Conceptacle of Erysiphe graminis, D.C., bursting in
spring, xlOO 133
CORN MILDEW — SPRING RUST AND MILDEW.
62. Fragment of wheat leaf invaded by Uredo Rubigo-
vera, D.C., x 3 136
63. Pustules or soil of Uredo Rubigo-vera, D.C., x 25 . 137
64. Section through a sorus of Uredo Rubigo-vera, D.C.,
x200 137
65. Germinating spore of Uredo Rubigo-vera, D.C.,
xlOOO 133
66. Fragment of wheat stem invaded by Puccinia
Rubigo-vera, D.C., x 5 139
67. Soil of Puccinia Rubiyo-vera, D.C., x 25 . . 139
68. Section through sorus of Puccinia Rubigo-vera, D.C.,
x200 140
69. Teleutospore of Puccinia Rubigo-vera, D.C., ger-
minating in spring, x 1000 . . . . .141
LIST OF ILLUSTRATIONS.
BORAGE BLIGHT.
FIG. PAGE
70. Leaf of Tuberous Comfrey, Symphytum tuberosum,
L., invaded by ^Ecidium asperifolii, Pers., natural
size. ^Ecidium cups, x 10 144
71. ^cidium asperifolii, Pers. Single cup x 50 ; spore,
xlOOO 145
CORN MILDEW— SUMMER RUST AND MILDEW.
72. Fragment of leaf invaded by Uredo linearis, Pers.,
x3 148
73. Pustules or sori of Uredo linearis, Pers., x 25 . 148
74. Transverse section through half a pustule or sorus
of Uredo linearis, Pers., x 200 . . . .149
75. Spores of Uredo linearis, Pers., x 1000 . . .150
76. Two spores of Uredo linearis, Pers., germinating on
a fragment of the epidermis of a wheat leaf,
x400 151
77. Fragment of wheat stem invaded by Puccinia
graminis, Pers., x 5 . . . . .153
78. Pustules or sori of Puccinia graminis, Pers., x 25 . 153
79. Transverse section through half a pustule or sorus
of Puccinia graminis, Pers., x 200 . . .154
80. Teleutospores of Puccinia graminis, Pers., x 1000 . 155
81. Teleutospores of Puccinia graminis, Pers., ger-
minating and producing pro-mycelium spores,
xlOOO 156
BARBERRY BLIGHT.
82. Barberry leaves invaded by sEcidium Berberidis,
Pers. Natural size . . . . . .160
83. Section through a Barberry leaf, showing the cups
of ^Ecidium Berberidis, Pers., below, and the
spermogones above, x 50 . . . . .161
84. Section through a cup of ^Ecidium Berberidis, Pers.,
x!50 163
85. Section through a spermogonium of ^Ecidium Ber-
beridis, Pers., x 300 165
LIST OF ILLUSTRATIONS.
86. Germinating spore of ^aWiwml?0r&m£Ws,Pers., x 1000 166
87. Berries of Mahonia Aquifolium, Lindl., invaded by
d&ddium Berberidis, Pers., x 5 . . . .167
88. Spores of Uromyces appendiculatus, Lev., and
dBcidium Euphorbice-sylvaticce, D.C., germinating 190
89. Development of teleutospores in Puccinia Rubigo-
vera, D.C., x 500 192
90. Teleutospores of Puccinia tragopogonis, Corda, and
P. sparsa, Ck., x 500 207
NEW DISEASES OF WHEAT, BARLEY, AND RYE-GRASS.
91. Upper part of ear of wheat invaded by Fusisporium
cwZmmm, W.Sm. Natural size .. . . .209
92. Fusisporium culmorum, "W.Sm., x 400; spore, x 1000 210
93. Red corns of barley, with. Fusisporium hordei,
W.Sm., x2 210
94. Fusisporium hordei, W.Sm., x 400 ; spore, x 1000 . 211
95. Spike of Lolium perenne, L., invaded by Ergot and
Fusisporium Lolii, W. Sm. Natural size . . 212
96. Fusisporium Lolii, W.Sm., x 400 ; spore, 1000 . 213
ERGOT. — Claviceps purpurea, Tul.
97. Spike of rye, Secale cereale, L.,with Ergots. Natu-
ral size ........ 215
98. Ergots from rye and section, x 2; lodiculesof rye, x 5 216
99. Microscopic structure of Ergot, x 400 . . . 217
100. Ergot germinating and producing Claviceps pur-
purea, Tul. Natural size 219
101. Claviceps purpurea, Tul, x 5, and section through
upper part, x 20 219
102. Conceptacle of Claviceps purpurea, Tul., x 200 . 220
103. Ascus and sporidiurn of Claviceps purpurea, Tul.,
x 500, and 1000 221
104. Half-grown Ergot crowned with its Sphacelia, x 5 . 224
105. Microscopic structure of the Sphacelia, x 400 . 225
106. Sphacelia spore germinating and producing a second
spore similar with itself, x 1000 .... 226
LIST OF ILLUSTRATIONS.
WILSON'S variety of Claviceps on Ergot.
FIG. PAGE
107. Claviceps purpurea, Tul., var. Wilsoni, W.Sm.
Natural size 234
108. Claviceps purpurea, Tul., var. Wilsoni, "W.Sm., x5 234
109. Capitulum of Claviceps purpurea, Tul. , var. Wilsoni,
W.Sm., x20 235
110. Conceptacle of Claviceps purpurea, Tul. , var. Wilsoni,
W.Sm., x200 236
111. ASCIIS and sporidium of Claviceps purpurea, Tul.,
var. Wilsoni, W.Sm., x 500 and 1000 . . 237
MILDEW OF PARSNIPS.
112. Peronospora nivea, Ung., and oospores, x 400 ; con-
idia and oospores, x 1000 ..... 240
113. Protomyces macrosporus, Ung. , x 400 . . . 242
BUNT OF WHEAT.— Tilletia Caries, Tul.
114. Grains of wheat, with Tilletia Caries, Tul., x5 .246
115. Spores of Tilletia Caries, Tul., x 400 . . . 247
116. Germination of Tilletia Caries, Tul., x 1000 . . 248
SMUT OF CORN. — Ustilago carlo, Tul.
117. Panicle of oats, attacked by Smut. Natural size . 255
118. Section through a spikelet of oats slightly affected
with Smut, x 5 256
119. Fragment of glume of oats, showing the epidermis
burst by the Ustilago within, x 25 . . . 257
120. Spores of Ustilago carlo, Tul., x 400 . . . 258
121. Spores in various stages of germination, x 1000 . 259
TARE OR VETCH AND PEA MOULD.
122. Peronospora vicice, Berk, and oospore, x 400 ; coni-
dium, xlOOO 264
PEA MILDEW.
123. Erysiphe Martii, Lk., conceptacle, x 100 . . 266
124. Erysiphe Martii, Lk., ascus with sporidia, x 500 . 267
MILDEW OF LETTUCES.
125. Peronospora ganglioniformis, Berk., x 400 ; coni-
dium, xlOOO 270
xxiv LIST OF ILLUSTRATIONS.
FIG. PAGE
126. Peronospora ganglioniformis, Berk., oospores with
spiral vessel of Lettuce Leaf, x 400 . . . 273
POTATO DISEASE. — Peronospora infestans, Mont.
127. Section through a fragment of potato leaf, with the
fungus of the Potato Disease, Peronospora infestans,
Mont, in situ, x 100 284
128. Fragment of upper part of conidiophore of fungus
of Potato Disease, with conidia and zoospores ger-
minating, x400 287
129. Conidium and zoospore of the fungus of Potato
Disease, x 1000 288
130. The sexual organs, or oogonia and antheridia of
Peronospora infestans, Mont., x 400 . . .297
131. Oogonium and antheridium of Peronospora infestans,
Mont., xlOOO 298
132. Pijthium vexans, D.By., x 400 .... 304
133. Section through a fragment of a decayed potato leaf,
with ripe resting-spores of Peronospora infestans,
Mont, in situ, x 100 305
134. Section through a fragment of diseased potato tuber,
with oospores of Peronospora infestans, Mont. , in
situ, x 400 306
135. Peronospora infestans, Mont., oospores in British
Museum, x 400 307
136. Peronospora infestans, Mont., oospores germinating,
x400 308
137. Artotrogus as illustrated by Professor De Bary in 1881 313
138. Darwin's Potato, Solanum Maglia, Sch. (Frontispiece).
PARASITIC FUNGI AS FOUND IN A FOSSIL STATE.
139. Peronosporites antiquarius, W.Sm., x 400 . . 332
140. Protomycites protogenes, W.Sm., x 400 . . . 333
141. Multiplication by cell-division in Cryptogams . 334
142. Foot of common house-fly, with various fungus
spores engraved to same scale, x 100 . . . 337
143. Green fly from potato, natural size, and wing dusted
with conidia of the potato fungus, x 20 . . 338
DISEASES OF
FIELD AND GARDEN CEOPS.
CHAPTEE I
INTRODUCTORY.
IN preparing the following addresses we have endeavoured
to keep three objects clearly in view. First, the descrip-
tion only of such diseases as are of economic importance.
Second, the definition of all the phenomena of the diseases
in familiar words, such as, with proper attention, may be
understood by all ; this has been done without sacrificing
scientific accuracy, as all botanical terms in common use
are adverted to and explained. Third, the consideration
of the best means of preventing the attacks of plant
diseases.
Many diseases of field and garden crops are too trivial
in their effects to deserve notice : these will be either
entirely passed over or but briefly referred to.
We do not propose to describe any diseases caused by
members of the animal kingdom, with the exception of
one or two caused by the attacks of Nematodes or micro-
scopic worms. The diseases described are chiefly of
vegetable origin, and mostly such as are caused by the
parasitic fungi popularly termed mildews, moulds, smuts,
blights, and rusts. The life history of some of these
parasites is intricate : these will require close attention ;
others are more simple, and these simpler forms of disease
*' 8> B
2 DISEASES OF FIELD & GAKDEN CROPS. [CH.
are taken first, so that the simpler examples gradually
lead up to the more involved ones.
The great question of the prevention, palliation, or cure
of plant diseases is, as a rule, almost entirely overlooked
by botanists ; but from a practical point of view, — and these
notes are specially prepared for practical agriculturists, —
the prevention, palliation, or cure of plant diseases should
surely be an object to be kept chiefly in view.
A knowledge of the structure and vital phenomena of
field and garden plants should be used as a sure stepping-
stone to vegetable pathology. It must be confessed,
however, that in the same way as surgery is often more
precise and certain than medicine, so, much more is at
present known of vegetable physiology and anatomy than
the nature of disease and its prevention. We clearly
know the nature of some diseases of plants ; but as regards
the treatment of plants when invaded by parasites which
are too often the sole cause of disease, we frequently know
nothing. The reason for this defective information is
clear : there are no special teachers of vegetable pathology
in this country, and the few men who have made the
subject more or less a speciality, have not the time or
opportunity for extensive and continued experiment and
research. Field crops under disease are rarely or never
examined by competent observers. As nearly every
known disease of the animal kingdom is susceptible of
preventive, palliative, or curative treatment, it is only
reasonable to assume that the diseases peculiar to the
vegetable kingdom are also susceptible of similar manage-
ment. Of late years the spread of disease in the animal
kingdom has been greatly curtailed, and in the human
family the death-rate of towns has been much reduced.
These results have been entirely brought about by the
acquisition of an exact knowledge of the diseases peculiar
to animals, and of the circumstances favourable to the
spread or extinction of disease. Sanitary improvements
have considerably extended the average length of human
I.] INTRODUCTORY.
life.. It is therefore only reasonable to believe that when
we completely understand the nature of plant diseases, and
the circumstances which aid their spread or tend to their
curtailment, we shall be more or less able to cope with them
by rendering surrounding circumstances unfavourable for
their extension. In the majority of instances it is almost
futile to expect cures : the knowledge to be sought for
must be the facts which will indicate some mode of pre-
vention, or some method of detecting and treating disease
in its earliest stages. In the same way as dwellers in
towns have been of late aroused from their apathy and
made to understand something of what is necessary for
health, so all agriculturists should, if possible, arouse
themselves and learn something of the nature and sur-
roundings of plant diseases. Till this knowledge is
acquired, and till agriculturists become alive to the pos-
sibility of saving their crops from disease, little progress
can be hoped for. We do not say that it is necessary for
every farmer to be a complete master of the anatomy and
physiology of all the plants he grows, or to be perfectly
familiar with the life history of every assailing parasitic
fungus or destructive animal, any more than a house-
holder should know all about the exact nature of typhus,
or diphtheria, or bacteria, bacilli, and disease germs ; but
as every householder at length begins to know, amongst
other facts, that an open drain is likely to prove fatal
to life, so every farmer should know, amongst other things,
that imperfectly-drained fields and rotting vegetable-refuse
mean disease and destruction to his crops.
No sane healthy person would remain in a place tainted
with the contagia of dead and diseased animals, and it is
equally unsafe to place sound plants, tubers, or seeds,
amongst dead or diseased vegetable-refuse. In one case,
as in the other, certain individuals may perchance escape ;
but the general result is, the healthy organisms are at
length destroyed by the dead or diseased ones..
In regard to the illustrations prepared for this work,
4 DISEASES OF FIELD & GARDEN CROPS. [OH.
they are original, and have all been drawn direct from
nature to uniform scales, and engraved by ourselves, so
that the comparative degrees, of largeness or smallness
of the different parts of all the fungi described may be
understood at a glance. Nothing is more wretched than
the copying and recopying of book illustrations, too often
bad ones, without examination or verification. To such
an extent is this copyism at times extended, that in some
instances, only one original drawing has ever been made,
and every succeeding drawing, whether English, French,
or German, is a mere slavish copy. The description, too,
as well as the drawing, of one person is too often taken
on trust alone for an indefinite period of time. The com-
prehension of some published illustrations is sometimes
made difficult by the diverse and odd powers of magni-
fication, such as xlV, x!31, x316, etc., so that after
the examination of a few illustrations a student's mind
becomes greatly confused as to the relative sizes of the
objects illustrated. All magnified figures should be in
tens, hundreds, or thousands. Sometimes certain authors
have illustrated fungi and omitted the amount of mag-
nification upon the plates ; they have simply inserted
"slightly enlarged," "greatly enlarged," "still further
enlarged," etc. ; and the fact is certainly not creditable to
the copyists when we say that every illustration copied
from certain works with which we are acquainted repro-
duces these almost unmeaning terms. The fact shows that
the whole phenomena described by certain writers have
simply been taken by their successors on faith, and that
no single copyist has taken the trouble to measure and
verify for himself.
Students of nature should take very little on trust, for
the sharpest observer is liable to make a mistake in what
he thinks he sees, or in the meaning he attaches to what
he sees, or fancies he sees. Therefore, as far as possible,
every one should observe and think for himself, not with
a view towards finding fault with other observers, but to
i.] INTRODUCTORY.
confirm, extend, modify, and check the observations of
other men. Confirmations are not always to be trusted,
for it often happens that a beginner is over anxious to
confirm the statements of a master, as by that means the
pupil hopes to secure some of the credit belonging to the
original teacher. It is not only necessary to know what
is confirmed, but who confirms it. Some older views of
our own, in which we have now no belief, have been re-
peatedly " confirmed."
All phenomena which on the face of them are unusual
should be carefully examined and re-examined, and con-
stantly tested and retested. Eeasonable statements may
be more readily accepted than unreasonable, but it often
happens that the more wonderful and unreasonable a
phenomenon is — according to the descriptions — the more
avidiously it is accepted, especially by beginners.
Opinions often vary as to the meaning of the pheno-
mena connected with disease, as in the appearances pre-
sented by the fungi of corn mildew, of the potato murrain,
and some other diseases. In these instances we shall not
disrespectfully advance our own views to the disadvantage
of other observers, but shall clearly and impartially state
both sides of any disputed question. We shall, however,
consider it our duty to say how our mind has been im-
pressed by the evidence. Although certain facts are them-
selves often undisputed, yet the deductions made from
them are hotly contr^Vl.
In concluding t* ^^SjjfiJ introductory remarks, we
strongly advise sur ° °^ p^enome^ as nave tne opportunity,
to carefully exar^ Accept notfta hereafter mentioned
for themselves an v°wiH sift and rding on mereaents
Any new obse ve?s ^Mch appear to be'fiifkaBJnable will
of fact or de^ctio-.^ to science.
be doing a &p
CHAPTER II.
CLOVER SICKNESS — CLOVER MILDEW.
Peronospora trifoliorum, D.By.
THE well-known weakly growth of clover, termed clover
sickness, is said — perhaps on insufficient grounds — to be
due to a deficiency of potash in the soil, especially the
soluble salts of potassium in the subsoil. When clover is
grown too frequently in the same fields, and without alter-
nation of crops, the ground becomes " clover sick." Two
nematoid or thread worms (nema, a thread) of minute
size, and allied to the so-called " worms " of stale vinegar
and paste, and to the Nematode which causes ear-cockle
in wheat, oats, and rye, have been described as attacking
clover. These thread worms have been described under
the names of Tylenclius devastatrix and T. Havensteinii ;
but the impoverished condition of clover when due to
these parasites is said to be distinguishable from clover-
sickness proper. Some observers have said that the ail-
ment is due to the presence of a fun- \s known as SpTiceria
herbarum, Pers., sometimes des,^^1 Vs Pleospora, and
more frequently as Clado$§f an(j ' . ">; fungus is over-
1 'ciyiened with synonyneous plaT]f ',1 a^ames, and is so
to be tire* all herbaojf clover sic-rnes< '* ^ hardly likely
named Peziza Uoorioides, Fr., it is a^' ^Vond fungus,
the cause of this ailment, but pro°L^edlV with being
evidence, although, from what we havt 01A ^sufficient
of the attacks of the spawn or mycelium alei% learned
living potato plants, the presence of P. cibo^feKiza uPon
clover deserves attention. A third fun cms !f ' -Fr>> on
Sed Pha-
CH. ii.] CLOVER SICKNESS— CLOVER MILDEW. 7
cidium medicaginis, Desm., has also with, insufficient reason
been referred to as the cause of the disease. It is almost
impossible to say what may be the chief cause of clover
sickness, and there may be several forms of the disease.
The spawn of fungi is sometimes confined to the interior
of plants, where it causes serious disturbance, and this
spawn or mycelium, if without fruit, even when seen
under the microscope, is commonly so indefinite in char-
acter that no one can say for certain what fungus it is
destined to produce.
A frequent fungus on dying clover leaves is Ascdbolus
trifolii, Biv., which is the same with Phacidium trifolii,
Boud.; and another is Polythrincium trifolii, Kze., which
is said by some authors to be a second condition of Do-
thidea trifolii, Fr. A rust fungus named Uromyces append-
iculata, Lev., is also at times very prevalent on the pea-
flower tribe. The fungus parasites of the Leguminosce, to
the pea-flower tribe of which our clovers belong, are but
few in number, although their individual power for de-
struction is great. The best known are the mildews of
our garden and field peas named Erysiphe Martii, Link.,
and E. communis, Schl., the latter of which also occurs
upon the Ranunculacece and the vine. A close ally named
E. graminis, D.C., is parasitic on grasses.
A very frequent parasite of clovers in Britain, and one
to which we are inclined to refer a great deal of clover
sickness, is Peronospora trifoliorum, D.By., a pest which
appears to have attracted little or no attention in this
country till late years.
It, like all other species of Peronospora, attacks living
plants ; it is common on purple clover, Trifolium medium,
L. ; T. alpestre, L. ; crimson clover, T. incarnatum, L. \ -on
Lucern, Medicago sativa, L., and other plants. There is
some diversity of opinion as to the meaning of the name
Peronospora, as Corda, the botanist who first used the jname,
gave no explanation of its derivation. Corda probably
had in view the word peronao (Tre/oovaw — Homer and
8 DISEASES OF FIELD & GARDEN CROPS. [en.
Theocritus) — to pierce, pin, or transfix, or perone, the pin
or tongue of a buckle, a pointed or piercing object, and
spora (<nropa), a seed or spore. He probably used the
word in reference to the power of the fungus to pierce the
tissues of the plant it attacks as distinguished from other
fungi which have no such power. The specific name
trifoliorum explains itself. Peronospora trifoliorum, D.By.,
is closely allied to the potato fungus, and it grows within
and upon the under surface of the leaves of the plants in-
vaded. By this habit of growth, and its putrefactive power,
it not only chokes up the organs of transpiration of the host
plant, but causes decomposition of the tissues by contact.
It is remarkable for the profuse production of its minute,
oval, transparent spores or conidia. A spore in fungi is a
reproductive body, answering to the seed of flowering
plants, but with no embryo or rudimentary plant within.
Certain spores in Peronospora and in many other fungi are
often called conidia from fcom'a, dust, to distinguish them as
secondary spores, or spores of an inferior class, the fungus
itself being capable, under favourable circumstances, of
producing other spores of a much higher order and more
complex structure. The conidia in Peronospora, as the
name indicates, are like fine, generally transparent dust.
These conidia are filled with colourless protoplasm, or vital
material, and they do not readily germinate except in
water. When a conidium of Peronospora trifoliorum,
D.By., falls upon any damp surface it bursts at the side,
and the protoplasm exudes somewhat in the form of an
amoeba, one of the simplest animal organisms. From this
irregular amoeba-like form, other fertile stems of Perono-
spora trifoliorum, D.By., speedily arise.
The disease spots on the leaves, as caused by the Perono-
spora, are at first white, and speedily become pallid or
brownish. At length the corroded fragments drop from
the leaf to the ground. This species of Peronospora pro-
duces oospores, egg -like spores or resting - spores ; these
fall to the ground in the autumn, and rest in a hibernat-
ii.] CLOVER SICKNESS— CLOVER MILDEW. 9
ing state till the following summer, when they germinate,
and produce threads carrying secondary spores or conidia;
these conidia drop off from the parent plant, sail through
the air, and are carried in different directions by currents
of wind. Such spores as light upon clover plants cause
the production of the mildew ; such as fall on unsuitable
places perish. A description of how resting-spores are
produced, and their nature, is given farther on in this
work.
The illustration of this fungus (Fig. 1, A) shows the
parasite enlarged 400 diameters, growing from the under-
surface of the foliage of Medicago saliva L., whilst one of
the very pale gray spores or conidia is enlarged to 1000
diameters at B. In the illustration the fungus is really in-
verted so that its characters may be more easily understood.
All species of Peronospora usually grow from the under
surface of leaves, where they may be seen by the unaided
eye as small white cottony masses. They commonly
burst through the organs of transpiration — s tomata ;
sometimes, however, the fungi push the leaf cells aside
and so get access to the air from the interior of the leaf.
Two stomata are seen in section at C, D. The fertile
threads which carry the spores are termed by botanists
conidiophores or conidia-bearers. The spawn of the fungus
causes putrefaction of the tissues by mere contact. The
spores, on falling on the foliage and bursting, also cause
putrefaction of the leaf. The destruction of invaded
clover is further aided by the conidiophores or steins of the
fungi obliterating the organs of transpiration.
Imperfect drainage and thick planting favours the
growth of all the Peronosporece, whereas a free circulation
of dry air is often fatal to them. When once they make
their attack, it must be remembered that they establish
themselves within the tissues of the invaded plants, in a
position where it is impossible to reach them with any
curative material. Attention, therefore, to the mode of
cultivation may tend to stop the spread, if not to prevent
10
DISEASES OF FIELD & GARDEN CROPS. [OH.
400-
FIG. 1. — CLOVEK MILDEW.
Peronospora trifoliorum, D.By.
Enlarged 400 diameters. Spore enlarged 1000 diameters.
the original attack of the foe. As the fungus hibernates
in decaying clover plants in the winter, it is obvious that
ii.] CLOVER SICKNESS— CLOVER MILDEW. 11
the best mode of preventing attacks is, where possible, to
destroy all clover refuse with fire, and not allow any to
rot on the surface of the ground. This treatment will, of
course, not only destroy the eggs of any nematoid worms,
but the spawn of the other fungi so frequently seen on
sickly clover plants in the summer and autumn.
The Nematodes appear to be attracted to the decayed
spots caused by the Peronospora, for in these positions, in
company with resting spores of the parasite, which un-
doubtedly causes one form of putrescence, we have com-
monly found them.
CHAPTER III
PERONOSPORA EXIGUA, W.SM.
ANOTHER Peronospora, with the habit of P. grisea, Ung.,
a parasite of Veronica, is very frequent upon clovers in
Britain, is equally destructive with P. trifoliorum,
FIG. 2.— CLOVER MILDEW.
Peronospora emgua, W.Sm.
Enlarged 400 diameters. Spore enlarged 1000 diameters.
D.By., and is a minute and hitherto undescribed species.
This plant, Peronospora exigua, W.Sm. (exiguus, small), is
illustrated, enlarged 400 diameters at A, Fig. 2 • and a
CH. m.] PERONOSPOKA EXIGUA. 13
single spore, or conidium, is enlarged 1000 diameters at
B. The parasite causes putrescence, and grows within
the leaf and in effused patches on the surface, as does
P. trifoliorum, D.By. The illustration shows a some-
what large group of fruiting fungus threads, growing
upon a fragment of a leaf hair of Lotus corniculatus, L. ;
the mycelium has caused a gouty discoloured swelling to
appear on the hair. The spore - supporting threads are
frequently simple, or sometimes once, twice, or three times
branched and furnished with a few joints, stops, or septa ;
the spores are borne on minute, often lateral spicules ;
and as the branches grow, the spores have a tendency to
drop off, as in the Peronospora of the potato disease. The
spores are slightly oval, almost round, and burst at the
side on germination. P. exigua, W.Sm., sometimes grows
on clovers in company with P. trifoliorum} D.By., but
it is immediately distinguished by its very much smaller
size, as well as by its specific characters. Though com-
mon, this species has hitherto been overlooked, probably
because its appearance on clover leaves to the unaided eye
is precisely the same with that of P. trifoliorum, D.By.
PERONOSPORA EXIGUA, W.Sm. — Minute, conidiophores
simple or slightly branched, slender, sparingly septate,
conidia oval, almost globose, very small, very pale gray,
sometimes borne on one side of the conidiophore only,
non-papillate, bursting at the side on germination. On
Leguminosce, often in company with P. trifoliorum,
D.By.
The " New Clover Disease," described by P. Mouille-
fert in the Journal d' Agriculture Pratique, 1874, pp. 667,
670, and translated by Mr. William Carruthers, F.K.S.,
in the Journal of the Royal Agricultural Society of England,
Vol. x., Part ii., 1874, is possibly one form of clover
mildew, caused either by Peronospora trifoliorum, D.By.,
or P. exigua, W.Sm. In the disease described by P.
Mouillefert the fungus appears to be chiefly confined to
the base of the stems of clover. The author says he has
14 DISEASES OF FIELD & GARDEN" CROPS, [on. HI.
not been able to detect the fungus in the leaf, but,
curiously enough, he gives an illustration of it in that
position ; he has not illustrated the spores, perhaps
because they so readily fall from their little pedicels in
Peronospora, and are easily overlooked. As this fungus
doubtlessly hibernates in the same manner as the last, the
only plan for lessening its ravages is to burn all decaying
clover material.
This parasite is very near Ovularia (Ramularia) sphce-
•roidea, Sacc. ; but if the published descriptions are correct,
our plant differs materially from the German examples.
CHAPTER IV.
NEW DISEASE OF POTATOES.
Peziza postuma, Berk, and Wils.
IN the beginning of the month of August 1880, Mr.
Ambrose Balfe, secretary to the Boyal Horticultural
Society of Ireland, reported to us a disease then invading
certain crops of potatoes in the west of Ireland, in a
manner hitherto unknown to him. The potatoes had
been bought as " Champions," and planted in land which
had been reclaimed from bog eight years previous to the
outbreak of the disease. When the ground was reclaimed
a coat of clay was spread over and incorporated with the
soil. For the first three years potatoes were grown, fol-
lowed by a year of oats, next the ground was sown with
grass and meadowed, and lastly "champion" potatoes
were planted. In preparing the ground for the potatoes
sea-weed was first spread over the grass, and ten days
afterwards it was covered with farmyard manure. The
potato sets were laid on the manure, and then covered.
Ridge planting was adopted. No doubt the mode of
culture was defective, as it is bad in practice to place
potato sets in immediate contact with decaying vegetable
matter and farmyard manure ; such materials always con-
tain an immense number of disease germs both of animal
and vegetable origin. The manure used for potatoes
should always be old and thoroughly decayed, and it is
perhaps best that the cut faces of the sets should be
allowed to dry before they are planted. Some planters
pass the cut surfaces rapidly across a hot iron with good
effect, but others maintain that it is better to place the
16 DISEASES OF FIELD & GARDEN CROPS. [CH.
freshly-cut sets in the soil immediately after cutting, and
whilst the wound is still quite fresh. Planting in rank
undecayed material is not only destructive to the material
which is stored up within the tuber or set (material
which is the food of the future plant), but it is also in-
jurious to the young shoots and rootlets, for any hot,
fermenting material acts as a poison to these growths, and
diminishes the vigour of the infant plant. The conditions
of planting in the instance here adverted to may, how-
ever, have had nothing to do with the disease which
followed in the summer.
Until the attack now under description, the potato
plants, as far as outward appearances went, were free from
any taint of the fungus of the potato disease proper,
named Peronospora infestans, Mont. The disease was first
noticed in the beginning of July, at the time the potato
flowers were opening ; but there can be little doubt that
it was in or upon the plants several weeks previously, as
by its nature it would not attract much attention at first.
It is strange that other potatoes named " Protestants,"
growing close to the " Champions," were not attacked.
The appearance of the diseased plants was peculiar ; they
were covered within and without with a thick felt of
white fungus spawn or mycelium. The growth of this
spawn was so rapid and profuse that in a week or two the
whole of the stems and leaves were reduced to tinder, the
entire moisture belonging to the stems and leaves being
exhausted by the fungus. Leaves are of such vital im-
portance to plants that the destruction of them is
synonymous with a cessation of the plant's growth. If
the parts of a potato plant which are above ground get
seriously injured or destroyed, there will be little or no
further growth in the tubers. In the Peziza disease, now
under description, the mycelium was not a putrefactive
one as in Peronospora. It merely caused a sudden cessa-
tion of growth in the tubers.
Immersed in the thick felt of white fungus spawn,
NEW DISEASE OF POTATOES.
17
FIG. 3. — NEW DISEASE OF POTATOES.
Peziza postuma, B. and Wils. Potato stems with Sclerotia.
Natural size.
when it had reached its maximum of growth on the
potato stems, there were thousands of small black nodular
C
18 DISEASES OF FIELD & GARDEN CROPS. [CH.
bodies, varying in size from a grain of sand to that of a
small bean. An affected potato stein is illustrated,
natural size, at A, Fig. 3, and a section through a part of
a similar stem is illustrated at B to show the black
nodular growths in situ. The small black bodies here
drawn are at first white, at length they become externally
brown, and ultimately black ; they are hard and compact,
and, owing to their hardness, they have been termed
Sclerotia from sJderos, hard. One of these bodies sur-
rounded by spawn threads is shown, twice its natural
size, at Fig. 4. These nodular growths when examined
-X-2-
FIG. 4.
Sclerotium of Peziza postuma, B. and Wils.
Twice the natural size.
with the microscope are found to consist of highly con-
densed and compacted spawn cells or mycelium, white
in the centre and gradually getting black (through brown)
towards the outside. When an excessively thin slice is
taken off a cut surface of one of these nodules and magni-
fied 400 diameters, the appearance is similar with the
illustration at Fig. 5 ; here the gradual change of colour
from the white internal cells to the black thick -walled
outer ones is illustrated, together with the felted statum
of white mycelial threads, on the top of illustration, in
which the Sclerotia are embedded.
Sclerotia, or compacted masses of fungus spawn in a
resting state, are common amongst fungi ; some examples
iv.] NEW DISEASE OF POTATOES. 19
never attain a larger size than a grain of gunpowder,
others -are as large as peas or small beans. The " Native
Bread" or Mylitta of Australia, which often measures
several inches across, may be a Sclerotium. The edible
Americo-Indian Tuckahoo, which is dug out of the ground
in large masses, is not really a fungus, although so esteemed
•X'4-OO-
FIG. 5.
Section through outer surface of Sclerotium of Pesiza Postuma, B. and W.
Enlarged 400 diameters.
by Fries, and named by him Pachyma cocos. Some Sclerotia
are sphaerical in shape, whilst others are elongated irregular
ovals. Sclerotia are not confined to one order or genus of
fungi, but they possibly occur throughout the entire family.
Some forms are much less compact than others, and the
looser forms germinate after a comparatively short rest.
By means of Sclerotia certain fungi which would prob-
ably perish during drought or severe frost are preserved
alive through inclement seasons. The spawn naturally
compacts itself into these little hard masses and falls to
the ground ; it there remains, like a seed, uninjured by
continued cold or dryness, whereas the vitality of uncom-
20 DISEASES OF FIELD & GARDEN CROPS. [CH.
pacted spawn would be destroyed under similar conditions
in a few hours. Sclerotia vary in their period of hiberna-
tion from a few days to a year or more ; they are some-
what erratic as to the required amount of rest ; favour-
able circumstances will hasten on germination, whilst
unfavourable ones will retard it ; a very common period
of hibernation is from nine months to a year. When
germination at length takes place a perfect fungus is pro-
duced ; this perfect fungus at maturity produces spores
which, on germination, again produce spawn or mycelium.
Sometimes this spawn will at once reproduce the perfect
fungus, but in other instances it grows profusely, and at
length gives direct rise to the little resting nodosities just
described as Sclerotia.
Some Sclerotia on germinating only produce moulds or
mildews, whereas others may produce a tall club-shaped
fungus termed Typhula, from Typha, the reed-mace ; some
give rise to mushroom -like fungi, true Agarics, whilst
others produce cup-shaped fungi which may be either
sessile or supported on a long stalk ; these latter fungi
are termed Pezizce.
Many Sclerotia have received specific names, but such
names are almost valueless. For instance, two so-called
different species of Sclerotia have been known to give rise
to the same perfect fungus, — Typhula phacorrhiza, Fr.
(phacos, a lentil, and rhiza, a root), grows either from
Sclerotium complanatum, Tode, or S. scutellatum, A. and S.
Both Sclerotia are found on dead leaves, and although
they have been described as distinct they must be the
same with each other. Polyactis cinerea, B., and Peziza
Fuckeliana, D.By., both spring either from Sclerotium
durum, P., or S. echinatum. For this reason some botan-
ists esteem the Polyactis to be an early state of the Peziza.
Whilst respecting the opinions of the botanists who have
advocated this startling view, we are inclined to sus-
pend judgment and wait for confirmation. Polyactis
cinerea, B., not only grows on Sclerotia, but is extremely
iv.] NEW DISEASE OF POTATOES. 21
common on all decaying vegetable matter when no
Sclerotia are present. No one can tell for certain, by
mere examination, what any given Sclerotium will produce
on germination. Sometimes a clue is given to what a
Sclerotium may possibly produce by observing its habitat.
For instance there is one Sclerotium named S. fungo?°um,
P., commonly found in dead examples of certain members
of the mushroom tribe, chiefly found under the gen era named
Agaricus and Eussula, and sometimes inside the large decay-
ing corky fungi, named Polyporus, of trees ; these Sclerotia
invariably produce a small mushroom-like fungus named
Agaricus tuberosus, Bull. A closely allied Agaric, named
A. cirrhatus, Sch., also springs from a not dissimilar
Sclerotium. Another and very small Sclerotium found in
decaying onions and named S. cepcevorum, B., produces a
minute mould named Mucor subtilissimus, B. It com-
monly happens, however, that Sclerotia may be found on
the surface of the ground near where the supporting plant
has decayed. When gathered from such positions it is
impossible to say what they will produce under culture,
and sometimes they so closely resemble small truffles
(which, too, are often found on the surface of the ground),
that without a microscopical examination it is impossible
to distinguish one from the other. The perfect state of
some Sclerotia is unknown, as S. stipitatum, Fr., found in
the nests of white ants in India.
When the large black Sclerotia were found in Irish
potatoes they appeared to us to differ from the Sclerotia
we had hitherto noticed, and although it is never safe to
guess at what an unfamiliar Sclerotium may produce, yet
in our printed report, published in the Gardeners' Chronicle
for 20th August 1880, we hesitated to refer the bodies to
any already described form of Sclerotium. Sclerotia are by
no means uncommon in herbaceous stems, in cabbage
stumps, and even in potato stalks ; but the new bodies did
not appear to us to be the same with any others we had
previously observed. Black Sclerotia are not uncommon
22 DISEASES OF FIELD & GARDEN CROPS. [CH.
in dead potato stems in the spring, but not on living
potato plants in the summer.
Great efforts were made both by ourselves and numer-
ous friends to make the new potato Sclerotia germinate,
but in every instance that year without result. Many
enquiries were made of potato growers, but the Sclerotia
appeared to be lost again to this country, and nothing more
was heard of them till a paragraph appeared in Nature
for 19th July 1883. This described, from Natureen, a so-
called hitherto unknown form of the potato disease, which
had been making slow but steady progress near Stavanger
during the previous ten or twelve years, and was then, so
said the report, showing increased energy. The potato
stalks were reported to be the first parts affected, and
here Herr Anda had discovered small white fungoid
growths, which, after attaining the size of a small bean,
finally assumed a black colour. It appears that the
fungus at Stavanger, like the same growth on the west of
Ireland, rapidly increased at the expense of the supporting
plant, first reducing the interior of the potato stem to a
pulpy condition, and then shrivelling and hollowing it
out until nothing was left but the mere outer shell.
About the end of July or the beginning of August the
destruction caused by the fungus is seen at its worst,
and at this period whole fields of potato plants are often
reduced to the condition of withered straw.
Now this disease at Stavanger, which was more or less
lost sight of in Britain for three years, was clearly identi-
cal with the one reported on by us ; it agreed in every
way, and we at once wrote a note to Nature expressing
our regret that although the germinating Sclerotia had
apparently been seen by Herr Anda, yet the perfect plant
had not been identified ; so that, so far as Herr Anda's
report went, we were as much in the dark as before as
to the fungus which had caused the disease. A few days
after our letter was published, our friend Mr. A. Stephen
Wilson, of North Kinmundy, Summerhill, by Aberdeen
iv.] NEW DISEASE OF POTATOES. 23
— who, it may be said, had not seen our note in Nature —
sent us a box enclosing numerous living examples of the
desired black Sclerotia from potatoes grown near Aberdeen.
Mr. Wilson informed us in an accompanying letter that
he had hundreds of similar examples from the previous
year's potato stalks then germinating, and that, judging
from the appearance they presented at that time, they
would probably produce a Peziza allied to Peziza tuberosa,
Bull. At the same time a third letter appeared in Nature,
this time from Professor Blytt, stating that he had for-
warded examples of the Sclerotia to Professor De Bary,
and that they had germinated with him. Professor Blytt
stated that the Sclerotiutn had produced a fungus identified
by Professor De Bary as Peziza sclerotiorum, Lib., a species
which had not at that time been recorded as British. At
the period when this discussion was going on we received
other living examples of Sclerotia from Mr. Thomas
Carroll of the Gilbert Institution, Glasnevin, Dublin.
Mr. Carroll stated that the disease was prevalent on the
west coast of Ireland, especially on land too heavily
manured. On making further enquiries we learned that
the disease was spread generally over the counties of
Sligo, Mayo, and Galloway, and that it was by no means
confined to any special variety of potato ; that the
manuring with seaweed had nothing to do with the
disease, but that large patches of potato plants twelve to
fifteen yards in diameter were destroyed by the spawn of
the Sclerotium, and the most luxuriant crops were appar-
ently the most affected. The apparent luxuriance of the
affected crops was perhaps not real, for it is well known
that when some plants are first attacked by fungoid
diseases they put on a spurious appearance of luxuriance,
and are exqited into a sudden quick growth. It has often
been remarked that the apparently most healthy potato
plants fall first before the disease caused by Peronospora
infestans, Mont. The reason is found in the fact of the
spawn of the Peronospora being capable of exciting a sudden
24 DISEASES OF FIELD & GARDEN CROPS. [CH.
abnormal growth of chlorophyll or leaf -green, and in
other ways causing a morbid enlargement of the tissues,
termed hypertrophy. A transitory and spurious appear-
ance of unusual health and vigour is not peculiar to the
attacks of fungi, for insect injuries often excite the same
appearances. It is well known that the largest and- best-
looking pears in orchards in the early summer are often
swarming with larvse. The larvae stimulate the growth
of the young pears ; probably this abnormal growth is
an effort of nature to repair an injury. The larger
fruits fall to the ground and decay about the end of
May.
Mr. A. Stephen Wilson succeeded in raising such a
large crop of Pezizce from his Sclerotia that he supplied
us, as well as the Rev. M. J. Berkeley, and several other
botanists, with living examples. They came up in large
numbers in a garden bed where the Sclerotia had been
thrown the previous year. The Pezizce were all attached
to the Sclerotia, and Mr. Wilson even detected the Pezizce
growing from potato stems with the Sclerotia in situ.
Mr. Wilson considered the Sclerotium to be S. varium, P.,
a well-known pest of carrots, parsnips, cabbages, Jerusalem
artichokes, etc. Mr. Berkeley came to the conclusion that
the Peziza was a new species, to which he gave the name
P. postuma, Berk, and Wils. in the Gardeners' Chronicle for
1 5th Sept 1883. Whatever the name of the Peziza should
be, there is one thing quite certain, and that is, the plant
is new to the scientific observers of this country. The
genus Peziza derives its name from Pezica, a word used
by Pliny to denote a fungus ; and Messrs. Berkeley and
Wilson termed the plant now under description postuma.
because the perfect plant is produced at a much later
period than the Sclerotium.
There are at least thirteen or fourteen of these long-
stemmed Pezizce described, and the present plant comes so
near Peziza ciborioides, Fr., P. sclerotiacea, Ces., and P.
sclerotiorum, Lib., that it is almost impossible to distin-
iv.j NEW DISEASE OF POTATOES. 25
guish it from them. The rest of the described species
have smaller sporidia.
A few of the germinated examples forwarded by Mr.
Wilson are illustrated, natural size, at Fig. 6. It will be
FIG. 6.
Peziza postuma, B. and W.
Natural size.
seen that the Sclerotia give rise to long, slender, tortuous
stems, and that each stem at length bears a shallow pallid
head, which ultimately becomes flat or slightly recurved.
The stems are about two inches high, and the cup-like
26 DISEASES OF FIELD & GARDEN CROPS. [CH.
expansion at the top about lialf-an-incli across. On closely
observing these expanded tops in sunlight, especially when
held in favourable positions, as against a black background,
a slight sudden cloud resembling a puff of smoke or steam
may be seen to gradually sail away through the air from
the top surface. This almost invisible cloud really con-
sists of millions of minute spores, in this instance techni-
cally termed sporidia for a reason to be mentioned imme-
diately.
The whole interest of the fungus now centres on the
expanded top, and especially to its surface, whence the
clouds of sporidia sail away. If we cut one of the cups
in two, and look on the cut surface with a magnifying
power of five, and twenty diameters, we shall see the
structure as shown at A and B, Fig. 7. We now notice
FIG. 7,
Peziza postuma, B. and W. Section through cup.
Enlarged 5 and 20 diameters.
that the whole of the upper stratum of the expanded top
consists of elongated perpendicular cells as illustrated,
whilst the under surface is a mass of spherical cells of
various sizes ; but to see this curious structure well a
much higher magnifying power is required, and a small
fragment only of the top must be examined in section, as
at Fig. 8, magnified 400 diameters. We now distinctly
see the basal stratum of transparent globular cells of
various sizes, and a few of the hundreds of thousands of
vertical transparent bladders forming the top stratum,
and from which the cloud of dust consisting of oval trans-
parent sporidia arises. With a lancet we will now remove
a few of these vertical asci or long bladders, and magnify
NEW DISEASE OF POTATOES.
27
500 diameters. Each colourless bladder, sack, or bottle,
is termed an ascus, from askos, a bag, bladder, or bottle.
Each ascus, as at Fig. 9, contains eight oval spores or
lia. These bodies, one of which is farther enlarged
•x-400-
FIG. 8.
Peziza postuma, B. and "W.
Section through fragment of cup.
Enlarged 400 diameters.
X-500
FIG. 9.
Peziza postuma, B. ai:>l \v".
Asci, paraphysis, and sporidia.
Enlarged 500 diameters.
to 1000 diameters at B, are termed sporidia because they
are not supported on threads or fine branches like many
other spores, but are carried free in an ascus or transparent
bladder, in which position they have arisen from a dif-
28 DISEASES OF FIELD & GARDEN CROPS. [CH.
ferentiation of the protoplasmic contents of the ascus.
Mixed with the asci are numerous slender, often septate
organisms, termed paraphyses (as at A, Fig. 9), or organs
which grow about or in company with organs of greater
importance. The word is derived from para, about, and
phud, I grow. The right hand ascus in the illustration is
shown in the act of discharging its sporidia into the air.
At a given moment, depending upon unknown conditions,
possibly of the air, of light or heat, the ascus opens at the
top, as illustrated (in some fungi an operculum or lid flies
off), and discharges the eight sporidia which it invariably
contains into the air. In the genus of fungi named
Ascobolus, the ascus itself, with its contained spores, as
the name indicates, is shot into the air. Each transparent
sporidium is furnished with two or three lustrous spots.
The asci are so inconceivably small, slender, and attenu-
ated that there are more than 300,000 packed side by
side on the top of each expanded cap, which on the
average measures about half-an-inch in diameter ; and as
each ascus contains eight sporidia there are no less than
2,500,000 sporidia produced by every cap. Now, as
every infected potato plant will produce at least fifty
Sclerotia, it follows that a plant killed by this new disease
is capable, by means of its germinating Sclerotia during
the following season, of discharging more than 100,000,000
reproductive bodies into the air. It must now be specially
noted that after a year's rest the Sclerotia germinate on
the ground, and there produce their sporidia exactly at
the time in July when potatoes are making their best
growth. A vast number of the sporidia must perish, but
such as fall upon potato plants (and possibly some other
plants, as carrots) germinate at once, cover the stems with
spawn, obliterate the organs of transpiration, and speedily
reduce the haulm either to a mass of putrescence or to
dry tinder. During this rapid and exhaustive growth the
spawn again gradually compacts itself into the black
nodules of condensed mycelium termed Sclerotia, and
iv.] NEW DISEASE OF POTATOES. 29
these nodules are destined to germinate and produce the
Peziza disease the following year.
The remedy for this state of disease is obvious. No
infected stems should be allowed to rot in the fields, but
all should be carefully gathered together and burnt. If
the stems are allowed to rot on the ground the disease is
almost certain to recur ; but if they are burnt not only
will the Sclerotia of the Peziza be destroyed, but the
spawn, germs, eggs, and spores belonging to numerous
other parasites, perhaps equally bad with the Peziza
itself, will be destroyed at the same time. If, on an
examination of the potato stems, it is found that many of
the Sclerotia have dropped from them, the top surface of
the ground should if possible be raked and burnt.
Nothing is more common than to find hibernating repro-
ductive bodies falling readily to the ground. This is
clearly a natural provision for their preservation.
OHAPTEE V.
FUSISPORIUM DISEASE OF POTATOES.
Fusisporium Solani, Mart.
WHEN potatoes are destroyed by parasitic fungi in the
autumn, it does not always happen that the parasite is
the dreaded putrefactive fungus of the murrain, named
Peronospora infestans, Mont. Another highly destructive
fungus, named Fusisporium Solani, Mart., is sometimes
equally damaging to potatoes with the Peronospora itself.
The name Fusisporium refers to the spindle-shaped crescent
form of the spores, — fusus, a spindle ; Solani, of course,
refers to the genus Solanum, to which the potato belongs.
It often happens that Fusisporium grows in company
with Peronospora on potatoes, at other times the two
fungi grow apart ; exactly the same phenomenon of con-
sortism occurs with Peronospora Schleideniana, Ung., and
Fusisporium atro-virens, B., on onions. The same con-
sortism also occurs with Peronospora parasitica, Pers., and
Fusisporium aurantiacum, Lk., on the cabbage tribe.
Fusisporium Solani, Mart., although very common in
the southern and midland counties of England, has not
been recorded from Wales or Scotland. Two allied species
attacking the parsnip and turnip have been noticed in
the latter places ; and it appears strange that the Fusis-
porium of the potato should have been overlooked if it
really occurs. But however rare the fungus may be in
the north and north-west, it is certainly an extremely
common and highly destructive pest of potatoes over the
greater part of England.
Like many other fungi, Fusisporium Solani, Mart.,
CH.V.] FUSISPORIUM DISEASE OF POTATOES. 31
occurs in more than one form. One condition of the pest,
named Periola tomentosa, Fr., was described in 1836 as
an assailant of potatoes in the midland counties by the
Rev. M. J. Berkeley. This form is probably no other
than masses of compacted villons mycelium or spawn,
from which the Fusisporium at length arises. Another
and early condition of Fusisporium Solani, Mart., was
also described by Mr. Berkeley under the name of Dacty-
lium tenuissimum. A. distinct rose-coloured species of
Fusisporium, named F. roseolum, Steph., also grows on
decayed potatoes.
Fusisporium Solani, Mart., is not peculiar to decaying
potatoes, for it grows with rapidity on potatoes which are
apparently undiseased, if bruised or cut. To the unaided
eye its growth bears such a close general resemblance to
Peronospora that it might readily be mistaken for it ;
indeed, the use of a lens, even with practised observers,
is often necessary to distinguish between the two. The
fungus is simple in structure, and consists of threads and
compound spores ; the spore- bearing threads springing
direct from a floccose, sometimes mucilaginous, bed of
spawn or mycelium, as illustrated at A, Fig. 10, enlarged
400 diameters. The fruiting threads carry spores attached
somewhat obliquely, as illustrated at B, but more clearly
seen in .Fig. 11, where they are enlarged to 1000 dia-
meters. The spores are spindle-shaped — that is, attenuated
at both ends ; and curved like a crescent : each spore, at
first simple, is at length furnished with about three joints
or septa, and each of the four pieces of the compound
spore exhibits one or more lustrous spots. Sometimes
all four parts of a spore germinate whilst still attached to
the supporting threads, as at C, Figs. 1 0 and 1 1 ; but it
often happens that at maturity the spores fall into four
pieces, as at D, Fig. 11, and each of the fallen pieces
sometimes germinates at once. This rapid germination,
by its extension of the parasite, accelerates the destruction
of the host plant. The fallen pieces of the spore are at
32 DISEASES OF FIELD & GARDEN CROPS. [OH,
FIG. 10. — DISEASE OF POTATOES.
Fusisporium Solani, Mart. Enlarged 400 diameters.
first squarish, but on alighting on any moist surface they
speedily take a spherical form. Sometimes these little
bodies do not germinate at once, but hibernate for a short
FUSISPORIUM DISEASE OF POTATOES.
time, generally varying from three weeks to three months
(two months is a common time), and during this period
they become slightly spinulose, and faintly tinted with a
brownish hue. These little bodies, therefore, hibernate
after the manner of resting-spores, and it is possible that
many of them rest during the entire winter. A spore
which has hibernated is shown at E, Tig. 11. The ovate
FIG. 11.
Fusisporium Solani, Mart. Spores in different stages of growth.
Enlarged 1000 diameters.
bodies at F, Fig. 10, are grains of starch within a cell of
.the potato tuber.
The destruction of potatoes is complete when the Fusis-
porium works in company with the Peronospora, and
when the spawn-threads of both fungi are interlaced, a
condition very often observed in the midland and southern
counties.
In experimenting on the growth of this fungus, it is
very easy to transfer the flocculent or semi-mucilaginous
D
34 DISEASES OF FIELD & GARDEN CROPS. [OH. v.
spawn from a diseased potato to a thin slice cut from an
apparently sound one. If a very thin transparent slice
of a sound potato is placed on a glass slide, and some of
the spores and threads of the Fusisporium placed near the
edge of the apparently healthy living slice, as illustrated
in Fig. 10, the rapid progress of growth in the fungus
can be observed with ease if the material under experi-
ment is kept moist. The growth of the Fusisporium is
extremely rapid, and the production of the compound
spores most profuse. The small hibernating spores burst
and produce a perfect Fusisporium in six Jiours. The
mycelium appears to have the property of breaking up
the cell walls, of injuring the contained starch, and of
speedily reducing the potato to a loathsome mass of put-
ridity. In certain instances the presence of the Fusispo-
rium appears to cause the substance of the invaded potato
to rot and become dry. The fungi found under Fusisporium
are not generally considered to be capable of producing
putrescence of tissues, but F. Solani, Mart., is an exceptional
species ; the mycelial threads and the supporting threads
of the spores differ from typical species of the genus. A
profuse growth of Fusisporium Solani, Mart., when seen
under the microscope, looks like the surface of a field of
corn, the ears being represented by the closely -packed
Fusisporium spores. For lessening the attacks of this
fungus, the only known plan is to destroy all affected
material with fire, and not allow any decayed potato-
refuse to remain in the fields.
CHAPTER VI.
SMUT OP POTATOES.
Tubercinia scabies, B.
THE smut which produces one form of scab in potatoes
is caused by an olive-green or brownish fungus, of sub-
terranean habit, named Tubercinia scabies, B. ; from tuber,
an ancient Roman name for a fungus ; cineres, ashes ; and
scabies, the itch. This is the same with Bhizosporium
Solani, Rab., and is the Protomyces of Martius. The smut
fungus, which is very common upon potatoes, is supposed
to be allied to the bunt of wheat, Tilletia caries, Tul. ; and
the smut of corn, Ustilago carlo, Tul. It was described
and illustrated by Martius, Die Kartoffel-Epidemie, p. 23,
tab. 2, Figs. 9-13, tab. 3, Figs. 36-38, and afterwards by
the Rev. M. J. Berkeley in the Journal of the Royal
Horticultural Society, vol. i., 1846, page 33, Figs. 30 and
31 . The spores are compound, and are composed of minute
cells, which together form a hollow globe with one or more
apertures. In this character Tubercinia agrees well with
the allied genus Urocystis (formerly Polycystis, in reference
to the spores being composed of many cells), to which the
well-known smuts of rye and violets belong. The fungus
grows beneath the bark of the tuber, where it forms a
thin dark greenish-brown stratum, often extending over
the greater part of the external surface of the potato. The
presence of the fungus may be detected by discoloured
blotches on the bark. As in the disease caused by the
Peronospora, it often happens that no trace of the fungus
is to be seen at the time of harvesting. It frequently
shows itself during the winter in stored potatoes, which,
36 DISEASES OF FIELD & GARDEN CROPS. [CH.VI.
at the time of digging, were apparently quite sound. In
bad cases small discoloured spots first appear on the bark
of the tuber ; these increase in size and become confluent,
till at length the entire skin of the potato is discoloured.
The cuticle then bursts in many places, and the olive-green
spores are liberated. Like many other plant diseases,
potato smut is in some seasons rare, at other times it is
very common and destructive. No potatoes showing
traces of smut should be planted as seed.
CHAPTEE VII.
SCAB AND CRACKING OF POTATOES.
THE brownish scab on the skin or bark, and the cracking
of the bark in potatoes, are probably due to one and the
same cause, and that a mechanical one. Scab and crack-
ing are often confounded with smut in potatoes, but
whereas the latter ailment is invariably caused by a
fungus, there is seldom any fungus present (unless it be
one of the species which so commonly follow injuries) in
scab and cracking. Cracked and scabbed potatoes are
not marketable, and as the fissures in the bark expose the
inner substance of the potato to the earth it is generally
said that tubers so injured possess an earthy and dis-
agreeable taste foreign to uncracked and unscabbed ex-
amples.
Scabbing and cracking begin at a very early stage of
growth in the tuber, and both are at first seen as small
corroded spots, or minute open pustules ; in bad cases the
spots and cracks become confluent, and the whole bark
of the potato presents an unsightly appearance. When
the inner substance of the potato is once exposed minute
insects, and sometimes fungi, add to the injury. If such
fungi as Peronospora or Fusisporium light on the exposed
places, destruction of the tuber is soon complete.
Scab and cracking are said, in the first instance, to be
due to the presence of some irritating or corrosive sub-
stance in the soil. Continued drought, and possibly
sudden and superabundant moisture, will also cause one
form of scab. A natural effort is made by the potato to
repair the injury, and so a hard scab originates ; when
38 DISEASES OF FIELD & GARDEN CROPS. [CH.VII.
insects and fungi light on the injured parts, repeated
efforts are made to repair the damage, and so the bark is
brought into a scabbed and cracked condition. Lime
rubbish, builders' refuse, refuse from ashpits, and other
materials of the same class, are said to cause corrosion,
scab, and cracking of the bark of potato tubers by contact.
When the bark is carefully removed small depressions
are clearly seen, answering to the scabs removed with the
bark. In bad cases the pits beneath the scabs are exca-
vated deeply into the substance of the potato, and when
the bark is removed the substance of the tuber, though
frequently slightly discoloured, is left intact.
It generally happens that a portion only of a crop of
potatoes is scabbed, and this portion can be often dis-
tinctly traced to one part of the field whence the potatoes
were derived. On visiting this position the irritating
substances in the soil will usually be seen. When scab
and cracking can be thus traced the remedy is obvious.
CHAPTEE VIII.
NEW DISEASE OF ONIONS.
Puccinia mixta, Fl.
THERE are few kitchen-garden crops more liable to dis-
ease than onions, and in the best managed fields and
kitchen gardens, and in dry as well as wet seasons, whole
crops of onions, and all varieties alike, are liable to be
swept off by the attacks of fungi.
During the summer of 1883 great attention was directed
to a fungus named Puccinia mixta, FL, found growing
on chives, near Shrewsbury. The name of the genus Puc-
cinia was given in honour of Puccini, a Florentine professor.
When we remember how completely Puccinia malvacearum,
Mont., has, during the last few years, destroyed all our
best garden hollyhocks, we may well feel some anxiety as
to the course this new pest of onions may pursue. Mr.
William Phillips, F.L.S., of Shrewsbury, was the first to
detect the onion parasite, named Puccinia mixta, FL, grow-
ing in a garden. Mr. Phillips recorded its occurrence in
the Gardeners' Chronicle for 14th July 1883, and there
stated that the parasite was growing on chives, Allium
Schcenoprasum, L., and the crop, he said, was in a de-
plorable condition of disease, the leaves and scapes, or
naked flower-stems, being covered with yellow and brown
spots, and presenting a miserable appearance. Mr. Phillips
was good enough to forward some specimens to us at the
time of finding, and from these examples the illustrations
have been made.
Chives are perennial and indigenous to Britain. They
are grown to no great extent in England, but in Scotland
40 DISEASES OF FIELD & GARDEN CROPS. [OH.
they appear in nearly every kitchen-garden, where they
are grown for flavouring soups, and as ingredients for
salads. They are much milder than " scallions," i.e.,
thickly-sown onions, which have no room allowed for the
formation of bulbs.
At Fig. 1 2 is engraved a fragment of a scape or flower-
stem of an affected plant of chives,
enlarged five diameters, to illustrate
the enormous number of disease
pustules with which the flower stem,
together with every other part of a
diseased plant, is covered. It is
curious that the scapes are first and
most affected. When these pustules
or ulcers are examined with a strong
lens they look like little masses of
brownish dust distinctly growing
beneath the epidermis of the host
plant. As the disease runs its
course, and these pustules enlarge
in size, they burst open the epi-
dermis, and the brown or blackish
•X'5- matter, in the form of fine dust, is
FIG. 12 get free in the air r^ pustuies
NEW DISEASE OF ONIONS. „ , .. , .,
Pucdnia mixta, Fi. Frag- or masses of brown dust-like spores,
ment of flower scape of for such they are, are technically
chives. Enlarged 5 dia- cajle^ ^ from tte Greek, SOWS, a
meters. ,
heap.
To understand the nature of the brown dust contained in
the sori, a very small sorus must be selected and removed
from the leaf ; then, with a keen lancet, it must be cut
either transversely or longitudinally in two, and the ex-
posed surface examined. The illustration at Fig. 1 3 repre-
sents a transverse section through one of the smallest sori,
viz. a disease speck, only one-hundredth of an inch across,
magnified 200 diameters. The transparent epidermis of
the scape is seen broken at AA, and some of the constituent
VIIL] NEW DISEASE OF ONIONS. 41
cells of the scape are illustrated at BB ; between these
positions the fungus named Puccinia mixta, FL, is seen.
The parasite consists of club-shaped bodies, each consisting
of a brown head supported on a transparent peduncle or
stem, the whole growing from a matted base of brownish
fungus spawn or mycelium within the substance of the
invaded plant. It will be noticed that the brownish
bodies supported on the slender stems are of two classes, —
one perfectly simple, consisting of a single cell ; the other
of two conjoined cells, or of a larger cell with a distinct
transverse joint or septum across the middle. These
B
•X-20Q-
FIG. 13.
Pucdnia mixta, Fl. Section through a sorws.
Enlarged 200 diameters.
bodies are spores of a peculiar class, possessing the func-
tions of res ting-spores, and called technically teleutospores ;
the latter term meaning "finishing spores," or the last
spores produced ; from teleutao, finishing. From the oc-
currence of the two sorts of spores in each sorus the fungus
under description has been specifically distinguished under
the name of mixta. The two varieties of teleutospore are
farther enlarged to 1000 diameters at Fig. 14 to show
more clearly the cell- walls. Under the microscope the
colour of the spores is rich transparent yellowish-brown ;
the supporting threads are almost colourless. In some
42
DISEASES OF FIELD & GARDEN CROPS. [CH.
species of Puccinia these teleutospores germinate without
rest ; but in the majority of instances they hibernate or
rest in a quiescent state for eight, nine, or ten months ;
FIG. 14.
Pitccinia mixta, Fl. Teleutospores or resting-spores.
Enlarged 1000 diameters.
they then germinate in the peculiar way described
further on under Puccinia graminis, Pers., one of the
fungi of corn mildew. The teleutospores of P. mixta.
vin.] NEW DISEASE OF ONIONS. 43
FL, have not yet been seen to germinate in this country,
and the different experiments made in that direction by
Mr. Phillips all failed, as he at the time foresaw they
would do. The teleutospores of Puccinia mixta, FL, as
found in this country in July 1882, will probably not
germinate before the forthcoming spring or summer ; we
shall then know whether the species is here capable of
being artificially spread on to other members of the genus
Allium. It is supposed that the fungus, long known as
British under the name of Gar lie Rust, — Uromyces alliorum,
D.C., and sometimes as Uredo alliorum, D.C., or Uredo
porri, Sow., is possibly one form of the plant now before
us. It grew with the Puccinia on the same plants of
chives at Shrewsbury. On the Continent there is a
Puccinia named P. allii, Rud., found on Wild Garlic, —
A Ilium ursinum, L. ; and another bearing the same name
is found on A. oleraceum, L., which is the same as A.
virescens, D.C. It is probable that neither of these two
fungi (each named Puccinia allii) are the same with each
other or with our P. mixta, Fl.
On the Continent Puccinia mixta, Fl., is said to grow in
three distinct forms on the same host plants. These forms
have been termed the dEcidium, the Uredo, and the Puccinia
forms ; dEcidium and Uredo are fully described under Puc-
cinia graminis, Pers., one of the fungi of corn mildew. At
present the ^cidium, supposed to belong to P. mixta, FL,
has not been found in Britain. Elsewhere in Europe P.
mixta, FL, grows on various species of Allium, inclusive of
chives, — A. schcenoprasum, L. It has been recorded on
A. acutangulum, Schrad. ; on the Garlic, A. sativum, L. ;
the Rocambole, A. ophioscorodon, Don. ; the Wild Rocam-
bole, A. scorodoprasum, L. ; the Leek, A. Porrum, L. ; A.
rotundum, L. ; the Onion, A. Cepa, L. ; the Welsh Onion,
A. fistulosum, L.; A. carinatum, L. ; A. palustre, Pourr. ;
A. flavum, L. ; A. stellatum, GawL, and no doubt others.
As we have all these plants in our gardens, it is of course
possible, and perhaps probable, that they may soon act as
44 DISEASES OF FIELD & GARDEN" CROPS. [CH.VIII.
host plants here for Puccinia tnixta, FL, as they do on
the Continent, and that we may be unfortunate enough to
speedily see our onions, leeks, and garlic in the condition
of the Shrewsbury chives.
There is an jtEcidium, — the earliest stage, according to
many observers, of a Puccinia, — found abundantly on our
wild garlic, Allium ursinum, L., and named ^Jcidium
Allii, Grev.; which is said to give rise, not to a Puccinia
on Allium, as might have been expected, but to a Puccinia
on Phalaris or Bigraphis arundinacea, Trin., one of our
common river-side grasses. It seems strange that al-
though we have the JEcidium in the greatest abundance,
and the grass upon which its ultimate or Puccinia con-
dition should grow, yet Puccinia sessilis, Schum., — for such
is the name of its other supposed form, — has not yet been
recorded as British.
The only method at present known for preventing
attacks of Puccinia is to burn all affected plants and all
field and kitchen -garden refuse on which the Puccinia
has grown. In decaying refuse the spores hibernate, and
they should be killed during the period of hibernation.
CHAPTER IX.
MILDEW OF ONIONS.
Peronospora Schleideniana, Ung.
ONE of the best known fungus pests of Onions is the
dreaded onion mildew caused by the fungus named
Peronospora Schleideniana, Ung. As its name indicates,
it is a close ally of the fungi of clover mildew, Perono-
spora trifoliorum, D.By., and P. exigua, W.Sm., already
described, and of the fungus of the potato disease, Perono-
spora infestans, Mont.
Peronospora Schleideniana, Ung. , is named in honour of
the botanist Schleiden. The fungus is illustrated, enlarged
200 diameters, at Fig. 15 ; it is shown as growing from
the base of a leaf near the collar of the bulb ; at this
position there is but little leaf -green or chlorophyll in
the cells of the leaves. The spawn or mycelium of the
fungus ramifies amongst the loose cells of the leaf and
sets up decomposition in its progress. At Fig. 1 5, A, the
threads of the fungus may be seen emerging through an
organ of transpiration into the air. The minute open-
ings, " mouths," organs of transpiration or stomata, occur
abundantly on most plants, — generally on the under
surface of the leaves, but also in various other positions.
The general habit of the different members of the genus
Peronospora is to grow within the leaves and stems, and
send their fruiting branches through the stomata into the
air. This habit is fatal to the growth of the host plant,
for the spawn not only causes putrefaction of the inner
cells of the leaf and stem by contact, but the fertile
threads choke up the organs of transpiration and prevent
46 DISEASES OF FIELD & GARDEN CROPS. [CH.
FIG. 15. — MILDEW OF ONIONS.
Peronospora Schleideniana, Ung. Enlarged 200 diameters ; restiug-spores,
400 diameters ; single spore, 1000 diameters.
rx.] MILDEW OF ONIONS. 47
the evaporation of water in the form of vapour from the
plant attacked. So potent are the different members of
the genus for evil that their spawn threads are capable of
pushing aside the plant-cells of the plants attacked either
from without .or within. The mycelium of some species
has also the power of piercing through the cell-walls and
traversing the interior of a plant, not by creeping be-
tween the cells, but by breaking down the cell-walls in
its progress. Putrefaction attends the whole progress of
growth of the invading Peronospora. The complete tree-
like habit of the fungus of onion mildew is shown in
Fig. 15. It will be noticed that the fungus repeatedly
branches and rebranches, and at the ends of all the
minor branchlets the ovate spores or acrospores termed
conidia are borne. These are shown at B, and a single
conidium is farther enlarged to 1000 diameters at C.
The conidia are pale gray or pale lavender in colour, and
are very large in comparison with the conidia belonging
to other species of the genus Peronospora. At the time
of germination the spores usually burst at the side. At
Fig. 16 part of the Peronospora is enlarged to 400
diameters, so that the different illustrations of the genus
in this work may be presented uniform in size. The
difference in size between Peronospora Schleideniana, Ung.,
Fig. 16, and P. exigua, W.Sm., Fig. 2, is very great.
From the irregular mass of protoplasm exuded at the time
of germination mycelial threads and fruiting branches
quickly arise. The fungi of clover mildew are trans-
parent and almost colourless in all their parts, whereas
the onion fungus is more or less tinted with a pale
reddish-gray, a brownish, or a dull violet hue throughout.
This is especially noticeable in the comparatively large
conidia. The Rev. J. E. Vize, of Forden, Welshpool,
has found the resting-spores of this species in decaying
patches upon onions, where the Peronospora in its conidium-
bearing state previously existed. The oogonium, or unim-
pregnated oospore, or resting-spore, is shown at D (Fig. 1 5),
48 DISEASES OF FIELD & GARDEN CROPS. [OH.
•X-400
FIG. 16.— MILDEW OF ONIONS.
Peronospora Schleideniana, Ung. Enlarged 400 diameters.
and mature fertilised oospores or resting-spores at E and
F, enlarged 400 diameters. These bodies carry on the life
of the fungus in a hibernating state during the winter
ix.] MILDEW OF ONIONS. 49
months. Their mode of formation is described under
Peronospora infestans, Mont.
Peronospora Schleideniana, Ung., is specially common
on the garden onion, Allium Cepa, L., and on the Eock,
Welsh onion, or stone leek, named Allium fistulosum, L.
The latter plant does not form a true bulb, but is sown
to form small green onions for spring salads. The
fungus attacks numerous other species of Allium, and is
even suspected on lilies and amaryllids. It is sometimes
extremely common and virulent, attacking the onions
early in the season, and so preventing the bulbs from
reaching perfection. The chief point of attack is the
leaves in an early stage of growth, and to such an ex-
tent is the work of destruction sometimes carried on that
the entire onion plant may be seen covered with one mass
of whitish-gray semi-transparent bloom. In bad cases the
onions are left as one offensive mass of putrescence.
One form of mildew is caused by the attack of a
Fusisporium named F. atrovirens, Berk. This fungus is
not unlike F. Solani, Mart., already described, but the
spores are grayish -green or greenish-black in colour and
more decidedly curved. The disease begins with the
exhibition of grayish gelatinous spots, which speedily be-
come confluent. The Fusisporium often accompanies the
Peronospora. .Both mildews materially affect the crop of
seed. Practical growers say mildews are favoured by an
extremely dry season, as in the mildew of peas and roses,
and also by a wet and cold season, as well as by bad culti-
vation. Deep trenching is generally advocated for the
prevention of onion mildew.
A good plan for the avoidance of mildew in onions is
to sow the seeds in autumn ; by following this plan the
onions are able to make good strong growths before the
appearance of the mildew in the following spring. This
is perhaps the only reliable plan for obtaining sound
onions of a large size in districts subject to attacks of
mildew. Autumn sowing is advocated by onion growers.
E
50 DISEASES OF FIELD & GARDEN CROPS. [CH. ix.
This is said to prevent the attacks of the onion-fly, for it
is believed that the flies select the younger onions for the
deposition of their eggs in preference to those of more
mature growth.
Where possible all mildewed material should be burnt,
for no fact is better known than that mycelium or fungus
spawn is often perennial.
Like many other plant ailments, the diseases of onions
require investigation. Several forms of disease are known
to growers, of which no explanation has at present been
forthcoming. In one form the whole crop turns sickly
yellow just before ripening, the tops soften, the bulb be-
comes detached, the roots decay, and the entire growth
soon becomes rotten. Another disease causes the onion
to become thick and soft-necked, the bulb in proportion
being small. Sometimes mildew attacks the full -sized
and apparently not thoroughly -ripened bulb after it is
harvested, and commences from the outside.
CHAPTEE X.
MOULD OF ONIONS.
Mucor subtilissimus, B.
THE fungi known as Mucors or Moulds (muJces, mould)
are extremely common on decaying bulbs, fruits, pro-
visions, etc. They are said not to be the immediate cause
of decay, but there can be no doubt that they greatly
accelerate putrescence when they grow upon exposed or
injured places. Sometimes they grow in the inner sub-
stance of plants, like the one under description, which has
been named by the Kev. M. J. Berkeley, in reference to its
extreme smallness and delicacy, Mucor subtilissimus.
A species of fungus closely allied to the onion Mucor is
the common Mucor mucedo, L., so frequent on paste, jam4
damaged fruit, etc. The onion Mucor differs from all its
allies, t in its extremely small size ; it is said to be the
most microscopic of all fungi found in Great Britain.
In the Mucor disease the whole substance of the neck
of the onion near the bulb, and sometimes the bulb itself,
is traversed by fine threads of mycelium, and in the midst
of this mass of spawn may be seen innumerable black
atoms like minute grains of gunpowder. These little
grains have been described as fungi under the name of
Sclerotium Gepce and Sclerotium cepcevorum, B. ; the nature
of Sclerotia is described under Peziza postuma, B. and
Wils. The present Sclerotia not only differ from the
potato Sclerotia in their much smaller, almost microscopic
size, but also in their less compact and more filamentous
structure. The less compact a Sclerotium is, the more
readily it will germinate, and in the present instance the
52 DISEASES OF FIELD & GARDEN CROPS. [OH.
Sclerotium will protrude germ tubes in a few hours if
placed in a drop of water. The whole process of growth
may be easily watched under the microscope. On germi-
nation in water, the filamentous mycelium of which each
Sclerotium is formed, protrudes, elongates, and branches in
a flexuous manner in all directions ; this spawn is some-
times jointed and sometimes free from joints, and whilst
in water no farther progress is made in growth beyond
the protrusion of these threads. As soon, however, as the
mycelium reaches the edge of the water, a change takes
place, and the branches become furnished on their tips
with minute globose heads, technically termed Sporangia
or spore cases. These extremely minute spherical heads
at length become filled with elliptic spores or sporidia
(so called because they are produced within a spore case),
which, on the natural bursting of the Sporangium, or spore
case, are set free in the air. The little oval sporidia now
soon germinate and reproduce the species. The small
mycelial threads protruded from the sporidia are capable
of forming little white knots, which at length become, on
exposure to the air, the minute black granules termed
onion Sclerotia.
Now, although these small dark-coloured granules will
germinate very readily under favourable conditions in
water, it must not be supposed that every example germi-
nates after a few hours' rest. There can be no doubt that
this Sclerotium condition answers precisely the same purpose
in the onion as it does in the very large Sclerotium of the
potato ; that is, it carries on the life of the fungus in a
hibernating state through the winter.
Individual fungi vary in their habits of growth pre-
cisely in the same way as they vary in their specific
characters. No hard and fast line can bind down every
individual fungus to specific characters or to habits of
growth. For instance, in Puccinia mixta, FL, already
described, although nine hundred and ninety-nine teleuto-
spores out of a thousand may go to rest for several months,
x.] MOULD OF ONION'S. 53
the thousandth example will germinate at once ; and, in
the allied fungus of the Hollyhock disease, Puccinia
malvacearum, Mont., where nearly all the teleutospores
germinate as soon as ripe without any rest, it may con-
stantly be observed that a certain number of erratic ex-
amples do certainly hibernate. Provisions of this nature
are favourable to the existence of fungi, for, if plants were
strictly bound by inexorable laws as to habit of growth,
they might, by some untoward circumstance, be all sud-
denly swept out of existence. When unfavourable con-
ditions arise, a species may be kept in existence by indi-
viduals of erratic habit.
No doubt the attacks of the onion Mucor may be
palliated by the destruction by fire of infected plants
and refuse. By this means all Sclerotia and perennial
spawn will be destroyed.
At present Mucor suUilissimus, B., has only been re-
corded as growing upon the cultivated onion, but it
doubtlessly grows upon other and allied plants.
CHAPTER XL
ONION SMUT.
Urocystis cepulce, Far.
IN 1879 a disease of onions appeared in France, known
as the " American Onion Smut," Urocystis cepulce, Farlow,
a disease which had hitherto "been confined to America.
Fungi belonging to Urocystis are parasitic on living plants,
and familiar allies in this country are the bunt of wheat,
Tilletia caries, Tul. ; corn smut, Ustilago carlo, Tul. ; the
common black smuts of violets and colchicum, named
Urocystis violoe, B. and Br., and U. colchici, Tul. ; potato
smut, Tubercinia scabies, B. and Br., and many others.
It is probable that we already have this disease in Britain,
as onion growers have sorely complained of late of their
onions falling into a dusty black mass after harvesting.
Whether this fungus is really distinct from the common
smut of colchicum, U. colchici, Tul., seems somewhat un-
certain. The name of the parasite is derived from uro
to burn, Jcystis a bladder, and ccepa the onion plant.
CHAPTEE XII.
NEW DISEASE OF GEASS.
Isaria fuciformis, Berk.
DURING the last few years the grass of the southern
counties of England has been attacked, as far as this
country is concerned, by a curious and possibly new form
of disease. Some farmers, however, state that they have
noticed the malady for many years past ; in one instance
twenty years has been mentioned. The disease has ap-
peared chiefly on sandy and chalky soils, and is apparently
absent from clay districts. The grasses chiefly attacked
by the disorder are the Festucas, and notably Festuca
ovina, L., a valuable pasture grass especially relished by
sheep. The fungus which causes the disease does not
generally grow on young grass, but the growths appear
to be almost peculiar to the old grass, first appearing in
September, and continuing in ordinary seasons till the
following January. In mild winters the fungus may be
seen on the grass till March.
The Rev. M. J. Berkeley originally described this
fungus on grass in the Journal of the Linnean Society for
1873, vol. xiii. p. 175, under the name of Isaria fuci-
formis. The word Isaria is derived from the Greek, and
simply means equal, in reference to the simple equal
growth of many of the species ; and fuciformis is founded
on the word phukos, the Greek name for sea- weed ; the
word fuciformis therefore means, like a Fucus or sea-weed.
The specific name is an appropriate one, for the resem-
blance of Isaria fuciformis, B., to some of the small linear
red sea -weeds is strong. Mr. Berkeley describes the
56 DISEASES OF FIELD & GARDEN CHOPS. [CH.
fungus as growing en
some germinating cereal from
Mount Gambier in Australia ;
his description is very brief.
He writes : "Pallid, slender,
filiform, sparingly branched,
branches acute, spores very
minute, globular."
The general appearance of
the fungus, as seen growing
upon a panicle of sheep's
fescue, Festuca ovina, L., is
shown twice the natural size
at Fig. 17. The fungus tufts
spring from an effused, muc-
ous, pinkish base of spawn or
mycelium, which has a tend-
ency to glue different parts of
the grass together as shown
at Fig. 18, where three grass
stems invaded by the fungus
are illustrated five times the
natural size. The tufts grow
on the stems, leaves, and every
other part of the grasses
affected, and the growths show
a marked tendency when ripe
to drop off and fall to the
ground. There can be little
doubt that this falling of the
fungus tufts to the ground is
one of the means by which
the propagation of the pest is
FIG. 17.— NEW DISEASE OF GRASS, aided. The Colour of the fun-
Panicle of Sheep's Fescue Grass in- gus is sometimes extremely
vaded by Isaria fuciformis, Berk.
Twice the size of nature.
similar with pink coral.
X 2
rangng n tint
from blood-red to a pink hue
Were it not for this bright
xri.] NEW DISEASE OF GKASS. 57
colour the fungus might be easily overlooked, and it must
be remembered that the original description states the
plant to be " pallid." A pallid condition of this fungus
may therefore possibly be widespread and unnoticed.
In some places and seasons the fungus may be always
pallid, and so virtually invisible. When the fungus
FIG. 18.
Stems of Sheep's Fescue Grass, with Isaria fuciformis, Berk.
Enlarged 5 diameters.
exhibits the scarlet colour, the tint appears to be per-
manent, for no change has been observed in the colour of
our dried herbarium examples. In this permanency of
colour it resembles the beautiful and closely-allied orange-
coloured Anthina flammea, Fr., so common on the dead
beech leaves of our autumn woods.
In reaching England from Australia this fungus has
taken the same course as the Pucdnia of our hollyhocks
and the Capnodium of our Thuyas.
The structure of 'Isaria fuciformis, B., is very simple,
as is the case with all imperfect fungi. The whole sub-
stance of the parasite is one compacted mass of minute
cells or exceedingly small transparent bladders, as illus-
58 DISEASES OF FIELD & GARDEN CROPS. [en.
X-IOO-
FIG. 19.
Tip of one of the minor branches of Isaria fuciformis, B,
Enlarged 100 diameters.
x- 1,000
FIG. 20.
Tip of branchlet of Isaria fuciformis, B. , with conidia.
Enlarged 1000 diameters.
trated at Fig. 19, enlarged 100 diameters ; the larger
cells occupy the centre of the branches, and the smaller
cells form the outside stratum. As the smaller outside
XIL] NEW DISEASE OF GRASS. 59
cells gradually reach the tips of the branches they break
into innumerable extremely small globose spores or con-
idia, as shown at A, Fig. 20, enlarged 1000 diameters.
It should be noticed here how closely these Isaria
spores or conidia resemble in size the Torrubia, spores at
J, Fig. 22, and how extremely small they are as compared
with the spores or conidia of the mildew of onions, Per-
onospora Schleideniana, Ung., illustrated to the same scale
at C, Fig. 15.
Isaria fuciformis, Berk., is a remarkable fungus, for no
other British species of Isaria is known to grow on a
living plant. Some species grow on dead flowers and
dead twigs and stems, others on decaying fungi, one upon
cat's dung, and another on dead spiders. Some species of
FIG. 21.
Living Wasp, with fungus growths. Twice the size of nature.
Isaria grow upon dead hymenopterous insects, others grow
upon living bees and wasps. One species of Isaria grows
in this country on dead pupce, another on dead moths ;
but it is known that these creatures are attacked by the
60 DISEASES OF FIELD & GARDEN CROPS. [OH.
fungus whilst they are still alive. The illustration at Fig.
21 shows a wasp, twice the natural size, as caught in a
living state in this country, languidly flying about with a
fungus burden, which may either be a Stilbum, as figured
by Charles Eobin in his Histoire Naturelle des Vty&aux
Parasites (Paris, 1853), or an abnormal form of the
Isaria condition of the fungus known as Torrubia spheco-
cephala, Kl.
The genus Isaria amongst fungi represents an early or
larval state of another and more perfect genus of fungi
named Cordiceps, from the Greek Jcordyle, in allusion to its
clublike shape, a genus termed in more modern books
TorruUa.
Now the species belonging to Cordiceps or Torrubia
(certainly in many instances the perfect state of Isaria)
are equally curious in their habits with Isaria itself;
some grow on larvae and pupse, even when buried in the
ground ; two grow on subterranean truffles, and it is
remarkable that two distinct species of Torrubia attack
two equally distinct truffles ; another species grows on
Wych elm twigs. It is well to mention at this place
that Torrubia is very closely allied to the fungus named
Claviceps, which is the perfect condition of the dangerous
ergot of rye.
As it is quite possible that Isaria fuciformis, B., may
be an early condition of a Torrubia belonging to an insect
or plant host, a brief description of one of the species of
Torrubia may be useful. The genus Torrubia is named
in honour of a Spanish botanist who wrote a work on
" vegetable wasps." A Torrubia parasitic on the truffle
named Elaphomyces muricatus, Vitt., is fairly com-
mon. This curious parasite is named Torrubia ophio-
glossoides, Tul., the specific name having reference to a
fanciful resemblance between the head of the perfect
fungus and a serpent's tongue. This Torrubia is repre-
sented, natural size, at A, Fig. 22, attached to its peculiar
host or truffle, B. The club-shaped top of the Torrubia
XII.]
NEW DISEASE OF GEASS.
61
FIG. 22,
Perfect condition of Torrubia ophioglossoides, Tul. ; parasitic on the
truffle Elaphomyces variegatus, Vitt.
Natural size, and enlarged 5, 200, and 1000 diameters.
62 DISEASES OF FIELD & GARDEN CROPS. [OH.
at A is shown in section. This upper part of the fungus
is technically termed the stroma, meaning a coverlet ; and
immediately under the surface of this clublike stroma or
coverlet are numerous minute embedded flasks, as illus-
trated. These flasks are termed perithecia, in reference to
their function, which is to enclose a number of transparent
bladders or thecce. The number of contained bladders in
each perithecium is about one hundred. At C some of
the perithecia embedded in the stroma are enlarged five
diameters, and the upper perithecium is shown in the act
of discharging the contained spores. If, with the point of a
needle, we remove a few of the transparent thecae, bladders,
or asci, and magnify them 200 diameters, we shall see them
as at D E F. An ascus is represented, packed with its
eight spores, in situ at D ; the top of the open ascus is
shown at E with the eight hairlike spores escaping, and
F shows an empty ascus after all the spores have been
expelled. If we now leave the Torrubia and examine the
truffle, we shall find its inner mass densely packed with
its own spores, also contained in transparent bladders ;
but the truffle spores are spherical in form, blackish-brown
in colour, and packed in twos, threes, or fours in the asci
(not in eights as in the Torrubia}. Some of the Elapho-
myces spores in an ascus are illustrated at G, enlarged,
like the asci and spores of the Torrubia, to 200 diameters.
The spores, or necklace-like chains of sporidia, of Tor-
rubia ophioglossoides, Tul., are amongst the most wonderful
objects of the vegetable kingdom. One of the eight chains
from an ascus is enlarged to 1000 diameters at H.
To sum up the characters, the Isaria is perfected early
in the season, and is capable of reproducing itself by its
own spores or conidia, as shown in Figs. 19 and 20.
Later in the season the mycelium of Isaria often produces
a Torrubia, A, Fig. 22 ; and the latter plant, instead of
producing free, dustlike spores from the naked apex of
its branches, as in the Isaria, produces spores or sporidia
in the form of long chains enclosed in transparent flasks,
XIL] NEW DISEASE OF GRASS. 63
these flasks being enclosed in larger bladders termed
perithecia, and the whole embedded in the club-end or
stronia of the Torrubia. It is obvious that in the latter
position the sporidia are well protected by three different
enclosing walls, one within the other, and there can be
no doubt that these contrivances aid the contained spores
in tiding over vicissitudes of rain, drought, and frost during
winter. When the spring conies the stroma softens, the
mouths of the perithecia open, the asci sail out and burst,
and the chains of spores are set free in the air. These
chains speedily fall to pieces, as illustrated at J, enlarged
1000 diameters, and each fragment or sporidium on
germinating is capable of producing, not a Torrubia but
an Isaria. How small these numerous reproductive
bodies are may be judged from the fact that it would
require two hundred millions of them to cover a super-
ficial inch. Every plant of Torrubia ophioglossoides, Till.,
sets free at least ten millions of these reproductive bodies
every spring.
Before dismissing Torrubia ophioglossoides, Tul., a curious
fact regarding it may be mentioned. As it is parasitic
on an underground truffle named Elaphomyces muricatus,
Vitt, the question presents itself, How can the exceedingly
minute spores of the Torrubia reach the subterranean
truffle, buried, as it is, some four or five inches beneath
the ground, in places often thickly covered with brambles,
ferns, and moss ? The explanation of this phenomenon is :
The spawn from which the Torrubia springs grows in the
first instance, over a common moss named Mnium hornum,
Hedw., sometimes given as Bryum hornum, Sw., illus-
trated natural size at Fig. 23. The spawn or mycelium
of the Torrubia is yellowish, and when this yellow spawn
once fixes on the moss it goes from leaf to leaf, from stem
to stem, and from root to root, till sometimes a large
patch of this common moss is covered with the yellow
sticky threads of the curious Torrubia mycelium, which is
really now running from plant to plant in search of truffles.
64 DISEASES OF FIELD & GARDEN CROPS. [OH.
It is an above-ground truffle-hunting mycelium. If there
are no truffles in the wood, the mycelium, of course,
perishes ; but in some other wood where the moss grows,
the truffles will certainly occur. When the moss does
grow in the same wood with the truffle, the parasite is
certain to descend by the roots, and so find the subterra-
Fio. 23.
Mnium Jiornum, Hedw., invaded by the mycelium of Torrubia
ophioglossoides, Tul.
nean Elaphomyces. The truffle usually grows some four
or five inches beneath the surface of the ground, and
seldom so near as two inches, as shown in illustration at
Fig. 22, where KK shows the ground line, and LL the
spawn belonging to the TorruUa connected with the truffle
below at B.
It is a curious fact that field mice, and probably several
other small mammals, are extremely fond of the truffles
named Elaphomyces (from elaphas, an elephant, and
mukes, a fungus), and these little animals are continually
xii.] NEW DISEASE OF GRASS. 65
scratching about for the truffles. There is a kind of race
then between the mammal and the mycelium for the
underground fungus.
Botanists, who search for truffles always know where
the Elaphomyces is to be found by looking for the spawn
of the Torrubia on the moss. The spawn-covered moss
points out the position of the truffle, as surely as the
bird named the Great Honey Guide, Indicator major,
Steph., leads hunters to bees' nests in Africa.
A closely allied species of Torrubia, named T. militaris,
Tul. (in the older books Cordiceps or Sphceria), of bright
scarlet colour, is extremely common in Britain in the
autumn and early winter, growing from dead pupae buried
in the ground. The fungus resembles a scarlet club,
about an inch or an inch and a half high. It grows in
pastures and grassy places, where larvae have buried them-
selves. The mycelium grows within the body of the
pupa, and the scarlet club commonly grows from the first
joint behind the head. The anatomical characters of T.
militaris, Tul., agree generally with T. ophioglossoides, Tul.
The Isaria disease possesses considerable interest on
account of the popular belief that it is capable of greatly
injuring, and indeed of killing the cattle that feed upon the
grass infected with it. In September 1880 an instance
occurred where two cows died in an Jsarm-infected dis-
trict from an affection of the lungs ; and when a post-
mortem examination was made it was found that the lungs
were covered with a fungus-like growth, not unlike, it is
said, the appearance presented by the throat in diphtheria.
The veterinary surgeon who conducted the examination
declared his opinion that the fatal ailment had been con-
tracted from the /sano-infected grass. The same medical
practitioner is said to have fed two rabbits on infected
grass only, and that they both died therefrom. It is,
perhaps, unnecessary to say, that these cases are far from
being proved. At the same time, it would not be wise to
immediately say that such cases are impossible. In favour
66 DISEASES OF FIELD & GARDEN CROPS. [OH.
of the doctor's view it may be called to mind that several
members of the germs of fungi to which Isaria belongs,
grow naturally on living animals.
It is not usual for fungi to grow on living or, indeed,
dead animal material, yet botanists are familiar with
several such examples other than the ones already cited.
The familiar fungus of the salmon disease, Saprolegnia
ferax, Kutz., is one, and the white dusty fungus named
Empusa, muscce, Cohn., so common on flies on our window
panes in autumn, another. This is said by some observers
to be merely a second condition of the Saprolegnia. The
ringworm fungus Oidium porriginis, Mont., is another
example ; Microcera coccophila, Desm., a parasite of Cocci,
— insects of the American blight class, — is a third ;
Onygena equina, Pers., which grows on the hoofs of dead
horses, is a fourth, and the list might be greatly extended.
Fries has described an Agaricus, named by him A.
nauseosus, growing on the carcass of a wolf, and A.
ostreatus, Jacq., has been seen in this country by Mr. C.
B. Plowright growing on the dead body of a stranded
whale ; Onygena apus, B. and Br., is not uncommon on
bones. These instances are not mentioned with the view
of showing that Isaria fuciformis, B., is capable of killing
cattle, but to indicate that other and, in some instances,
closely allied fungi can support themselves on living and
dead animal substances.
It is extremely difficult to suggest any means for the
destruction of the Isaria^ partly because the fungus falls
from the grass to the ground on the slightest touch, and
partly because so very little is known of the fungus or its
habits. The whole subject requires investigation. Some
crops might be saved by removing the greater part of the
grass before September, or by the substitution of some
crop on which the Isaria could not grow. As humidity
probably favours the growth of the pest, good and careful
drainage might prevent its spreading. No doubt many
insects carry the conidia, or spores, from place to place
XIL] NEW DISEASE OF GRASS. 67
on their bodies, and so infect previously untainted
districts.
Towards the end of 1883 Mr. Greenwood Pirn, M.A.,
F.L.S., and Dr. E. P. Wright, A.M., F.L.S., detected Isaria
fuciformis, B., growing in a new position, viz. on grass
belonging to a silo at the Albert Model Farm, Glasnevin,
Co. Dublin. Mr. Pirn kindly forwarded examples to us,
and he soon afterwards published an illustrated account
of the discovery in the Gardeners' Chronicle for 22d Decem-
ber 1883. Mr. Pirn's examples were remarkable for
being infested with a parasitic fungus, and one apparently
till now undescribed. The parasite grows on the Isaria,
breaks up its tissues, and more or less absorbs its crim-
son colour. The parasite is a Saprolegnia allied to S.ferax,
Kutz., of the salmon disease, but different in many im-
portant characters.
The new parasite, which may be termed Saprolegnia
pliilomukes, W.Sm. (from sapros, decayed ; legnon, a fringe
or border ; phileo, I love ; and muJces, a fungus), is illustrated
at Fig. 24, enlarged 400 diameters. The circular bodies
are sporangia, zoosporangia, or spore -cases of unusually
large size, and filled with small motile spores or zoospores.
In the largest sporangium illustrated it will be seen that
the zoospores are germinating within the sporangium, and
protruding their germ tubes through its gelatinous wall.
A remarkable character in this parasite is found in the
septate or jointed mycelium, an unusual character in the
Saprolegniece, in the mycelium carrying numerous conidia,
as at AA, and in the sporangia and mycelial threads often
becoming confluent, as at B, C. In the Dublin examples,
the sporangia were so abundant that all parts of the
Isaria threads were covered, they were so crowded together
that they took pentagonal and hexagonal instead of cir-
cular forms. Many sporangia were sessile, or intercalated
in the mycelium, whilst others were shortly stalked.
Antheridia (male organs described under the fungus of
the potato disease), as at D, were rare : the jointed my-
68 DISEASES OF FIELD & GARDEN CROPS. [CH.XII.
celium formed a dense transparent stratum over the host
plant. In some places the parasite was colourless, like
FIG. 24.
Parasite of Isaria fuciformis, B. ; Saprolegnia philomukes, W.Sm.
Enlarged 400 diameters.
the better known species of Saprolegniece ; in other places
it was rose-coloured, from its absorbing the colour of the
red Isaria.
Saprolegnia philomukes, W.Sm., zoosporangia, very large,
thick-walled, sessile, shortly stalked, or intercalated in the
mycelium, sometimes confluent with each other, all bear-
ing zoospores, which often germinate whilst still in situ.
Antheridia elongated, rare ; mycelium profusely septate,
somewhat torulose, filled with colourless or rose-coloured
protoplasm, and bearing many small abortive sporangia
or conidia.
We have seen a similar plant on fungi, with oospores
as well as zoospores.
CHAPTER XIII.
STRAW BLIGHT.
UNDER the name of Straw Blight agriculturists are well
acquainted with a peculiar diseased condition of the living
x-5
FIG. 25.— STRAW BLIGHT.
Fragments of diseased Wheat Steins. Enlarged 5 diameters.
stems of wheat, barley, rye, and other grasses, which
70 DISEASES OF FIELD & GARDEN CROPS. [CH.
commonly makes itself apparent in midsummer by the
impoverished appearance of the flower-spike. Sometimes
the disease occurs in spring, and then it often proves fatal
to the plants attacked.
Straw blight is caused by the attack of a fungus on
and in the straw at a point close to or very near the
ground. The fungus growth seldom reaches so far up the
stem as the second or third joint, the attack being more
frequent below the first joint from the root, and close to the
ground. It is superficially recognised by brownish disease-
spots outside the straw, as illustrated at AA, Fig. 25,
enlarged 5 diameters ; but if the straw is carefully cut
longitudinally with a sharp knife, it will be seen that the
disease is by no means superficial. The disease-spot goes
through the solid wall of the hollow stem, and in typical
examples the hollow part will be more or less filled with
loose flocculent material, as illustrated at BB. This floccu-
lence is really the mycelium or spawn of a fungus. Hav-
ing now obtained a clue to the nature of the so-called
blight, an excessively thin and transparent atom must be
sliced off from the exposed surface of one of the brown
disease-spots with a lancet, and this slice highly magnified.
If we enlarge this atom 200 diameters, and examine at it
as a transparent object under the microscope, we shall prob-
ably see it as illustrated at Fig. 26. The base of the
illustration shows the cells of the solid part of the straw
in transverse section, whilst the main part of the illustra-
tion shows the stem in longitudinal section : the bottom, in
fact, represents the base of the minute transparent atom
sliced off. We now see the spawn threads distinctly ; they
are transparent or nearly so, and so fine and attenuated
that (as may be seen by the thinner lines of the illustration)
they are less in thickness than the walls of the microscopic
cells, of which the straw stem itself is built up. They will
be seen to branch, and apparently pierce the cell-walls
both vertically and horizontally, and in old examples to
almost fill the hollow of the stem. A close examination
XIII.]
STRAW BLIGHT.
71
will show that in a few places there are transverse joints,
stops, or septa in the threads of spawn, as illustrated in
the separate thread on the left of illustration, enlarged
\ff
G. 26.— STRAW BLIGHT.
Fragment of diseased Wheat Stern.
Enlarged 200 diameters.
1000 diameters. It is a most unusual thing to see any
fungus spawn without these stops, although they are
72 DISEASES OF FIELD & GARDEN CHOPS. [CH.
mucli more common in some fungi than in others. The
stops or transverse partitions represent a slight temporary
rest in the growth of the threads. For a brief period
the spawn has exhausted its powers of extension, and a
septum or transverse wall is formed across the thread,
and from this point a new and vigorous start is com-
monly made.
As the fungus -spawn of straw blight apparently
possesses the power of piercing the walls of the cells
of which the straw is built up, it follows that all
parts of the straw are liable to be infested and
pierced. This piercing causes a fatal injury to the cell-
structure, and every farmer knows that if his plants are
attacked by this blight whilst the crops are still young,
the growth will be stopped. The walls being pierced the
flow of sap is arrested, and the upper part of the plant
perishes from want of nutriment. In the older plants
the stem is often observed to throw out new roots from
the joint above the diseased part ; and if these new roots
are able to reach the earth, they sometimes carry on the
life of the plant in place of the old roots, which are more
or less cut off from the stem. The new roots are never
quite effectual in keeping up the supply of food and life,
and a stem once attacked by straw blight is said to never
entirely recover.
It is curious that no one has at present recorded the
perfect form of the fungus which must, under favourable
circumstances, arise from this spawn. Such a barren
condition in mycelia is, not uncommon ; the fact often
holds good with fungi that there is an enormous
development of mycelium but no perfect fungus. There
are many more or less barren mycelia well known to
botanists, such as the orange-coloured fungoid growths
known as Ozonium; Byssus, Rhizomorplia, and many
others. In some instances, as in the grape mildew, the
spawn proceeds one step farther and produces what is
termed an Oidium, which, like the Isaria last described,
xiii.] STRAW BLIGHT. 73
is really a kind of larval condition of some more per-
fect fungus, the ultimate form of which has perhaps
never been seen, or, if seen, has not been recognised.
The nature of Oidium is described under the Oidium
of the turnip and under grass blight, Erysiphe graminis,
D.C.
Grasses both wild and cultivated, living and dead, are
subject to the attacks of so many fungoid assailants that
it would be almost useless to guess at what the perfect
form of the fungus of grass blight might be. The
Graminecs and Cyperacece are unusually subject to the
attacks of fungi. Many of the pests are, however, super-
ficial, and do not possess the power of piercing and
traversing the cellular tissue. All, however, are ob-
jectionable, as they not only reduce the crop, but more
or less lessen its value as food. The phenomenon of
piercing the cells in the fungus of straw blight reminds
us of the corrosive mycelium of the fungi of the potato
disease found under Peronospora. Several species of
Fusisporium have been detected on cereals, but the
mycelia of these as seen by us do not well agree
with the spawn of straw blight. A fungus named
Fusisporium insidiosum, Berk., a parasite of the grass
named Agrostis pulchella, Kunth., is by no means well
known, and requires further attention.
The loss to farmers from straw blight ranges from one-
half to one -fiftieth part of the crop, according to the
virulence of the attack ; but the blight is erratic in its
appearance, sometimes temporarily vanishing, and then
returning with great activity. Like some other mildews
and blights, but not all, straw blight is fostered by a
continuance of warm, wet weather.
As the growth of straw blight is promoted by moisture,
it seems probable that if the quantity of water about our
cereal crops in wet seasons could be lessened by perfect
drainage, the amount of destruction from straw blight
would be less. Wheat and barley are not generally
74 DISEASES OF FIELD & GARDEN CROPS. [CH.XIII.
grown in wet, peaty places, yet in the alluvial flats be-
longing to some rivers, and where good drainage is
difficult, these crops may often be seen. No doubt per-
fect drainage would considerably lessen the losses com-
monly entailed by attacks of straw blight.
CHAPTEE XIV.
SURFACE MILDEW OF TURNIPS.
Oidium Balsamii, Mont.
THE mildew of turnips, named Oidium Balsamii, Mont.,
is often confounded with the true putrefactive mildew of
the cabbage tribe, named Peronospora parasitica, Pers.
The two are indeed so much alike to the unaided eye
that it is often impossible for even an experienced ob-
server, without a lens, to distinguish one from the other.
As if to make the subject still more involved, it fre-
quently happens that the two fungi grow in company on
the same host plant. They are, however, wholly dis-
tinct from each other, both in habit and structure.
The name Oidium is derived from the Greek oon, an
egg, and eidos, resemblance, and refers to the usual egg-
shaped form of the spores or conidia. In the present
instance the generic name is not very appropriate, for the
conidia are somewhat barrel-shaped. The specific name
Balsamii was given in honour of Balsamo, a Milanese
gentleman, who first noticed the species. When first
detected the fungus was growing on a Continental species
of Mullein, named Verbascum montanum, Schrad. We
have this plant in our gardens, but the fungus is more
common here on the Black Mullein, Verbascum nig-
rum, L. ; it also grows on cultivated strawberries. In
the latter case the fungus makes its first attack on the
leaves, and then speedily invades with increased vigour
the flowers and footstalks, ultimately inducing the
wretched appearance so well known in connection with
grape vines when attacked by the allied fungus named
76 DISEASES OF FIELD & GARDEN CROPS. [OH.
Oidium Tuckeri, B. Although several species of Oidium
have been recorded from Scotland, Oidium Balsamii,
Mont., has not yet been detected there. If it really
grows so far north, it could hardly have been over-
looked, as it is a remarkable species.
Oidium Balsamii, Mont., first attracted attention as a
pest of turnips in September 1880, when Prof. James
Buckman, F.L.S., of Bradford Abbas, Dorsetshire, saw the
fungus growing in such profusion over hundreds of acres
of Swede turnips that the boots and clothes of persons
walking through the turnip fields were whitened with
the spores. Until 1880 the fungus was not supposed
to be common in Britain ; and it is remarkable that the
same fungus should be found growing upon three different
natural orders of plants, viz. the Scrophulariacece, the
Rosacece, and the Cruciferce. Some farmers say the plants
produced from early sown seeds are the most subject to
this mildew. It first attacks the lowermost leaves, and
then quickly covers every part of the affected plant.
The presence of this pest, which is now known to be a
common and injurious mildew of turnips, generally fore-
shadows a deficiency of roots.
To the unaided eye the foliage of affected Swedes is
white on both sides when attacked by the mildew ; but
when seen under a low power of the microscope this
white coating resolves itself into a dense felted mass of
spider-web -like threads, dotted all over with innumerable
barrel-shaped spores.
The higher powers of the microscope are required to
show the exact nature of the Oidium of turnips. A
minute fragment must be cut from an infected place on
a turnip leaf, and from this fragment an exceedingly thin
transparent slice should be cut. When placed in a dry
state under the microscope it must be specially noticed
that the fungus growth is wholly superficial, and that no
spawn threads belonging to the Oidium occur within the
leaf. In this respect the Oidium essentially differs from
XIV.]
SURFACE MILDEW OF TURNIPS.
77
the Peronospora, next described in this work. When a
minute slice, as just mentioned, is examined under the
microscope and enlarged 400 diameters, it will be seen as
illustrated at Fig. 27. The fungus grows on both sur-
faces of the leaf, and springs in both positions from a
X-400
FIG. 27.— SURFACE MILDEW OF TURNIPS.
Oidiim Balsamii, Mont. Enlarged 400 diameters.
dense stratum of matted and jointed spawn, as shown at
AA. From these horizontal spawn - threads arise in-
numerable vertical club-shaped growths, each club being
furnished with three joints and surmounted by a barrel-
shaped spore or conidium, as shown at BB. The cells at
78 DISEASES OF FIELD & GARDEN CROPS. [OH.
CC, represent the lower cuticle of the leaf, whilst the
openings at D D D, show the stomata or organs of trans-
piration. It will be observed that the spawn of the
invading fungus does not enter the stomata or traverse the
intercellular spaces of the leaf, such as are shown at
E E E. The barrel -shaped spores or conidia of this
fungus are so numerous that more than 10,000 are
produced on every square inch of leaf surface, and every
FIG. 28.
Oidium Balsamii, Mont.
Germinating Spore enlarged 1000 diameters.
turnip leaf will carry on its two surfaces a million or
more of these reproductive bodies.
The spores germinate very readily, for they have only
to be dusted on to clean glass and kept moist, under a
bell-glass, when they will be seen to germinate at once as
illustrated at Fig. 28, enlarged 1000 diameters. The
germinating conidium bursts at one corner as at A, and
from this corner the contained protoplasm or vital
xiv.] SURFACE MILDEW OF TURNIPS. 79
material streams in threadlike form, and from this thread
new clubs immediately arise. Four of these bodies, in
different stages of growth, are shown at B, C, D, and E.
The surface of one of the organs of transpiration is shown
at F, over the opening of which a thread of mycelium has
This Oidium chiefly injures the turnip by weaving a
thick web of mycelium over the organs of transpiration.
The spawn effectually stops the passage of watery vapour
from the interior of the affected plant, and so puts an
end to one of its chief vital functions. The general
result is an arrest of growth, and ultimately a poor crop
of roots.
Oidium Balsamii, Mont., is supposed to be an early
condition of some more perfect fungus, probably an
Erysiphe, such as one sees on the hop, on roses, on mil-
dewed grass, and on peas. The two latter species of
Erysiphe are referred to in detail farther on in this work.
Important as this Oidiuin is to agriculturists, no one at
present has worked out its life history or knows whence
it comes, where it goes, what other form it takes, or how
it hibernates through the winter. The fungus is more
prevalent when a humid September follows on a dry
August.
CHAPTER XV.
PUTREFACTIVE MILDEW OF TURNIPS AND CABBAGES.
Peronospora parasitica, Pers.
THE fungus which causes this disease frequently accom-
panies Oidium Balsamii, Mont., already described, but it
differs entirely from it both in anatomy and nature.
When a Peronospora infected leaf is examined with the
unaided eye, the thick white bloom on both sides of the
leaves, as in Oidium Balsamiij Mont., is never seen.
The Peronospora appears as a thinner, more scattered bloom
on the under side of the leaves only, and generally
borders pallid, discoloured, and decomposed patches on
the leaf. A profuse growth of Peronospora is not to be
distinguished from a slight growth of Oidium, without
the aid of a lens.
If a small piece of the leaf of a turnip infected with
Peronospora parasitica, Pers., corresponding in size and
thinness with the Oidium infected slice already described,
is placed under the microscope and examined, it will be
seen, if enlarged to a scale one-half that of the last, viz.
200 diameters, like the drawing at Fig. 29. The first
point to be especially noticed is, that the spawn which
gives rise to the fruiting threads of the Peronospora mil-
dew is inside the leaf, as shown between the letters A and
B. These letters indicate the upper and lower surface
of the leaf. The spawn threads are stout as compared
with many other mycelia, and have very few septa or
stops ; they are notably furnished with numerous
haustoria, — from haustor, a drawer, — or little suckers, as
shown at C. The suckers attach themselves to the
CH.XV.] MILDEW OF TURKU'S AMD CABBAGES. 81
x-aoo-
FIG. 29.— PUTREFACTIVE MILDEW OF TURNIPS AND CABBAGES.
Peronosp&ra parasitica, Pers. Enlarged 200 diameters.
constituent cells of the leaf, and the mycelium sets up
decomposition in every part of the leaf with which it
comes into contact.
G
82 DISEASES OF FIELD & GARDEN CROPS. [CH.
The upper surface of a turnip leaf is shown at A, and
the lower surface at B, and it will be seen at once that
the spawn threads within the leaf are of too great a
diameter to emerge through the little lancet-shaped
orifices of the stomata. When the fungus threads from
within approach the little openings of transpiration in
their growth outwards, they slightly contract in dia-
meter, form a stop or joint, and then, instead of emerging
through the stomata with a pointed end, they present
a chisel edge to the mouth of the pore, which exactly
suits the shape of the little lanceolate opening. This
mode of emergence is shown, enlarged 1000 diameters,
on the left hand lower illustration of Fig. 30. The stems
of the fruiting threads are therefore not truly cylindrical,
but, when seen in section, present a flattened oval form as
illustrated. As the stem now gradually grows upwards
it usually twists round once upon itself. This twisting
habit is slightly retained by all the numerous branches and
branchlets of the fungus. The upper part of each stem of
Peronospora parasitica, Pers., is very much branched and
rebranched, and each ^ittle branchlet carries a compara-
tively large ovate, almost globular spore, as shown, enlarged
1000 diameters, in Fig. 30. The spores or conidia
usually germinate by bursting at the side, and the pro-
truded vital material or germ tube has the power of
piercing the cuticle of cruciferous plants. Although repre-
sented in the drawing as growing in an upright fashion,
the real growth of every Peronospora is of course down-
wards from the under surface of the leaf. Now, if the
illustrative drawing is turned upside down to present the
Peronospora, in a really natural manner, the resemblance
of the fungus to a minute bunch of grapes is a striking
one. From this resemblance the genus now termed
was for many years known as Botrytis — from
a bunch, in reference to the resemblance to a
bunch of grapes.
No zoospores have been detected in this species, but,
xv.j MILDEW OF TURNIPS AND CABBAGES.
FIG. 30.
Peronospora parasitica, Pers.
Spore and Conidiophore emerging through an organ of transpiration.
Enlarged 1000 diameters.
like the Peronospora of the potato, it produces oospores or
resting-spores ; the resting-spores act as seeds, and carry
on the life of the fungus in a hibernating state through
84 DISEASES OF FIELD & GARDEN CROPS. [CH.
the winter. The resting -spores, which were first de-
tected in turnips in 1849 by Mr. C. Edmund Broome,
M.A., F.L.S., of Batheaston, Bath, were figured by Dr.
Montagne, and named by him (like the similar bodies
found in diseased potatoes) Artotrogus. The resting-
spores or oospores of Peronospora parasitica, Pers., are
often extremely common in rotten turnips and mangel-
wurzels, as found in the fields in autumn. These roots
are often destroyed by a combined attack of the putre-
factive mildew of turnips and the fungus of club -root,
described further on in this work. In order to see the
resting-spores, portions of the brown decayed substance of
the diseased root should be looked over till the oospores
are found. The less ripe examples are smooth outside
or slightly granular, and the more mature specimens are
beautifully echinulate, as illustrated, enlarged 400 dia-
X-40O-
FIG. 31.
Resting-spores or Oospores of Peronospora parasitica, Pers.
Enlarged 400 diameters.
meters, at Fig. 31. Dr. Montagne and Mr. C. E. Broome
first observed this fact in 1849, for on the original draw-
ing in the possession of the Eev. M. J. Berkeley the
smooth form is labelled " sp. jun." and the nodulose form
"matur" In colour the resting-spores are yellowish-
brown. The examples found by Mr. Broome in 1849 ;
those illustrated by Professor de Bary in 1863, Ann. Sc.
Nat,, 4 ser., vol. xx. — again by him in the Beitrage zur
Morphologic und Physiologic der Pilze, 1881, pi. 1, under
the name of Artotrogus hydnosporus, Mont, (see also Gar-
dener's Chronicle, April 26, 1884, p. 544) ; and the speci-
mens shown at Fig. 31, agree in size, character, and colour.
xv.] MILDEW OF TURNIPS AND CABBAGES. 85
The resting-spores of this fungus have also been detected
in the wallflower, Cheiranthus Cheiri, L. ; the Shepherd's-
purse, Oapsella Bursa-pastoris, D.C. ; Gamelina sativa,
Cranz., and in other plants.
Peronospora parasitica, Pers., not only grows on the
wild and all the cultivated varieties of the cabbage and
turnip, Brassica oleracea, L., and B. campestris, L. ; but it
often grows on Whitlow Grass, Draba verna, L., and on
the Shepherd's -purse, Gapsella Bursa-pastoris, D.C., in
company with one of the white-rust fungi named Cystopus
candidus, Lev., described farther on. It also grows on
Garlic Mustard, Alliaria officinalis, D.C. ; Pennycress,
Thlaspi arvense, L. ; Tower mustard, Ardbis perfoliata, L. ;
Coral root, Dentaria bulbifera, L. ; D. heptaphyllos, Clus. ;
Neslia paniculata, Des. ; Hairy Bittercress, Cardamine
hirsuta, L. ; Narrow-leaved Bittercress, G. impatiens, L.,
and other plants.
The many common weeds just mentioned act as nurse
plants for the putrefactive mildew of our turnips and
cabbages. The fungus lives through the winter in a
hibernating state not only in rotten turnip and mangel
roots, but in the decaying remains of such extremely
common weeds as the Shepherd's-purse and other worth-
less plants. It is obvious, then, that it is not only desirable
to burn all fungus-infected turnip and cabbage material,
but as far as possible to keep the fields and hedgerows
clear from the cruciferous weeds just mentioned. It
may be answered that it is impossible to keep down the
weeds and burn the decaying cruciferous rubbish. This
may be partially true, but the moral to be drawn from
the life history of this fungus is, Do not let putrid refuse
and worthless and dangerous weeds interfere more than
is necessary with the healthy growth of food-plants.
CHAPTER XVI.
WHITE-RUST DISEASE OF CABBAGES, ETC.
Cystopus candidus, Lev.
THERE is no more familiar parasite of cruciferous plants
than the fungus of White Rust, Cystopus candidus, Lev.
The generic name is derived from Jcystis, a bladder, and
pous, a foot ; candidus, of course, refers to the white
colour of the fungus ; the name is intended to indicate the
white pustular appearance of the fungus on the attacked
plants. White rust is extremely common on cabbages,
excessively so on the common Shepherd's-purse, Capsella
Bursa-pastoris, D.C., and many other cruciferous weeds
and garden flowers. The appearance of the fungus is
known to every one who has walked in a kitchen garden.
Cabbages and cauliflowers are seen with their leaves and
stems swollen, distorted, and spotted with white streaks and
blotches, as if sprinkled over with whitewash. If typical
examples of the parasite are carefully examined on in-
vaded leaves, it will be noticed that the white splashes
are really somewhat elongated swollen pustules, often
arranged in a concentric or spiral manner, and measur-
ing half an inch or more across. On the leaf stalks and
flower stems the pustules are disposed in a more irregular
manner. The parasite invades every part of the host
plant above ground, sometimes sweeping off every seedling
in the earliest stages of growth, at other times attacking
the flowers, and so stopping the production of seeds. The
fungus, in whatever form it appears, reduces and damages
the produce of the plants attacked. Experienced observers
can detect the presence of white rust long before the
CH.XVI.] WHITE-RUST DISEASE OF CABBAGES, ETC. 87
white pustules are visible, by the swollen and distorted
appearance of the leaves and stems, caused by the presence
of the spawn of the parasite within the plant. As in
Peronospora, the mycelium of Cystopus traverses the host
plant by the intercellular passages. The spawn threads
resemble the mycelium of Peronospora parasitica, Pers.,
in being provided with suckers which become affixed to
the constituent cells within the leaves and stems of the
host. When the white pustules are examined with a
microscope they are found to be not dissimilar in
character although different in colour from the pustules
belonging to Puccinia mixta, FL, already described, or of
the rust fungus of corn, Uredo linearis, Pers., described
further on. Instead, however, of simple red Uredo spores
or compound blackish Puccinia spores being found within
the pustules, chains of almost colourless round or ovate
spores or conidia are seen in the white -rust fungus.
Chains of conidia or spores belonging to Cystopus candidus.,
Lev., are illustrated in different stages of growth, enlarged
400 diameters, at A, Fig. 32. The fungus grows beneath
the epidermis of the plant after the manner of Puccinia
mixta, Fl., already described ; the pustules produced by
the white -rust fungus are, however, very much larger
than the blisters of the Puccinia. The spores or conidia
grow in chains, a fact first pointed out by the Eev. M.
J. Berkeley, in the Journal of the Royal Horticultural
Society, vol. iii., p. 269, 1848. The spores or conidia are
formed in Cystopus in the following manner : — At first
simple clublike growths are produced as at B ; a con-
striction forms towards the apex of the club, which
speedily takes the form of a joint or septum as at C ; in
the process of growth another constriction occurs as at D,
which in turn speedily becomes a septum or joint. As
this process is repeated each club at length supports a
short chain of conidia, each conidium being attached to
the conidia next in order by joints as at DD. When
large numbers of conidia have been produced in this
DISEASES OF FIELD & GARDEN CROPS. [CH.
manner in the disease pustules, the epidermis of the host
plant bursts in an irregular manner, and the conidia are
set free. Each conidium is filled with finely granulated
protoplasm or vital material.
When the conidia approach maturity in damp air or
water, the interior substance of each may be seen divided
into a definite number of portions, generally from five to
eight ; each portion presenting a pentagonal or hexagonal
form bounded by a white line, precisely in the manner of
the conidia of the fungus of the potato disease described
X400
Pro. 32.— WHITE RUST OF CABBAGES.
Cystopus candidus, Lev. Enlarged 400 and 1000 diameters.
farther on in this work. Each of the contained portions
within the conidium is now really a secondary spore, and
the body which was at first a simple conidium has now
become a sporangium or spore case — technically, in this
instance, called a zoosporangium, or case containing zoo-
spores or spores endowed with an animal-like motion.
If we take a perfectly ripe conidium, sporangium, or
xvi. J WHITE-RUST DISEASE OF CABBAGES, ETC. 89
zoosporangium, and enlarge it 1000 diameters, we shall see
it as at E, Fig. 32 ; the former point of attachment is seen at
F, and the sporangium is shown in the act of discharging
its differentiated contents, in the form of zoospores, from
its apex. These secondary spores were at first the poly-
hedric contents of the sporangium ; but as they emerge in
water or on any damp surface the angles become rounded,
and they are at last expelled as minute ovoid bodies as
illustrated at G. At first these small secondary spores or
zoospores remain immovable at the mouth of the burst
sporangium ; soon, however, they begin to slightly oscillate,
and two excessively attenuated hairlike cilia are de-
veloped from beneath as at H. At a special moment the
foremost cilium is distended in a straight line as shown,
whilst the hindermost cilium at the same time suddenly
quivers, and the zoospore sails away over any moist
surface, as if endowed with animal life. Each zoospore
exhibits within one or more lustrous, perhaps contractile,
vacuoles.
The phenomena just described can only be seen when a
zoosporangium of -the white-rust fungus, has been placed
in water upon a glass slide, and viewed under a cover-
glass with a high power of the microscope. In dry air
no differentiation of the contents of the conidium takes
place. It is certain that rain, dew, or moisture of some
sort is essential for the bursting of the sporangia and the
expulsion of the zoospores. The bursting, as seen in
water, under the microscope takes place in an hour or
two after immersion ; the conidia retain the power of pro-
ducing zoospores for about a month. The zoospores are
able to swim about for several hours ; their cilia then
vanish, the zoospore retakes a globular tailless form,
bursts as at J, produces a germ tube, and this germ tube
is then a spawn thread of white rust capable of pro-
ducing a new series of clubs capped with zoospore-
bearing sporangia.
It is obvious from the above description that the white-
90 DISEASES OF FIELD & GARDEN CROPS. [OH.
rust fungus is carried from one leaf to another, from one
plant to its neighbouring plant, and from weeds to food-
plants, in damp, rainy, or misty weather, by the microscopic
zoospores sailing about over moist surfaces. There can
be no doubt that they are also carried about in damp air,
in currents of wind, and that birds, insects, and other
animals help to carry the living conidia and zoospores
from place to place.
As a rule conidia, zoospores, germ tubes, and fungus
spawn are very liable to perish ; too much dryness, a
superabundance of moisture or frost, will quickly destroy
them.
Three questions now present themselves to us — How
does the white-rust fungus tide over the winter ? Where
is it hidden ? How does it suddenly reappear in the
spring ? In the case of many plant diseases, as in the
surface mildew of turnips, Oidium Balsamii, Mont., already
described, no one is able to answer such questions ; but
with the white-rust fungus and several of its allies the
knowledge has been obtained, and a satisfactory answer
can be given. It has been already stated that the spawn
or mycelium of Cystopus grows within the leaves and stems
and burrows amongst the intercellular spaces of the host
plant. It not only bears the chains of spores already de-
scribed, which, when ripe, are blown away by the wind,
but it carries other bodies within the substance of the leaf.
These latter organs roughly answer to the pistils and anthers
of flowering plants. The first bodies are female, and are
termed oogonia ; these are large globular cells in which
the female reproductive bodies, or oospheres, or sometimes
zoospores, are formed. They generally grow on terminal
branches of the mycelium ; sometimes they are sessile
or nearly so, or they may be intercalated in the my-
celium itself. An oogonium is illustrated, enlarged 400
diameters, at A, Fig. 33. The oosphere, filled with
granular protoplasm or vital formative material, is seen
within. Other organs borne on the mycelium are male,
xvi.] WHITE-RUST DISEASE OF CABBAGES, ETC. 91
and termed, in reference to their nature, antheridia, or
organs answering to the anthers of flowering plants. In
the course of growth the antheridium comes in contact
with the oogonium as at B, and projects a fine "beak
through its wall, till it pierces the oosphere within as at
C. This is the act of fertilisation answering to the dis-
charge of pollen on to the stigma in flowering plants.
In the same way as an ovule becomes a seed after
FIG. 33. — WHITE RUST OF CABBAGES.
Oogonium with Antheridium and Resting-spores, or Oospores of Cystopits
candidus, Lev. Enlarged 400 diameters.
fertilisation in flowering plants, the oosphere becomes an
oospore or egglike spore, after the contact of the antheridium
with the oosphere. The now fertile oospore within the
oogonium grows and matures itself whilst still within the
supporting leaf or stem for many months, generally for
the greater part of a year, and it does not become perfectly
ripe till the host plant has decayed. Although the
92 DISEASES OF FIELD & GARDEN CROPS. [CH.
cabbage leaf which carried the Gystopus may be dead and
putrid, or reduced to tinder by drought and frost, the
oospores or resting-spores remain alive and uninjured, in
a dormant state. They change in colour and form, from
almost colourless smooth spheres to amber-coloured, warted,
globular bodies, as illustrated at D. They are best seen
in the putrid remains of plants which have been destroyed
by the white-rust fungus. In good material the amber-
coloured oospores will be seen closely packed together in
the decayed leaf or stem in enormous numbers. The best
plan for obtaining resting-spores is to collect leaves in-
fested with white rust and allow them to decay upon a
garden bed ; after the diseased leaves have perished the
oospores will be found during the winter or the following
spring in the decayed fragments of foliage.
The oospores germinate on the ground during wet
weather in the spring ; but the germination may be
easily observed in water under the microscope. After a
few ripe resting-spores have been placed in a drop of
water they will speedily burst, either at once, or in a day
or two, according to the state of their maturity. They
germinate by bursting, as illustrated at Fig. 33, E ; a
transparent inner membrane is protruded, and the con-
tained protoplasm, which at first is differentiated into
numerous polyhedric portions, at length resolves itself into
a large number of oval zoospores as at F. Soon the trans-
parent investing membrane is ruptured, and the zoospores
sail out as at G, thus repeating, after from six to ten months'
rest, the phenomenon described as belonging to the chains
of conidia illustrated in Fig. 32. The zoospores produced
Ly the conidia are precisely the same in size and habit with
those produced by the oospores ; in both instances they
germinate in the same manner after swimming about for
three or four hours in water. The difference in size of the
zoospores shown in Fig. 32 and Fig. 33 is owing to the
fact of the former being enlarged 1000 diameters, whilst
the latter illustration is only enlarged 400 diameters.
xvi.] WHITE-RUST DISEASE OF CABBAGES, ETC. 93
Cruciferous plants in the spring owe their infection
with white rust to the zoospores germinating upon the
seed-leaves. No one, of course, has seen such extremely
minute objects as zoospores with the unaided eye, so no
one has ever seen them naturally transferred from the
germinating oospores on the wet ground, to the young
seed-leaves of cabbages, cauliflowers, and other cruciferous
plants. It is said that the zoospores cannot effectually
, germinate and form mycelium upon and in leaves and
stems of cruciferous plants unless the latter are very
young. But as cruciferous weeds infected with Cystopus
are extremely common, the oospores must occur in pro-
fusion in all districts every spring. No doubt the little
motile zoospores are carried through moist air by currents
of wind, and distributed in every direction throughout
the country.
Alternation of crops must tend to diminish white rust.
Cabbages, cauliflowers, etc., should not be grown for two
years in succession where white rust has prevailed.
Cruciferous weeds should be gathered together and burnt,
especially when they exhibit the well-known white
sprinkling of the white-rust fungus. No cabbage, cauli-
flower, turnip, or mangel refuse should be allowed to
remain in a decaying state throughout the winter in the
fields, for in those positions not only the white-rust
fungus, but the putrefactive mildew of the cabbage tribe
and the fungus of club-root hibernate. Clean and intelli-
gent farming will greatly reduce the attacks of these two,
as well as of many other pests.
CHAPTER XVII.
CLUB-ROOT OF TURNIPS, CABBAGES, MANGELS, AND
ALLIED PLANTS.
Plasmodiophora Brassicce, Wor.
THE disease of turnips, cabbages, and allied plants, known
in some districts by the popular name of club-root, is
recognised in other places as anbury and finger and toe.
On the continent the disease is popularly known as hernia
or rupture.
Until the last six or seven years no one knew the cause
of club-root, but in 1876, after three years' constant atten-
tion, M. Woronin, a Russian botanist, as completely
explained the nature of club-root in turnip and cabbages,
as the Rev. M. J. Berkeley expounded the nature of the
murrain of potatoes in 1846.
The observations made by M. "Woronin, which have
several times been confirmed by others as well as ourselves,
seem to place the fact beyond all doubt that clubbing is
caused by a fungus named, by M. Woronin, Plasmodiophora
Brassicce. Plasmodiophora means a bearer or carrier of a
plasmodium, and a plasmodimn is an Amoeba-like mass of
protoplasm or vital formative material of changeable form ;
Brassicce, of course, means that the fungus is peculiar to
the turnip and cabbage class. The family to which the
fungus belongs is a remarkable one, and is known as the
Myxomycetes or family of slime-fungi. These fungi have
appeared so animal -like to some observers that, by a
misinterpretation of analogies, an attempt has been made
to transfer them to the Protozoic division of the animal
kingdom. With the same idea in view they have been
CH.XVII.] CLUB-EOOT OF TURNIPS, ETC. 95
termed by Professor A. De Bary Mycetozoa, or fungus-like
animals. No fungologists of repute, however, and very
few zoologists, hold either of these views at the present day.
When Professor De Bary termed these fungi Mycetozoa,
little or nothing had been learned of the production of
zoospores in fungi, a phenomenon now so well known in
Gystopus, Peronospora, and other genera.
The Myxomycetes are especially remarkable in the
fact that they do not form cells, cellwalls, tissues, or
mycelium, during the period of vegetation, but their
protoplasm remains during that time free, and collected
into small masses of various and changeable forms. At a
certain definite advanced period of growth the vital
material of a Myxomycete breaks up into small portions,
and these portions at length surround themselves with a
cellwall, and become either fruits, sporangia, or spores,
and in this condition the fungus remains at rest during
a certain definite period. If the spores are kept dry they
will retain their vitality for several years. After a period
of hibernation the sporangia sometimes coalesce, and the
spores germinate by the cellwall cracking, and the vital ma-
terial exuding as a small round or irregularly-shaped mass ;
this exuded mass speedily becomes furnished with one or
two highly-attenuated tails, vibrating hairs, or cilia, and
with the aid of these tails the little exuded masses are
enabled to creep about over any moist surface in an
Amoeba-like fashion. The exuded masses are capable of
multiplication by division, or (generally after a few days)
they will unite with each other, and so form a homogene-
ous mass of protoplasm of larger size, which mass also
possesses an Amoeba-like movement. This homogeneous
mass of combined Amoeba-like material ejected from the
spores is termed a plasmodium. The plasmodium is now
capable not only of coalescing with other neighbouring
plasmodia, but also of absorbing other Amoeba-like spore
contents. A plasmodium possesses the power of creeping
about by extending armlike processes from its margin,
96 DISEASES OF FIELD & GARDEN CROPS. [CH.
and by the vital material from the mass repeatedly press-
ing into the arms or processes. The plasmodium is enclosed
by a dense hyaline layer, and this in turn is surrounded
by a thin coat of mucilage, which mucilage is sometimes
left behind by the progressing plasmodium like a trail of
slime from a slug. Although our fields are at all times
saturated and traversed by the spore contents or plasmodia
of this destructive fungus, yet Plasmodiophora has not
hitherto appeared in our printed lists or handbooks.
The structure and habits of the members of the whole
family of the Myxomycetes, with its numerous genera, are
too involved and different from each other for any further
general description in this place.
Clubbing commonly commences at an early period in
the life of the seedling turnip, cabbage, or other crucifer-
ous plant. If we take a young seedling turnip, — one
which shows by its flagging foliage and dwindled growth
that it is out of health, — and examine the root, we shall
probably see it, if attacked by the club-root fungus, some-
thing like the illustration at Fig. 34, which is engraved
one-half the natural size. The example illustrated repre-
sents a seedling turnip two months old. The rootlets
will be seen to be swollen with spindle-shaped swellings,
generally with a smooth and flowing outline ; and this
peculiar smooth spindle form of the clubs distinguishes
true clubbing from all abrupt tuberous swellings and
excrescences, sometimes natural, at other times abnormal,
as when caused by the insects and larvaa so common on
cruciferous plants. Every swelling in true club-root is
not necessarily perfectly smooth or truly fusiform or
spindle-shaped, but in the majority of instances, and
especially in an early state of growth, this distinguishing
mark holds good and is characteristic. Some entomolo-
gists have ascribed the origin of club-root to the attacks
of Aphides ; but Mr. G. B. Buckton, F.R.S., in his "Mono-
graph of the British Aphides," published by the Ray
Society, rejects this idea, but thinks some clubbing may
xvii.] CLUB-ROOT OF TURNIPS, CABBAGES, ETC. 97
be due to insect punctures on the tap-root when the plant
is young. But excrescences caused by insect punctures
are quite distinct from true clubbing. Curtis, in his
Farm Insects, has rejected the idea of Aphides being the
cause of clubbing.
To see the nature of the fungus of club-root, one of the
smaller spindle-shaped swellings must be cut in two, as
on the line A, B, Fig. 34, and from one of the exposed
FIG. 34.— CLUB-ROOT DISEASE OF TURNIPS, ETC.
Root of young Turnip with Clubs.
One-half natural size.
surfaces a thin slice must be cut. If this is done in July,
and the slice is viewed as an opaque object and magni-
fied 10 diameters, it will be seen, as in Fig. 35, faintly and
curiously mottled and clouded. If an extremely thin
atom is now cut off and viewed as a transparent object
with a power of 200 diameters, it will be seen as at Fig.
36. The cause of the mottling will now be seen to be
due to the presence of a yellowish stringy slime or plasma,
H
98 DISEASES OF FIELD & GARDEN CROPS. [cir.
X-10
FIG. 35. — Cr.fB-RoOT DISEASE OF TURNIPS.
Section through a small Club. Enlarged 10 diameters.
FIG. 36.— CLUB-ROOT DISEASE OF TURNIPS.
Section through Cells of Turnip-root, showing the Plasma of Plasmodio-
phora Bfassicce, Wor. Enlarged 200 diameters.
xvii.] CLUB-ROOT OF TURNIPS, CABBAGES, ETC. 99
sometimes wholly filling certain cells, in other instances
appearing as strings of slimy protoplasm drawn across
from one side to the other. No true mycelial tubes can
be seen, and none of the familiar cells so common in most
fungi. One fact will strike the observer at once, and that
is, the affected cells will be noticed as much larger in size
than the ordinary cells of the rootlet — in many instances
enormously larger. This distention of the cells is a
common result of the attacks of parasitic fungi on leaves
and roots, and one can understand at once that if each
constituent cell of the infant turnip -root or rootlet is
distended to ten or one hundred times its normal size, a
clublike growth must result. It will be noted too that
the cells, though enormously distended, have not burst.
If a club is examined later in the season — say in
October — a very different appearance is presented, and
the change we then see has been gradually going on
during the autumn months. The protoplasm of the
summer has, by the late autumn, broken up into innu-
merable minute spherical portions, and the stringy, slimy
mycelium has been replaced by millions of excessively
minute spherical spores. These spores may now be dis-
tinctly seen to possess a cell wall. The cells of the turnip
are now, even more distended than before, and in many
instances they will be seen closely packed with the greatest
regularity by vast cohorts of the Plasmodiophora spores.
It will still be seen that most of the distended cells of
the turnip remain intact, and only a few are ruptured.
A little pressure of the covering glass of the microscopic
slide will, however, speedily break some of the cell walls,
and the spores will pour in enormous quantities through
the breach into the surrounding film of water. This
condition of the disease is illustrated at Fig. 37, enlarged
200 diameters, where the spores are seen pouring out
through the breaches in the cell walls. The spores are
farther enlarged to 1000 diameters at Fig. 38, so that
their size may be compared with other spores drawn to
100 DISEASES OF FIELD & GARDEN CROPS. [CH.
the same scale in the other illustrations given in this
work.
The three larger spherical brown bodies seen in Fig.
37, and the single example in Fig. 38, are resting-spores
of the putrefactive fungus of cabbages and turnips, named
Peronospora parasitica, Pers., and sometimes seen in great
•K-200 •
FIG. 37. — CLUB-ROOT DISEASE OF TURNIPS.
Spores of Plasmodiophora Brassicce, Wor., seen within the Cells of Turnip-
root. Enlarged 200 diameters.
abundance in turnip plants reduced to putridity by the
combined attacks of the Peronospora and the Plasmodio-
phora. These larger bodies are the second species of
Artotrogus (not A. hydnosporus] of Montagne.
The Plasmodiophora spores remain uninjured and in a
xvii.] CLUB-ROOT OF TURNIPS, CABBAGES, ETC. 101
resting state in affected turnips all through the winter ;
and in the spring, if another transparent slice is taken
from a small club formed during the previous year, the
X-IOOO
FIG. 38.— CLUB- ROOT DISEASE OF TURNIPS.
Small spores of PlasmodiopTiora, Brassicw, "Wor., and single Resting-spore
of Peronospora parasitica, Pers. Enlarged 1000 diameters.
spores will be found perfectly ripe and ready for germi-
nation. This germination takes place as illustrated in
FIG. 39.— CLUB-ROOT DISEASE OF TURNIPS.
Spores of PlasmodiopJiora Brassicce, Wor., germinating and producing
Amoeba-like zoospores.
the six spores, enlarged 1000 diameters, and numbered
from 1 to 6 on Fig. 39. The cell wall of the spore
102 DISEASES OF FIELD & GARDEN CROPS. [CH.
cracks, and the contained protoplasm creeps out, as illus-
trated in the successive figures. First a small protuberance
is seen, then a slightly branched arm, next the main
growth is attenuated, then the attenuation proceeds to an
almost invisible hairlike tail or flagellum ; at last the
minute speck of protoplasm flies out of the - spore wall,
often whirling the empty case to some little distance.
Each atom of protoplasm which has emerged from the
spore is now a free zoospore, or animal-like spore — capable,
aided by its hairlike tail, of creeping or sailing along, or
whirling round in any film of moisture, as illustrated in
the figures numbered from 7 to 12.
These zoospores, like all other zoospores, do not long
remain in the zoospore condition. In most fungi zoo-
spores speedily burst and protrude a thread of mycelium ;
but in Plasmodiophora the zoospores quickly coalesce, and
when a few have conjoined they form the growth already
described as a plasmodium. The viscid plasmodium formed
by one set of conjoined zoospores speedily comes in con-
tact with other and similarly-formed plasmodia, and so
larger examples are formed. These examples, large and
small, are, when in a state of nature, washed out of de-
caying club-root material into the ground by the spring
rains. There, on and in the moist ground, they are able,
by pushing out arms and prolongations, and by con-
tinually propelling their contained vital material into
these extensions, to move about in a sluggish Amoeba-
like fashion.
All practical agriculturists will now see that when
club - root refuse is left in the fields, or thrown on to
dung-heaps, and then distributed over the ground, the
most certain method is taken for propagating club -root
disease, for out of this decayed material innumerable
motile plasmodia will be washed into the ground. When
the seeds of turnips and cabbages grow and extend their
rootlets, the rootlets naturally come into contact with the
viscid watery plasmodia in the ground, and these plas-
XVIL] CLUB-ROOT OF TURNIPS, CABBAGES, ETC. 103
modia are absorbed into the young turnip plants by the
rootlets. It cannot be objected that a plasmodium is too
large to find entrance to a plant by the rootlets, for plas-
modia are capable of existing in a state of threadlike fine-
ness and watery attenuation beyond conception.
When once in the rootlets, the plasmodia are in the
position that best suits them, and in that position they
act as true parasites in the host plant, and by their
growth excite disease, unnatural distention of the cells,
and " club-root."
The proofs that old club -roots, with their contained
ripe spores, can really produce "club -root" disease in
growing turnips have many times been given of late
years, and the experiments have been several times re-
peated by ourselves. They amount shortly to this. If in
spring-time turnip seed is planted in pots in virgin mould,
the seedlings will come up unclubbed ; but if exactly
similar seeds are planted in earth in which old chopped-
up clubs have been incorporated, the seedlings will nearly
all be at an early period of growth fatally clubbed.
For the prevention of clubbing, an alternation of crops
for two or three years may reduce the disease, for, as far
as is at present known, Plasmodiophora Brassicce, Wor.,
is confined to cruciferous plants. As the spores of the
fungus can live for more than a year in dry material,
more than one season should elapse before turnips or
cabbages are again planted in tainted fields. As char-
lock is often badly clubbed, this, with other worthless
cruciferous weeds, should not be allowed, more than is
possible, to choke the hedge sides of fields under cultiva-
tion with cabbages, turnips, and mangels.
Beyond all other things, it is necessary that old club-
root should not be allowed to remain on the ground where
turnips or cabbages are to be grown. All the diseased ma-
terial should be gathered into a heap, and, if possible, burnt.
Prof. Jamieson, in the last annual report of the Sussex
Association for the Improvement of Agriculture, advised
104 DISEASES OF FIELD & GARDEN CROPS. [CH.XVII.
farmers not to use manures containing sulphur and chlo-
rine elements ; the one given usually as sulphuric acid and
sulphate in dissolved or soluble manures, the others given
in chloride, or muriate of potash, and in common salt.
Sulphuric acid is a characteristic ingredient in nearly all
vitriolated or phosphatic manures. Prof. Jamieson states
that the only crops which are uninjured by these manures
are the cereals, and that they should never be used for
other crops unless the soil is unusually black ; he con-
siders that club-root and sickness in turnips is aided by
sulphur, and that chlorine is injurious both to Swedes
and peas. Prof. Jamieson is of opinion that the sulphur
in the manure (in whatever form) accelerates the elabor-
ation of the delectable sulphurous material, in which the
dormant spore finds abundant sustenance. At the same
time, by the readily available form of the mineral food, a
flush of premature growth pervades the cultivated plant,
and consequent weakness. Simultaneously the fungoid
enemy, at the expense of the higher plant, increases in
myriads, war is waged, in which the assailing foe — the
fungus — is never subdued, but may have either a com-
plete victory in the death of the higher plant, or only
partial victory, resulting in a more or less clubbed root,
and a more or less normal bulb above.
In August 1883 experiments at Hassocks, in Sussex, a
conspicuously unhealthy appearance is said to have been
observable in all the turnips of the superphosphated plot ;
although it had been stated that the club-root disease had
previously been unknown in Sussex.
In October the plants were taken up from all the plots,
with the following result : —
MQTI, loAd Total diseased and
Manures used. knied Plants>
Ground coprolite . . 11.
Ground bone-ash
Superphosphate
No phosphate .
Steamed bone-flour .
Coprolite and steamed bone-flour
16.
51.
45.
16.
3.
CHAPTER XVIII.
EAR-COCKLE, PURPLES OR PEPPERCORN IN WHEAT,
OATS, AND RYE.
Tylenchus tritici, Bast.
FOR a clear comprehension of the disease of wheat, oats,
and rye, named ear-cockle, purples, or peppercorn, it is
necessary that the nature of an ear of wheat should be
well understood. We will therefore briefly describe the
structure of the inflorescence of wheat, so that the pecu-
liarities of ear-cockle may be made clear.
An ear of wheat .is technically termed a spike, and
the spike consists of a rachis (literally a backbone) or zig-
zag stem, on which are placed numerous little clusters of
grains with their chaffy scales. Each cluster is termed a
spikelet, and as a single cluster or spikelet taken from
any part of a spike is generally the same in arrangement
as all the other clusters on the spike, we will remove one
cluster or spikelet for careful observation.
At Fig. 40 is illustrated a spikelet of wheat enlarged
two diameters. It will be seen that the whole growth is
enclosed between two outer sheaths or bractlike scales, seen
at AA ; these are termed the two outer glumes — glume
merely meaning " chaff." In wheat it is common to see
the two outer glumes enclosing five other growths. One
of these is an aborted growth seen at B, and the two
clusters on each side at C, D, E and F, each include two
bractlike scales, one a flowering glume, and the other
a pale. Each of the two inner growths encloses a pistil,
three stamens, and two minute scales. The lowermost and
outer bract of these two is termed the flowering glume ;
106
DISEASES OF FIELD & GARDEN CROPS. [CH.
this has a central mid-rib, and the inner one is termed
the pale (meaning also chaff), with two distinct side ribs
and no mid rib.
If the parts of a ripe spikelet are compacted together so
' I 'X 2
FIG. 40.
Spikelet of Wheat. Twice the size of nature.
that they will not break away under the knife, and a
horizontal section is made through all the fruits or grains
with their enclosing chaffy scales, the parts of the spikelet
will be seen as in the section at Fig. 41, enlarged four
D C
FIG. 41.
Horizontal Section through a Spikelet of Wheat, showing the grains,
scales, etc. Enlarged 4 diameters.
diameters. We now see in horizontal section the central
abortive growth at A, and the four normal grains, 1, 2,
3, and 4, each enclosed within a pale, with its two side
xviii.] EAR-COCKLE IN" WHEAT, OATS, & RYE. 107
ribs, as at B, and a flowering glume with its central mid-
rib C. The flowering glume is often capped with a long
thread beard or awn, familiar in barley and oats. The
whole growth is embraced by the two outer glumes at
DD.
We will now closely examine one of the four groups,
with its two enclosing scales, the flowering glume and
pale. Arranged round each pistil (or in the ripe ear — the
grain) are three stamens, E, F, and G (Fig. 41), and two
beautiful fimbriated transparent, membranous, almost
microscopic scales at H, J; the three delicate drooping
stamens and two little scales all grow at the base of the
pistil, carpel, or grain. The two little transparent scales
are usually admitted to represent the perianth of more
perfect flowers. The different parts of a grass spikelet
possess considerable botanical interest, and the questions
are by no means settled as to the exact morphological
significance of the glumes, palae, and scales, and their mode
of attachment. The questions are, however, beyond our
province here, and need not be discussed in detail. It
is remarkable that the two little transparent scales at
the base of the pistil are persistent, and in this they differ
from the fugitive feathery stigmas and stamens.
As great attention has been directed by botanists to
the two minute scales, technically termed lodicules, grow-
ing at the base of the ovary, and their connection with
ear-cockle, a single grain of wheat detached from the
spikelet, but still enclosed within its pale, is shown
enlarged to five diameters at Fig. 42. The spectator
is supposed to be looking towards the interior of the
pale, and the furrow or cleft of the seed is away from the
spectator and towards the pale. Pendulous from the base
of the grain is a withered stamen. On either side of the
point of insertion of the stamen, and at the base of the
grain, the two minute lodicules are seen at A and B.
The wrinkled part of the base of the grain at G is the
spot whence the plumule and radicle of the young wheat-
108 DISEASES OF FIELD & GARDEN CROPS. [CH.
plant will emerge on the germination of the seed. A
single scale or lodicule is enlarged to fifty diameters at
Fig. 43 to clearly show its form. In text-books these
scales are usually described as fringed at the top, but in
nature they are also usually fringed down both sides ; and
a similar fringe belongs to both glume and pale towards
X-5 X<SO
FIG. 42.— Grain of Wheat FIG 43.
enclosed in its Pale. Lodicule from base of Wheat grain.
Enlarged 5 diameters. Enlarged 50 diameters.
their base, and a not dissimilar one to the grain at its
apex.
We will now leave the normal healthy spikelet, and
examine a diseased one. When a wheat plant is affected
with ear-cockle, the spikelets present a much thinner,
looser, and more open appearance than the healthy ones.
This appearance is shown at Fig. 44, enlarged two dia-
meters ; here the large normal grains are replaced by four
small peppercorns, shown solid black in the illustration.
xvni. ] EAR-COCKLE IN WHEAT, OATS, & RYE. 109
The small purplish -black grain -like growths found
within the flowering glume and pales in the disease known
as ear-cockle, are galls caused by the attack of a nematoid
or thread -worm, named, by Dr. H. C. Bastian, Tylenchus
tritici. Tylenchus is a compound word derived from the
Greek, and indicates the knoblike growth of the galls and
-the so-called "spear" of the Nematode — that is, the mus-
cular bag forming the back part of the mouth; tritici, of
course, refers to the classical name of the wheat plant.
The popular names ear-cockle, purples, and peppercorn
have reference to the form, size, and colour of the little
galls ; these galls are roughly comparable, on a small
X 2
FIG. 44.
Spikelet of Wheat, the grains replaced by the galls of Ear-Cockle.
Twice the size of nature.
scale, with the familiar galls found on the leaves of the
oak and other plants. The Nematode itself is a close
ally of the well-known " eels " of stale paste and vinegar.
In a Nematode-infested wheat spikelet, such as the one
illustrated at Fig. 44, the little blackish galls or pepper-
corns can generally be seen as illustrated between the ill-
grown glumes and pales. Four of these galls are shown
free at Fig. 45, A, B, 0, and D, enlarged five diameters.
These galls are commonly two bodies conjoined, seldom a
single body, and in rare instances three conjoined bodies,
always within the pale and flowering glume. For the
reason that the galls are commonly two conjoined bodies
110 DISEASES OF FIELD & GARDEX CROPS. [GH.
these growths have been associated by Mr. William
Carruthers, F.R.S., in the Journal of the Royal Agri-
cultural Society, vol. xviii., 1882, with the two minute
transparent scales or lodicules belonging to the base of
the pistil. In some instances these galls are, however,
single growths, with one or more furrows, and the ex-
ample illustrated at D shows the two persistent lodicules
present at its base, proving that the two lodicules which
form the perianth of the flower are not invariably the
organs which are replaced by the galls. As this particu-
lar gall-growth at D is a double one, it probably repre-
sents two of the three stamens. A gall of one cell
X5-
FIG. 45.
Galls of Ear-Cockle from a Wheat spikelet.
Enlarged 5 diameters.
represents the pistil, and galls with three cells represent
the three stamens. When a single example of pepper-
corn only is produced within the glume and pale, it agrees
well with the single central oblong carpel, with its downy
top. This view confirms Devaine's observation that the
gall is formed from any of the growths belonging to the
central part of the flower. On one occasion Devaine
detected a gall growing from one of the leaves of a wheat
plant.
The galls originate at a very early period of the de-
velopment of the flowers of wheat, at a time when the
different parts which are to compose the flower are repre-
xviii.] EAR-COCKLE IN WHEAT, OATS, & RYE. Ill
sented by mere minute swellings on the little lateral axis
destined to produce the spikelet.
The small worm, we suppose, attacks these swellings —
either the one belonging to the pistil, the three belonging
to the stamens, or the two belonging to the scales. The
parts attacked may, however, belong in part to more than
one series, — as one scale and one stamen, or the pistil and
one scale, etc. When the puncture is made, an unusual
flow of sap to the injured place — possibly a natural attempt
to repair the injury — is the result, an extremely common
phenomenon in plant injuries. This flow of sap causes
the first cells of the monstrous gall-growth to appear :
the abnormal growth is rapid in development, and in a
short time the assailing Nematode or Nematodes are en-
closed within an abnormally-grown cell wall. A section
of a mature gall is shown at Fig. 45, C, and a section
farther enlarged to forty diameters is shown at Fig. 46,
illustrating part of the wall and part of the enclosed
colony of Nematodes. Both sections show the compara-
tively thick nature of the wall of the gall, and the large
number of thick obscurely hexagonal cells of which it is
built up. It is remarkable that the fungus of corn mildew
and other fungi peculiar to corn have been seen growing
upon these galls. The fact probably shows how completely
the substance of the galls agrees in nature with the sub-
stance of the wheat plant from which they are derived.
When a young gall is cut in two and its interior ex-
amined, it is found filled with a cottony mass, which, on
enlargement with the microscope, becomes revealed as a
mass of semi-transparent nematoid worms of all sizes,
from mature individuals one -seventh of an inch long,
through semi-mature and infant individuals, to transparent
eggs, in which the little Nematodes may be seen coiled up.
A group of these worms, in all stages of growth, may be
seen at A, Fig. 46, drawn to the same scale, i.e. enlarged
forty diameters, as the adjoining wall of the gall at B.
By the time the gall is quite mature the mother worms
112 DISEASES OF FIELD & GARDEX CROPS. [CH.
have died, and the eggs have burst and produced
young.
In voL xxv. of the Transactions of the Linnean Society
Dr. H. C. Bastian has described several other species of
Nematode found on and in wheat, oats, and other grasses.
Dr. Bastian specially adverts to the tenacity of life belong-
ing to the species found under the genus Tylenchus, and
he attributes this vital tenacity in part to the structure
of the integument of the animals. This integument is of
such a nature that it enables the Nematodes to resist
FIG. 46.
Fragment of the wall of an Ear-Cockle gall, with part of a colony of
Nematoid worms, Tylenchus tritici, Bast.
Enlarged 40 diameters.
dessication, and prevents the evaporation of moisture
through their tissues. Whilst some Nematodes immedi-
ately shrivel up when immersed in gelatine, Dr. Bastian
has found that members of the genus Tylenchus will move
about in gelatine from fifteen to twenty minutes. The
power of remaining in a dormant deathlike state for a
long series of years Dr. Bastian attributes to some in-
herent peculiarities of the animals' tissues beyond the
reach of detection by optical instruments of even the
highest power. The same author say it is an estab-
lished fact that Tylenchus tritici, Bast., is capable of
xvrii.] EAR-COCKLE IN WHEAT, OATS, & RYE. 113
resuming activity after remaining dormant for twenty-
seven years.
Various observers have artificially infected wheat with
the Tylenchus by placing the living Nematodes taken from
a gall in the furrow or cleft of the grain, and then planting
the infected seeds in the soil in the usual manner. Or galls
may be planted in close contiguity with healthy grain ;
after a brief time the Nematodes will work their way
through the wall of the softened and decaying galls, and
come naturally in contact with the young leaves sprouting
from the healthy seed. The Nematodes then insert them-
selves between the sheaths of the leaves, gradually working
their way round till they come to the innermost, where
they remain till the rudiments of the future ear begin to
form. Several grasses, in addition to wheat, oats, and
rye, are assailed by Nematodes, notably Festuca elatior, L.,
maize, and different species of bent grass, Agrostis.
Other Nematodes attack cucumbers, melons, carnations,
and many different plants belonging to our greenhouses
and flower and kitchen-gardens. These have never been
scientifically described by zoologists, or even named.
Sometimes the Nematodes attack the roots, especially the
rootlets, and they cause little nutlike swellings to appear.
Sometimes the stem, and in other instances the leaves, are
made the point of attack, and the Nematodes cause dead
pallid patches to appear. On cutting a slice from a pallid
spot, or a slice from a nodule belonging to a rootlet, the
Nematodes and their eggs are almost invariably met with.
The worms are coiled in various ways within the eggs, and
after a definite period the eggs burst, and the young
thread-worms emerge.
As it has been clearly proved that ear-cockle can be
produced by planting the galls containing the Nematodes
with sound grain, the greatest care should be taken in
separating the galls from the seed-wheat. This should
be a very easy matter, as the galls are black in colour,
whilst the grain is yellow, and the galls are only one-half
I
114 DISEASES OF FIELD & GARDEN CROPS. [CH.XVIII.
the size of the grains of wheat, making the process of
sifting an easy one. Of course all galls should be care-
fully gathered together and burnt, for under any circum-
stances such growths and their contents are unpleasant
objects to be ground with corn as food. As the little
animals inside the galls can live, under favourable circum-
stances, in a deathlike state for more than a quarter of
a century, galls should never be stored with corn or
planted.
CHAPTEE XIX.
CLOVER DODDER.
Cuscuta Trifolii, Bab.
CLOVER DODDER is such a familiar, and, as some ob-
servers say, increasing pest in our fields, that any detailed
description of its superficial appearance and habits is
unnecessary. Clover dodder is probably perfectly familiar
to every observant person who has walked through clover
fields.
All the dodders, — and there are some forty or fifty
species — belong to one genus of parasitic plants termed
Cuscuta, a name said to be derived from Chassuth, the
Arabic name for dodder plants. The Kadytas of Theo-
phrastus and the Cassytas of Pliny are believed to be
dodder. These names, as well as the Arabic name, signify
to hold fast, to stitch, and to oppress. The popular name,
dodder, is an English form of the Dutch and German names
Dodern, Touteren, and Todern. Dodd signifies a bunch,
and dot, a tangled thread. Trifolii indicates that the
plant now under description invades clovers. The dodders
are commonly termed scald-weeds, hell-weeds, or strangle-
weeds, and in some districts devil's -guts ; the popular
names indicate the strong hatred rustics bear towards
these weeds.
Cuscutas are closely allied to the Convolvuli of our
gardens, and some botanists place them in the same
natural order with the convolvulus ; others relegate them
to a natural order by themselves, named Cuscutece.
Part of a plant of Cuscuta Trifolii, Bab., growing para-
sitically on clover, is shown at Fig. 47, twice the size of
116 DISEASES OF FIELD & GARDEN CROPS. [OH.
nature. Most of the characters of the parasite that are
visible to the unaided eye are given in this sketch.
Dodders are plants with yellowish or reddish leafless
threadlike stems, the leaves being represented by a few
small transparent scales. The small, usually pinkish,
bell-shaped, sometimes sweet-scented flowers, as in clover
dodder, are collected in little closely -packed heads or
FIG. 47.— CLOVER DODDER.
Cuscuta Trifolii, Bab., growing on Clover.
Twice the size of nature.
clusters, as shown at A, B, Fig. 47. Each floral perianth
is usually divided into four or five segments. The flowers
are commonly succeeded (but not in the case of clover
dodder in this country) by four small seeds. The thread-
like stems are furnished with numerous very small
suckers, as at C, D, with which the parasite attaches itself
to its host.
Dodders grow in all hot and temperate regions, and
xix.] CLOVER DODDER. 117
they fix upon a great variety of plants in addition to
field clovers and lucerne. The best known of these host
plants are flax, thyme, broom, heath, furze, cabbages,
nettles, hops, cranberry, rock-rose, centaury, scabious
grass, bracken, yellow -rattle, eyebright, bastard toad-flax,
yellow bedstraw, camomile, sow thistles, tomatoes, and
even the vine.
It is extremely common to find seeds of dodder amongst
impure clover seeds imported from the Continent. In
some instances it is easy to sift dodder away from the
larger-seeded varieties of clover, and we know that most
seed-merchants are very particular in this respect. In
other instances the dodder and clover seeds approach each
other so nearly in size that sifting one from the other is
impossible, and the No. 17 sifter becomes quite useless.
J^Ajf;>
mm
X 5
FIG. 48.
Seeds of Red Clover, Yellow Trefoil, Dutch Clover, and Clover Dodder.
Enlarged 5 diameters^
For the purpose of comparison, two seeds of perennial red
clover are illustrated at Fig. 48, A ; at B the seeds of
yellow trefoil ; at C of white Dutch clover ; and at D
the seeds of clover dodder, all to the same scale, viz. five
times the size of nature. All clovers in cultivation vary
in size between the limits shown by A and C in Fig. 48.
The examples for measurement were kindly forwarded by
Messrs. Sharpe and Co. of Sleaford, Messrs. Sutton and
Sons of Eeading, and Messrs. Edward Webb and Sons of
Wordsley, Stourbridge. The dodder seeds were sifted out
of impure foreign importations. In a pound of average
clover there are 250,000 seeds.
A dodder seed is brown, dull, and minutely granular
outside when seen with the aid of a strong lens, whereas
118 DISEASES OF FIELD & GARDEN CROPS. [OH.
clover seed, although often brown, is smooth and shining,
with a minute scar or protuberance at one point of the
circumference at E, Fig. 48. At this point the radicle,
the first or elementary root of the plant, emerges at the
time of germination. This scar is almost invisible in
dodder seeds. When seen in section a clover seed mate-
rially differs from a dodder seed. The interior of the
clover seed shows the presence of cotyledons or seed-
leaves, as at Fig. 49, A, enlarged ten diameters ; and the
first rudimentary rootlet or radicle at B. The illustra-
tion at 0 shows a clover seed after it has been planted
for three days or a week in moist sand. The testa, or
outer integument of the seed, has burst, and the first
PIG. 49.
Seeds of Perennial Bed Clover and Clover Dodder, seen in section, and
germinating. Enlarged 10 diameters.
rootlet of the infant clover plant at D is descending to
the sand. Clover-dodder seed is engraved to the same
scale at E and F ; at E the seed is shown in section ;
there are no seed leaves, but the young plant within con-
sists of a simple thread, spirally coiled round a little
central mass of fleshy albumen. At the period of germi-
nation the thread emerges with a dilated end as at G,
and the granular coat of the seed frequently breaks up as
shown. If a germinating seed in this condition is trans-
ferred to a slip of glass and held before a strong light,
the spiral embryo will be seen through the cracked testa
as here illustrated. Sometimes the cotyledons or seed
XIX.]
CLOVER DODDER.
119
leaves are represented by one or two very minute scales
in germinating dodder.
Clover and dodder seeds, farther advanced in growth,
are shown at Fig. 50, twice the size of nature. The clover
seeds on the left are sending their first roots deep into the
earth, whereas the dodder seeds on the right are sending their
threads into the air in search of a host on which to live
parasitically. The ground line is indicated at A. Clover
and dodder seeds generally germinate after the lapse of a
week or less ; but certain seeds of clover dodder from the
X 2
FIG. 50.
Clover and Dodder seeds germinating.
Twice the size of nature.
sample we experimented with did not germinate till two
months had passed. The variation in the time of rest in and
on the ground before germination is an obvious advantage to
the dodder, for, if the first dodder seedlings find no hosts
ready for them, other seedling dodders, during a period
of two months, still have a good chance. Clover often
remains without producing other than the two first seed
leaves for one or two weeks, and during that period the
young dodders often attach themselves to the clover seed-
lings, as illustrated at Fig. 51, enlarged ten diameters.
120 DISEASES OF FIELD & GAKDEN CROPS. [en.
10
PIG. 51
Germinating Dodder seed, with its threadlike stem fixing on the seed
leaves or Cotyledons of Clover seedling.
Enlarged 10 diameters.
FIG. 52.
Infant Dodder plant on a young Clover leaf.
Enlarged 10 diameters.
xix.'] CLOVER DODDER. 121
For the same period the dodder threads, as a rule, have
produced no suckers, although in one or two erratic ex-
amples we have seen suckers whilst the embryo thread
was still within the burst testa. Dodders bear minute
transparent scales on their threads as equivalents to
leaves, but we have not seen them in clover dodder till
the young plants have been several weeks old. When
dodder twines round a young seedling clover, the rapidly-
growing clover carries the dodder away from the ground,
the old withered testa being sometimes still attached to the
dodder thread for a week or two. A dodder plant a few
weeks old, on a young clover leaf, is shown enlarged ten
diameters in Fig. 52. As the clover grows the dodder
now grows with it, and the parasite is lifted higher and
higher from the ground. As the spring and summer
advance, the dodder flowers profusely, and as the clover
plants grow in size and come in contact with each other, the
dodder spreads from one host to another. The dodder, in
growing, repeatedly branches and rebranches, and throws
out long arms, so that during a single summer one or
two infested clover plants will help to spread the dodder
over a large area. The parasite cannot live on the remains
of the plants it has destroyed, so, in the process of growth,
it leaves the central clover plant for other plants at the
circumference of a dead circle of clover, which may be
many feet or even yards in diameter.
At Fig. 53 is shown a fragment of clover stem cut
from the top of the stem at E, as illustrated at Fig.
47. This illustration is enlarged fifteen diameters to
show the connection by suckers of the twining dodder
with the stem of the clover. On examination with the
microscope it is seen that not only has the dodder no
roots or true leaves, but it is destitute of green colouring
matter, the substance which helps to elaborate the food
of plants, and which occurs so abundantly in clover.
Dodder has none of the small mouths or organs of tran-
spiration so frequently adverted to in these notes, but its
122 DISEASES OF FIELD & GARDEN CROPS. [CH.
outer cells, seen at A, 011 the contrary, are somewhat like
the pallisade cells found on the upper surface of ordinary
leaves. In the centre of the dodder threads there is a
woody cylinder, as at B, surrounded by colourless, some-
what loose, cellular tissue, C. The suckers, D, proceed from
•X-15
FIG. 53.
Fragment of Clover stem with Dodder entwined to show the connection
Toy the suckers of Dodder. Enlarged 15 diameters.
the central woody portion of the threadlike stem, and are
surrounded by a wall of colourless cells. The suckers of
Cuscuta Trifolii, Bab., are pushed into the fine longitudi-
nal furrows EE (which are always present in clover stems),
xix.] CLOVER DODDER. 123
until they reach, the central pith FF, as illustrated.
The suckers could not penetrate the clover stem were it
not for the woody skeleton belonging to each sucker ;
this atom of hard pointed woody material pierces the stem
like a small thorn. Hairs of the clover stem are shown
at GG. The connection of a sucker of dodder with the
clover stem, as seen under the microscope, is illustrated,
farther enlarged to fifty diameters, in Fig. 54. The outer
pallisade-like cells of the dodder are shown at AA, AA.
The woody cylinder at BBB, the loose cellular tissue at
CCC, and the point of a sucker, inserted in a clover
stem, at D. The centre of the clover stem is shown by the
pith -cells at E, the cells of the clover bark at FF, and
two fibro-vascular bundles at GG. The parasitic life of
clover dodder commences with the insertion of the first
sucker into the host plant. When the pith is reached by
the suckers pushing themselves in between the cells of
the stem of the host, the cellular tissue of the dodder
comes into close contact with the living cells of the
clover, and the result is, the vital juices elaborated
by the clover pass through the cell walls of the clover
into the cells of the dodder, and so the sap of the clover
feeds the parasite by transfusion. The dodder grows
with such extraordinary rapidity when it has once fixed
on clover, and it produces so many branches and brach-
lets, with such a vast number of suckers, that the growth
of the parasite generally far exceeds that of the host.
The consequence is, the dodder completely drains out
the elaborated juices of the clover and kills it by ex-
haustion. The destruction of the clover is also hastened
by the great weight of the accumulated masses of en-
tangled dodder ; for one commonly sees the clover quite
prostrate on the ground, whilst most of the thickly-
matted dodder growth is on the top. The profuse
growth of dodder in some clover fields was well illus-
trated by a correspondent of the Agricultural Gazette, who
wrote on 9th July 1870, p. 941, that his men had found
124 DISEASES OF FIELD & GARDEN CROPS. [CH.
X-5O
FlG. 54.
The anatomical connection of Clover Dodder with a Clover stem.
Enlarged 50 diameters.
several patches in the fields near Bridport, from two to
three yards square. The labourers, for a joke, clothed
xix.] CLOVER DODDER. 125
themselves with the matted threads of the parasite, and
went home as if clothed in bear skins. Clover dodder
is generally said to be an annual ; but observers are not
wanting who have expressed a belief that it is often
perennial, and lives on from year to year irrespective of
the introduction of new seed, which, indeed, it very
seldom produces in Britain. The dodders are said to
be acrid and purgative, and mischievous to flocks and
herds. It is singular that the parasite should be capable
of elaborating acrid principles from the juices of a sweet
non-acrid host.
Dodders are still largely imported to Britain in unclean
foreign seed. Prof. Lindley has stated that both clover
dodder and flax dodder were first imported to this
country from Afghanistan so lately as 1843. Dodder is
so common now that Prof. Buckman, to whom we are
indebted for a most interesting and instructive essay on
dodders, records an instance of seventy bushels of flax
dodder seeds being sifted out of a single field of flax seed,
whilst a year or two afterwards almost as much was separ-
ated from a crop of flax grown at the Royal Agricultural
College.
On rare occasions clover dodder produces seeds in
Britain ; and as there is evidence that the threadlike
stems are sometimes perennial, dodder refuse should
never be left on the ground to rot. Every patch of
dodder should be carefully raked together and burnt,
and by this process and careful sifting its appearance in
the fields can generally be prevented. Some agriculturists,
on first seeing the yellow patches in the clover fields,
remove all the clover from the outer edges of the invaded
patch for a width of about eighteen inches ; this leaves
nothing for the dodder to prey upon, as the threadlike
stems cannot stretch across the eighteen inches of vacant
ground. The clover is removed because it is extremely
difficult to entirely remove dodder.
CHAPTER XX.
GRASS MILDEW.
Erysiphe graminis , D.C.
THERE are few fungi more common or injurious in warm
dry weather than the fungus of grass mildew, grass blight,
or wheat rust, — or white rust, as it is termed in America,
named Erysiphe graminis, D.C. It is possible that the
familiar straw blight of agriculturists, already described,
may be caused by the mycelium of the Erysiphe, and this
is an additional reason for directing special attention to
grass mildew or blight. The generic name Erysiphe was
the term given to mildew by the Greeks ; the specific
name graminis needs no explanation. The fine, creeping,
jointed mycelium of Erysiphe graminis, D.C., forms a
white superficial mildew on the living stems and leaves
of cereals and other grasses in the summer and autumn.
When the white mildew patches are examined in the
autumn with a very strong lens, they will be seen
sprinkled with minute black dots, as illustrated twice the
natural size on the wheat stem in Fig. 55. The spawn
of this mildew is generally supposed to be incapable of
penetrating the tissues of plants ; but suckers have been
described as belonging to the mycelia of some allied species
of mildew. With these minute suckers the Erysiphe
adheres to its host, if the suckers do not indeed pierce the
leaf cells and derive nourishment therefrom, after the
manner of Peronospora. In the summer the mycelium
gives rise to vast numbers of vertical moniliform, or neck-
lace-like groups or chains of conidia or spores, as illus-
trated at Fig. 56, enlarged 400 diameters. This peculiar
CH. XX.]
GRASS MILDEW.
127
fruit of the mycelium was at one time considered a perfect
fungus, and was described under the name of Oidium moni-
lioides, Lk. The meaning of Oidium has been already ex-
plained, and monilioides means necklace-like, and refers to
the growth of the fungus, which resembles a string of beads.
X-2
FIG. 55. — GRASS MILDEW.
Wheat Stem invaded
byErysiphe graminis,
D.C. Twice the na-
tural size.
X-400
FIG. 56.
Oidium monilioides, Lk. The early condition
of Erysiphe graminis, D.C., enlarged 400
diameters. Germinating conidium enlarged
1000 diameters.
The Oidium is extremely common on the Graminece in the
summer, and it may always be found on grass, and especially
rankly growing grass in damp positions. The necklace-
like growth of the beadlike conidia is so delicate that the
slightest touch or breath destroys their cliainlike arrange-
128 DISEASES OF FIELD & GARDEN CROPS. [OH.
ment. If, however, infected grass is kept in damp air, a
fragment of a leaf may generally be successfully cut,
examined without water with a low power of the micro-
scope, and the Oidium seen in a growing condition. When
thus examined the profuse chainlike growth may be easily
observed. If placed under a cover glass for examination
under the higher powers of the microscope, the beadlike
spores or conidia instantly break away from each other,
so inconceivably slight is their attachment. The monili-
form habit can, therefore, only be seen with a low power
applied to the dry living fungus whilst in situ. In water
or damp air the conidia, and especially the topmost coni-
dium, quickly germinate and produce thin threads, as
illustrated at Fig. 56, A, enlarged 1000 diameters. This
constant spore production and germination is incessantly
continued through the summer months, till at last a thick
grayish-white coat of mildew more or less covers all in-
fected plants. The conidia are so small that it would
take about a million to cover a square inch. The Oidium
state of grass blight may be compared with the Oidium
of the turnip, illustrated to the same scale at Figs. 27 and
28. The grass Oidium is somewhat taller, but the spores of
the turnip Oidium are more than twice the length of those
of our present plant. Oidium monilioides, Lk., is a typical
plant. The Oidium growth of the fungus, however, and
the production of the profuse mycelium is only a prepara-
tory stage of growth for the perfect Erysiphe which gene-
rally follows ; it has been observed that when the Oidium
does not appear till late in the summer, the Erysiphe or
perfect condition is never produced, and the whole growth
of the fungus is confined to the Oidium stage. Under
favourable conditions of growth, the Oidium threads of
the summer produce in the autumn little brown globose
bodies termed conceptacles. This condition of the mildew
is shown on the wheat stem in Fig. 55. When examined
even with a powerful lens, these little blackish dots, termed
conceptacles, appear less in size than fine grains of dust ;
XX.]
GRASS MILDEW.
129
but when magnified with a moderately high power of the
microscope, they appear as spherical bodies, furnished with
a large number of slender, curved, radiating, tentacle-like
arms or branches, the whole growth being partly buried
in the spawn, as illustrated in Fig. 57, enlarged 100
diameters. The limit of the page will not admit of a
higher magnification than 100 diameters; it must there-
Fig. 57.
Conceptacle of ErysipJie graminis, D.C.
Enlarged 100 diameters.
fore be borne in mind that this Erysiphe is a comparatively
large fungus when placed side by side with others which
are magnified 200 and 400 diameters. The difference
in size between this Erysiphe and Peronospora exigua,
W.Sm., illustrated, enlarged 400 diameters, in Fig. 2, is
very striking.
130 DISEASES OF FIELD & GARDEN CROPS. [CH.
The term conceptacle explains itself; it simply means
a receptacle formed of one valve. Sometimes these bodies
are termed perithecia, or boxes enclosing the bladders,
termed asci or thecce. These bladders always in turn
enclose spores or sporidia. The conceptacles arise, it has
been said, at a point where two specialised threads of
spawn cross each other, and where the enlarged ends of
two spawn or mycelium tubes come in contact in a
manner similar, it has been said, with the contact of the
anther with its pollen with the stigma in flowering plants.
•X-200-
FIG. 58.
Horizontal section through a Conceptacle of Erysiphe graminis, B.C.
Enlarged 200 diameters.
The conceptacle or perithecium, after it is once formed
on the mycelium, quickly grows in size, and speedily
acquires a brown or blackish tint. The wall of which it
is composed is built up of minute, firm, closely -com-
pacted cells, resembling externally, on a very small scale,
the irregular pattern so frequently seen belonging to the
epidermis of many leaves, as of the pea. Towards the
base of the perithecium certain privileged cells throw out
curved unbranched processes or tentacle-like filaments of
xx.] GRASS MILDEW. 131
different sizes, some very small, and others comparatively
long. The largest of the mature perithecia are, however,
so small that it would require more than 100,000 to
cover a square inch. To understand the structure of
these minute perithecia and their contents it is necessary
to cut them in two both horizontally and vertically. Two
sections of this nature are shown at Figs. 58 and 59,
enlarged 200 and 100 diameters. By referring to the
horizontal section, enlarged 200 diameters, at Fig. 58, it
will be seen that twenty-four closely-packed transparent
bladders have been cut across ; whilst in the vertical
section, enlarged 100 diameters, at Fig. 59, five of the
x-ioo
FIG. 59.
Vertical section through a Conceptacle of Erysiphe graminis, D.G
Enlarged 100 diameters
central bladders have been cut through vertically.
With a little careful manipulation these contained
bladders or asci may be squeezed out of the perithecium,
and then, upon the application of a higher power of the
microscope, it will be seen that each bladder contains
eight spores, technically termed sporidia. A single
bladder or ascus, with its spores in situ, is shown both in
Figs. 58 and 59. Three of these asci, farther enlarged to
500 diameters, are illustrated at Fig. 60. One ascus con-
taining the normal eight sporidia in situ is shown at A,
whilst the figure at B shows the sporidia emerging from
the top of the ruptured ascus. The transparent asci are
so small that it would take 2j millions of them, contain-
ing 21 millions of sporidia, to cover a square inch. The
peculiar shape of the oval sporidia, slightly flattened in
their longest dimensions like the egg of a tortoise, is
worthy of note. One of these bodies is shown, enlarged
132 DISEASES OF FIELD & GARDEN CROPS. [CH.
1000 diameters, in the act of germination at C. The
asci and sporidia of Peziza postuma (B. and Wils.), illus-
trated to the same scale in Fig. 9, may be referred to for
comparison of size and form.
The autumn-borne perithecia do not burst and set free
their contained sporidia during the autumn in which they
have been formed, but they fall to the ground with the
decaying grasses on which they have lived during the
summer. They rest on the ground during the winter,
x
FIG. 60.
Asci and Sporidia of Erysiphe graminis, B.C., enlarged 500 diameters.
Germinating Sporidium, enlarged 1000 diameters.
and the hard wall of the perithecium effectively protects
the contained asci, and the asci in turn protect the
tender sporidia against all ordinary frost, dryness, or damp.
No sign of life can be detected in the fallen perithecia till
the following spring or early summer, and then, if old
decaying grass or straw is searched over, the perfectly un-
injured perithecia may be found. In the early summer
these bodies burst on the ground, as illustrated at Fig. 61,
xx.] GRASS MILDEW. 133
enlarged 100 diameters. When they burst the contained
bladders or asci often burst at the same time, and the
living sporidia, after their six months' rest, fly into the air.
At other times the bladders or asci themselves fly out of
the perithecia, and sail, each with its little load of eight
sporidia, through the air. When in the air the asci or
bladders burst, and the spores are set free in the atmos-
phere. The facts just given can easily be seen when old
grass or straw infected with Erysiphe is kept in damp air
FIG. 61.
Concoptacle of Erysiphe graminis, "D.C., bursting in spring.
Enlarged 100 diameters.
under a bell-glass, if strips of glass smeared with glycerine
are suspended over the infected leaves. On to these
strips of glass the asci and sporidia will be ejected from
the perithecia. It is a curious fact that the little sporidia
are so ready for germination in the spring that they com-
monly burst and produce spawn threads as they sail
about in the air.
Such of the sporidia as alight on the Gramine® or
grasses attach themselves to their hosts by their spawn
threads, and speedily produce a necklace-like Oidium
134 DISEASES OF FIELD & GARDEN CROPS. [CH.XX.
similar with the one which, preceded their own Erysiphe
condition.
Dr. H. W. Harkness has described, hefore the Micro-
scopical Society of San Francisco, pycnidia upon the
spawn of this fungus as found in California. Pycnidia
are small conceptacles containing stylospores, or spores
after the nature of conidia. The "white rust," as it has
been termed in America, first appeared in California in
1877 on mature wheat. In that year it covered half a
million acres of wheat, destroying some of the crops.
We have taken pains to completely illustrate this com-
mon, curious, and destructive fungus, for, as far as we
know, no complete illustrations have hitherto been given.
LeVeille's illustration, generally referred to, in the Ann.
des. Sc. Nat., vol. xv., 1851, t. 10, f. 33, is very bad.
It shows the appendages branched and the asci without
sporidia ; the scale of magnification is also omitted.
Hops, peas, beans, roses, and many other plants, are
preyed upon by fungi very closely allied to the fungus of
grass blight.
Grass blight is synonymous with bad, impoverished
grass, with mildewed hay, and ill-nourished herds and
flocks. The life of the fungus which causes the blight
is preserved through the winter in decaying mildewed
grass and straw. These facts therefore teach us that all
mildewed material should, as far as possible, be gathered
together in the late autumn or winter and destroyed. If
all the autumn-borne perithecia are destroyed it is obvious
there can be no Oidium in the spring. The destruction
of all infected material is perhaps impossible, but all the
facts known in reference to the fungus of this disease
point to the great desirability of clean farming, and to
the necessity of destroying as much mildewed grass and
straw refuse as possible. If all farmers would agree to
one course of action the bad effects of diseases like the
one here described would be greatly lessened.
CHAPTER XXI.
CORN MILDEW — SPRING RUST AND MILDEW.
Puccinia JRubigo-vera, D.C.
THE disease of wheat, popularly known under the name
of corn mildew, is the best known and most widely-spread
of all plant diseases. Agriculturalists are familiar with the
reddish spots of '« rust " early in the season, and, later on,
the black spots of autumn and winter termed " mildew."
These spots are different conditions of the same parasitic
fungus, the "rusts" being early conditions of the "mildews."
Many observers believe that rusts and mildews have a
third condition of growth, and that the mildews of wheat
are capable of jumping over an apparent gap and causing
a blight of comfrey, bugloss, alkanet, or barberry ; and
the blights of these plants, termed jflcidia, in turn being
capable of causing new " rusts," and ultimately " mildew "
of corn.
The subject of the mildews of corn may be approached
from several points ; and as farmers first become acquainted
with the pests every year in the spring, when the stems
and leaves of their cereals become rusted, we will start
with the familiar rust of spring, premising that botanists
term the rust Uredo, and the mildew Puccinia. These
terms have been already explained.
There are two rusts of corn — one termed Uredo Rubigo-
vera, D.C., the other Uredo linearis, Pers. Both are
followed by a Puccinia, the first by P. Rubigo-vera, D.C.,
and the second by P. graminis, Pers. The first, Uredo,
with its Puccinia, is less known and less generally in-
jurious than the last ; but as it appears first in the season
136 DISEASES OF FIELD & GARDEN CROPS. [en.
and sometimes entails great losses, it may be more con-
venient to describe it first.
The spring rust of wheat appears in March, April, and
May, on grasses and cereals : it is common on soft grass,
Holcus lanatus, L. ; creeping soft grass, H. mollis, L. ;
barren Brome grass, Bromus sterilis, L. ; and wall barley,
Hordeum murinum, L. Its specific name, — Rubigo-vera,
means "true rust."
If a leaf of wheat invaded by "true rust" be taken
in the month of April or May and examined with a strong
lens, it will be seen, as on Fig. 62, enlarged three diameters.
The minute livid yellow
pustules are the sori of
the Uredo; in the bottom
left-hand corner of the
illustration a few black
spots will be seen, —
these are the advanced
or Puccinia condition of
the parasite. If we place
a fragment of the leaf
under a low power of
the microscope and mag-
nify twenty -five diam-
eters, we shall see the
-x-3- pustules as at Fig. 63.
FIG. 62. — Fragment of Wheat leaf in- It is now evident that
vacled by Vredo Riibigo-vera D.C. the fungUS within the
plant has, in reaching
maturity, burst the epidermis of the wheat leaf. The
fungus may be seen as a fine yellowish or orange
powder, which is set free by the slightest touch and
carried away in the air by the faintest breath. If we
now make a transverse section through a very small
pustule, cut off a transparent slice from the exposed sur-
face, and magnify 200 diameters, we shall see the interior
of the pustule as at Fig. 64. This is engraved to the
XXI.]
SPRING RUST AND MILDEW.
137
same scale as Pucdnia mixta, Fl., Fig. 13, which may be
referred to for comparison.
The yellow powder is now
seen as a mass of yellowish
sub-spherical or ovoid spores,
each spore filled with dense
granular protoplasm, is sup-
ported on a short stalk and
springs from an involved
stratum of mycelium. A single
spore is seen emerging through
the centre of a ruptured pus-
tule at A. The transparent
circles at BB are part of the
cellular tissue of the wheat
leaf. When the spores escape
in this matter, as they do in
inconceivable numbers, many
fall on to the leaves from
which they have grown. In
their new position they germi-
nate, as at Fig. 65, by the protrusion of a spawn-thread
from both sides, the vital material pours from the spore
11
X • 25
FIG. 63.— Pustules or Sori of Uredo
Bubigo-vera, B.C., on Wheat
leaf. Enlarged 25 diameters.
r
FIG. 64.
Transverse Section through a pustule or Sorus of Uredo Rubigo-vera, D.C.
Enlarged 200 diameters.
into the threads ; and as the spores get empty a septum
appears as at A, and cuts off the connection of the germ-
tube with the spore. The germinal threads now
138 DISEASES OF FIELD & GARDEN CROPS. [en.
the organs of transpiration of the leaf, and there form a
new stratum of mycelium, from which new Uredo pustules
arise. Under favourable circumstances this growth and
•X-IOOO
FIG. 65.
Germinating spore of Uredo Rubigo-vera, D.C.
Enlarged 1000 diameters.
regrowth of successive crops of spores goes on for several
weeks, until the whole plant, whether wheat, rye, or some
wild grass, is at length permeated by the spawn.
The above -described condition of spring rust is the
same with Uredo Rubigo, D.C., Uredo Rubigo-vera, Lev.,
Coeoma Rubigo, Lk., Trichobasis Rubigo-vera, Lev., and
Trichobasis glumarum, Lev.
As the autumn and winter approach, the pallid and
yellowish spots of the Uredo vanish, and black spots
appear, and this black condition is the mature fungus of
the mildew named Puccinia Rubigo-vera, D.C. A small
piece of wheat-stem is illustrated, enlarged five diameters,
at Fig. 66, the small black dots show the Puccinia
pustules. Reference may be here made to Fig. 12, where
similar pustules are shown, to the same scale, on a frag-
ment of a flower stem or scape of chives. The black sori
are farther enlarged to twenty-five diameters at Fig. 67,
to show that the pustules are almost identical in appear-
ance with the Uredo sori of Fig. 63 ; the only difference
SPKING RUST AND MILDEW.
139
is the Uredo produces simple oval yellow spores ; whilst
the Puccinia produces dark brown or blackish spores, each
of the latter having a joint or septum across its narrowest
diameter.
A section through a Puccinia
sorus is shown, enlarged 200 diam-
eters, at Fig. 68 ; the Puccinia
spores are pushing in the same
manner as the Uredo through the
ruptured epidermis of the wheat
leaf. The dark elongated bodies at
A A are said to be paraphyses — pos-
sibly in this case undeveloped
X-5
FIG. 66.— Fragment of Wheat stem
invaded \tyPucdnia Rubigo-rcra,
D.C. Enlarged o diameters.
-X-25
FIG. 07.— Pustules or Sori of P«c-
ciniaRubigo-vera, D.C. Enlarged
25 diameters.
Puccinia spores. The compound spores of Pwcinia
Rubigo-vera, D.C., act as resting -spores, for although
140 DISEASES OF FIELD & GARDEN CROPS. [CH.
developed in the autumn, they will not, as a rule,
germinate till the following spring ; they then burst in
the manner illustrated at Fig. 69, enlarged 1000 dia-
meters, upon old decaying grass and straw upon the
ground. In the warm damp weather of April and May
one or both segments of a teleutospore will burst and
produce a transparent thread of mycelium, termed by
botanists pro-mycelium, because it is the first mycelium
of a cycle of phenomena belonging to Puccinia. The
compound teleutospore is shown at AA, with its short
stalk B still attached. As the pro-mycelial threads, CO,
Fig. 68.
Transverse section through a pustule or Sorus of Puccinia Rubigo-vera, D.C.
Enlarged 200 diameters.
increase in length, the protoplasm pours from the spores
into the tubes. A series of septa then appear, two of
which are seen at DD, and these septa enclose the
protoplasm in the growing end of the tube ; from this
end two or three minute transparent or pale yellowish
spores, termed pro -mycelium spores, as at EEE, are
borne : these speedily fall from their slender supports,
and germinate very readily on damp surfaces, as shown
at F.
It might be considered reasonable to suppose that these
little pro -mycelium spores, as produced in the spring,
would, if placed on the leaves of grasses, reproduce the
Uredo first described ; but many botanists believe it to be
proved that they do not and cannot at once reproduce the
Uredo ; but when placed on the leaves of certain plants
SPRING RUST AND MILDEW.
141
belonging to the Borage family, as Lycopsis arvensis, L.,
Anchusa officinalis, L., Symphytum tuber osum, L., etc.,
they produce an apparently totally different fungus,
X-IOOO
FIG. 69.
Teleutospore of Puccinia Rubigo-vera, B.C., germinating in spring.
Enlarged 1000 diameters.
termed JEcidium asperifolii, Pers. When the JRd&ium
in due course produces its spores it is said that the
142 DISEASES OF FIELD & GARDEN CROPS. [CH.XXI.
spores give rise to the Uredo if placed on
certain grass leaves. In the next chapter will be found
a description and illustration of the ^Eddium.
Before dismissing Pucdnia Rubigo-vera, D.C., we may
say that Uredo and Pucdnia spores often, though not in
this species, grow together in the same pustules, and that
P. Rubigo-vera, D.C., and P. graminis, Pers., often grow in
company on the same leaves and stems. These parasites
divert the material which should go to the production of
good ears of grain for their own support ; they therefore
cause the general growth of corn to be weak and the ears
to be small in proportion to the virulence of the attack:
the straw, too, is not only damaged, but is made the
means of carrying the disease over the winter for the
following season. A variety of P. Rubigo-vera, D.C.,
termed P. simplex, Kor., also occurs. The teleutospores
in this plant consist chiefly of a single cell, similar to the
single-celled examples of P. m-ixta, FL, illustrated in the
left-hand spore in Fig. 14. P. simplex, Kor., has also
been described as a species under the name of P. Hordei,
Fl., and P. anomala, Rost.
Pucdnia Rubigo-vera, D.C., occurs in Europe on Cala-
magrostis ISpigejos, Roth. ; Arrhenatherum elatior, L. ; Holcus
mollis, L. ; H. lanatus, L. ; Avena flavescens, L. ; Festuca
elatior, L. ; Serrafalcus secalinus, Bab. ; Bromus mottis, L. ;
Secale cereale, Walld. ; B. arvensis, L. ; B. asper, L. ; Triti-
cum vulgare, VilL ; Lolium temulentum, L. ; Hordeum vul-
gare, L. ; H. distichum, L. ; H. murinum, L. ; and H.
secalinum, Trin.
Suggestions for preventing and destroying the mil-
dews of corn are adverted to in the chapter where the
evidence for the connection of Pucdnia and JEddium is
reviewed.
Pucdnia Rulngo-vera, D.C., is the same with P. stra-
minis, FL, and P. striasformis, West
CHAPTEE XXII.
BORAGE BLIGHT.
dEcidium asperifolii, Pers.
As the disease of barberries, believed by many botanists
to be one condition of the fungus of summer rust and
mildew of wheat, is popularly known as " barberry blight,"
it may be well to term the closely allied fungus of the
Borage family, which is similarly associated by many with
the spring mildew of corn, borage blight.
The name of the fungus blight found on various plants
belonging to the Borage family, and considered by many
observers to be one form of the spring rust of corn, is
^Ecidium asperifolii, Pers. The name JEcidiwm is derived
from the Greek, and should properly be written (Ecidium ;
the word means a little chamber, in reference to the form
of the fungus in an infant state, as shown at C in our
illustration of the ^Ecidium of summer mildew of corn,
Fig. 83 ;— asperifolii, refers to the nature of the coarse
hairy leaves of several of the Boraginaceous plants on
which the JEcidium grows.
The fungus of spring rust and mildew, already described,
is generally common in Britain, — so common that persons
walking through wheat fields have sometimes had their
boots and clothes covered with the orange spores. The
SEcidium, on the other hand, said to be one condition of the
Uredo and Puccinia, is one of the rarest of British plants.
JEddia grow within the tissues of plants, and in these
positions they form minute spherical balls, filled with
chains of whitish or yellowish, semi-transparent, generally
spherical, spores. In the process of growth the immersed
144 DISEASES OF FIELD & GARDEN CROPS. [on.
spheres burst through the cuticle of the host plant — gener-
ally from the under side of the leaves, and often through
the stem. As the JEcvdvwm cups mature they burst at the
exposed apex, and the fractured part turns back so as to
give the little fungus growths the form of minute cups
filled with spores.
The under side of a leaf of Tuberous Comfrey, Sym-
phytum tuberosum, L., is illustrated, natural size, at Fig.
70. Two groups of the cups belonging to
x-io
FIG. 70.
Leaf of Tuberous Comfrey, Symphytum tuberosum, L., invaded
asperifolii, Pers., natural size.
jficidium cups at B enlarged 10 diameters.
asperifolii, Pers., are shown at AA. Each cluster of
cups is surrounded by a large pallid disease patch, which
has been caused by the exhaustion of the vital material of
the leaf by the spawn of the fungus which at first grew
within. The fungus growth has also caused the leaf to
become torn. Five of the little AZcidium cups are shown,
enlarged to ten diameters, at B.
At A, Fig. 71, one of the mature Medium cups is
farther enlarged to fifty diameters ; the cup has burst, and
XXIL] BORAGE BLIGHT. 145
its frayed edges, made up of transparent polyhedral cells,
are outwardly curved. Springing from the interior are
the chains of globose (or slightly polyhedral) spores. The
cells of the wall of the cup and the contained spores
are nearly the same in size, the spores being some-
what larger than the cells ; the difference in the appear-
ance of the two under the microscope is very striking,
for the component cells of the cup have a thin wall,
and are filled with a watery fluid, whereas the spores have
two or even three coats, and are filled with granular and
lustrous protoplasm. The spores, unlike the constituent
cells of the ^Ecidium, are commonly studded with minute
1000
FIG. 71.
Mcidium asperifolii, Pers. Single cup enlarged 50 diameters ;
Spore enlarged 1000 diameters.
sub-globose particles as at B, enlarged 1000 diameters ;
these small bodies, as seen attached to the circumference
of the spore, are considered to have the same function as
pollen-grains, and are termed spermatia.
As the nature of ^Ecidium is fully explained under
sEcidium Berberidis, Pers., Chap. XXIV., it is not neces-
sary to describe it in this place, especially as A. asperifolii,
Pers., is not a completely typical species.
It is sufficient here to say that Professor A. de Bary of
Strasbourg states that the spores of this JScidiwn will
not reproduce an jflcidium on germination, but, on the
L
146 DISEASES OF FIELD & GARDEN CROPS. [OH.XXII.
contrary, will produce Uredo Rubigo-vera, D.C., if placed
on the leaves of various grasses and cereals. Professor
de Bary states, Neue Untersuchungen uber Uredineen, ii.
1866, that the germ tubes of the ^Ecidium spore, on
germination, bore into the cuticular cells of certain grasses,
and develop a mycelium in the body of the leaf, which
mycelium at length gives rise, after about eight days, to
Uredo Rubigo-vera, D.C.
JEcidium asperifolii, Pers., has been found in Europe
on Gynoglossum officinale, L. ; Borrago officinalis, Tour. ;
Anchusa officinalis, L. ; A. arvensis, Bieb. ; Nonnea pulla,
D.C. ; Symphijtum officinale, L. ; S. tuberosum, L. ; Cerinthe
minor, L. ; (7. alpina, Kit. ; Echium vulgare, L. ; Pulmon-
aria officinalis, L. ; P. tuberosa, Schrk. ; and Lithospermum
arvense, L.
CHAPTER XXIII.
CORN MILDEW — SUMMER RUST AND MILDEW.
Puccinia graminis, Pers.
THE summer rust and mildew of corn, as caused by the
fungus known as Puccinia graminis, Pers., is of greater
importance from an economical point than the spring
rust and mildew already described. The losses entailed
on farmers by attacks from Puccinia graminis, Pers., have
sometimes reached 50 or even 75 per cent on the whole
crop ; where there should have been 40 or 50 bushels
only 20 have been harvested, and in some instances only
12 have been recorded.
The first parasite may be found in its early or Uredo
state in March, April, and May, whilst the summer rust
is seldom seen till June or July.
As in the last, the early condition of summer mildew
is the rust stage, termed in this case Uredo linearis,
Pers. The generic name has already been explained ;
linearis refers to the elongated form of each dot or
disease pustule. The black condition of the fungus, as
found in autumn and winter, is the one termed Puccinia
graminis, Pers. The name Puccinia has been explained ;
graminis needs no explanation.
Botanists in former times believed the red and black
spots of rust and mildew to be caused by two distinct
fungi ; it is, however, very common to see both Uredo
and Puccinia spores in the same pustule.
Part of a wheat leaf suffering from summer rust, Uredo
linearis, Pers., is shown at Fig. 72, enlarged three
diameters. The Puccinia or mildew is replacing the
148 DISEASES OF FIELD & GARDEN CROPS. [CH.
Uredo in the black spots in the bottom left-hand corner
of the illustration. Two pustules (technically sori, or
spore cases), are further enlarged to twenty-five diameters
in Fig. 73. It will now be instructive to turn to Figs.
62 and 63, where the spring rust is illustrated to the
same scale. ,
•X-2S-
FIG. 73. — Pustules or Sori of
Uredo linearis, Pers. Enlarged
25 diameters.
•X-3-
FIG. 72.— Fragment of Wheat leaf in-
vaded "by Uredo linearis, Pers.
Enlarged 3 diameters.
Neither Uredo Rubigo-vera, D.C., or U. linearis, Pers.,
are confined to one side of the foliage of cereals ; they
grow on both sides, and (especially in U. linearis, Pers.)
attack the inflorescence with its glumes and pales. The
latter organs have been described under ear-cockle.
The Uredo of summer rust is less livid or yellow and
more orange or brown in colour than the Uredo of spring
rust, and being altogether larger and more robust in
growth it splits and lacerates the cuticle of the affected
plant more completely ; but the description of the pustules
of spring rust apply generally to the larger sori of the rust
of summer.
XXIIL ] SUMMER RUST AND MILDEW. 149
If we now cut across the larger of the two pustules
of summer rust, as illustrated at Fig. 73, and magnify a
thin transparent slice from the cut surface 200 diameters,
as was done with U. Eubigo-vera, D.C., we shall see it as
at Fig. 74. We now find that a typical pustule of
summer rust is so large that the page of our book is in-
sufficient for it. The engraving at Fig. 74 therefore shows
one-half the diameter of a sorus only, AB being the centre
line. The closely packed Uredo spores are supported on
somewhat longer pedicles than in U. Rubigo-vera, B.C.,
and the spores themselves differ not only in colour but also
A
; -x 200^
FIG. 74.
Transverse section through half a Pustule or Sorus of Uredo linearis, Pers.
Enlarged 200 diameters.
somewhat in shape and size. They spring from a stratum
of involved jointed brownish mycelium. The constituent
cells of the wheat leaf are shown at Fig. 74, C, and
two organs of transpiration belonging to the leaf are shown
at DD. Two typical Uredo spores, supported on their
transparent pedicles, are enlarged 1000 diameters in Fig.
75. No individual spore in a sorus exactly agrees in
shape and size with another ; they vary, however, within
well-defined limits. Germination has commenced in the
smaller of the two spores at A.
The Uredo spores fall very readily from their support-
150 DISEASES OF FIELD & GARDEN CROPS. [CH.
ing stems, and as they fall the faintest breath of air wafts
them away.
If Uredo spores are kept in moist air for a few hours
they readily germinate. This germination usually takes
place by the protrusion of two germ tubes or threads of
mycelium, one from each side, usually near the middle,
and rarely from or near the top of the spore. The spots
whence the tubes are destined to emerge from the spores
X-IOOO
FIG. 75.
Spores of Uredo linearis, Pers. Enlarged 1000 diameters.
can often be readily seen before germination takes place
as at BC, Fig. 75. These germinal spots are weak
places in the inner wall of the spore. Two spores,
enlarged 400 diameters, are shown germinating on a frag-
ment of the epidermis of a wheat plant in Fig. 76.
These two spores show the extreme limit of size. It
seldom happens that both germ tubes continue their
growth ; one usually remains effete, as at AA, whilst
XXIII.]
SUMMER RUST AND MILDEW.
151
the yellowish granular contents of the Uredo spore pour
into the other and more strongly growing tube, BB, Fig.
76. In a day, or a day
and night, this stronger
thread will have grown in
a convolved fashion, as at
C, and will have attained
many times the length of
the spore from which it
started, and the whole of
the contents of the spore
will now be in the ger-
minal tube. When this
stage of growth is attained
a septum or stop grows
across the my celial thread,
as at D, and the dead and
empty spore case is cut
off from the living thread.
If this process of ger-
mination is watched on
glass and not on a cuticle
stripped from a wheat leaf, it will be noticed that the
germ tube will flow into any little scratch or depression
on the glass, just as a brook gradually flows into the
lowest positions of a river valley, or as rain water makes
its way into our brooks by following little depressions in
the land surface.
If, on the other hand, the process of germination is
watched on a shred of transparent epidermis torn from
a wheat leaf, the thread of mycelium will be observed
to follow the minute depressions formed where the con-
stituent cells of the epidermis meet, and by following these
fine depressions, as at E, the germ tube at length
naturally arrives at one of the minute mouths, organs
of transpiration, or stomata, belonging to the plant, as at
F. These are the lowest points on a leaf surface, and the
B
X-400JJ
FIG. 76. — Two spores of Uredo linearis,
Pers., germinating on a fragment of
the epidermis of a Wheat leaf.
Enlarged 400 diameters.
152 DISEASES OF FIELD & GARDEN CROPS. [OH.
germ tube of the rust fungus enters the tissues of the
host plant through these apertures, just in the same way
as it would enter a small hole in a microscopic slide or go
over the edge of a slip of glass, as we have many times
seen it do. A stomate belonging to wheat is very large
in comparison with the size of a spore \ the length of an
organ of transpiration is from G to H.
When once the Uredo mycelium has found its way
amongst the tissues of the wheat plant, the germ tube is in
its natural position ; it now branches right and left, and
ramifies amongst the green constituent cells of the leaf. The
mycelium here quickly produces new Uredo pustules, which
burst through the wheat cuticle in fresh places, so that a
wheat leaf which may have had only a few pustules in
June, may have the number more than quadrupled by July
by the continued germination of the Uredo spores on the
leaf surface. No doubt the mycelium also spreads in the
leaf from the base of the original pustules.
Before proceeding farther it must be noted that these
Uredo spores are very short lived, for if they germinate on
any unsuitable material they quickly perish. When all
the vital material is once poured out of the spore into the
germinal tube, growth can proceed no farther on an un-
suitable matrix. It is only when the spores germinate
on grasses, and the germ tubes find their way through the
organs of transpiration into the tissues of the supporting
plant, that the plasma of the fungus is able to continue its
existence. It is therefore obvious that when the frosts of
winter arrive and grasses are for the most part dead, or,
if alive, sluggish to the last degree, that this fungus,
which has a much more slender hold on life than any
grass, must perish. Unless the mycelium of the rust
fungus has moderate warmth, sufficient moisture, and the
interior of a living grass, it probably always collapses and
perishes.
As in the spring rust of wheat, the life of the fungus of
summer rust and mildew is carried over the winter in the
XXIII.]
SUMMER EUST AND MILDEW.
153
following manner : As the summer advances the rust
mycelium within the leaf of the wheat plant gradually
ceases to produce rust spores, and, instead, produces dark
brown or blackish spores — jet black
to the unaided eye. A fragment of
a wheat stem is illustrated at Fig.
77, enlarged five diameters ; the
jet black pustules here shown repre-
sent the genuine corn mildew of
agriculturists, and this perfect con-
dition of the disease is the Puccinia
graminis, Pers., of botanists. The
X 5
FIG. 77.— Fragment of Wheat
stem invaded by Puccinia
graminis, Pers. Enlarged
5 diameters.
X-25
FIG. 78. — Pustules or Sori of Pucdnia
graminis, Pers. Enlarged 25 dia-
meters.
black disease spots are much larger in P. graminis, Pers.,
than in P. Rubigo-vera, D.C., as may be seen by referring to
154 DISEASES OF FIELD & GARDEN CROPS. [CH.
Fig. 66. Three pustules are farther enlarged to twenty-
five diameters in Fig. 78. A reference to Fig. 67, where
the pustules of P. Rubigo-vera, D.C., are illustrated to the
same scale, will show the difference in size of the sori
belonging to the two mildews. The familiar condition of
the burst epidermis of the leaf is seen in both, and it is
noticeable in P. graminis, Pers., that in bad cases the
black pustules commonly become confluent.
We will now take a transverse section on the line AA.,
though the smallest of the three pustules illustrated in
Fig. 78. If we magnify a thin transparent slice from the
FIG. 79.
Transverse section through half a Pustule or Sorus of Puccinia graminis,
Pers. Enlarged 200 diameters.
cut surface 200 diameters as before, we find the limits of
the page insufficient for it. One-half the section, there-
fore, only is illustrated in Fig. 79, the centre line being
shown at AB. If reference is now made to the section
through a complete pustule of Puccinia Rubigo-vera, D.C.,
as illustrated to the same scale in Fig. 68, the difference
in size between the two sori will be apparent. The
difference in form of the contained teleutospores will be
seen when the teleutospores of Puccinia graminis, Pers.,
Fig. 80, are compared with the germinating teleutospore
XXIIT.]
SUMMER RUST AND MILDEW.
155
of P. Rubigo-vera, B.C., illustrated in Fig. 69, both en-
larged 1000 diameters. The supporting stems of Puccinia
graminis, Pers., are larger in proportion, and are accom-
panied by no paraphyses
in the sori.
To see the germination
of the black Puccinia spores,
old Puccinia invaded straw
must be looked for in the
spring months, and the
Puccinia spores must be
taken from a pustule with
a small knife or needle, and
placed in a film of water
under a thin cover glass on
a slide, and kept in moist
air (to prevent evaporation)
under a bell glass. Ger-
mination usually takes
place, as in P. Rubigo-vera,
D.C., by the protrusion of
a thread from each of the
two cells of the Puccinia
spore, as illustrated at Fig.
81, enlarged 1000 diam-
eters. These two threads,
the first produced in the
spring, are the pro -my-
celium, or the first my-
celium, of one end of the
cycle springing from the X-IOOO"
" finishing spores " or tele- FIG. SO.— Teleutospores of Puccinia
Utospores belonging to the graminis, Pers., as borne in autumn
T m1 . ' and winter. Enlarged 1000 dia-
other end. This pro-my- meterg>
celium is seen at AA. The
pro-mycelium proceeds to no great length, but after some-
times attaining three or four times the length of the black
156 DISEASES OF FIELD & GARDEN CROPS. [CH.
FIG. 81.
Teleutospores of Puccinia graminis, Pers., germinating in early summer.
Enlarged 1000 diameters.
teleutospore belonging to the Puccinia, it usually coils
round somewliat like a shepherd's crook, produces about
xxiii.] SUMMER RUST AND MILDEW. 157
three stops, joints, or septa, as at B, C, D, and from each
of the three separate pieces three fine branches arise, and
these branches bear at their tops three irregular oval
transparent, very pale amber-coloured spores, as illustrated
at E, F, G. These spores are the third of the series.
First we have Uredo, or rust spores ; then Puccinia, or
black mildew spores ; last, spring or pro-mycelium spores.
Pro-mycelium spores germinate very readily in a film
of water on glass, as illustrated at H, by the protrusion
of a fine tube of mycelium.
In a state of nature the black Puccinia spores germi-
nate upon straw, as it rots on the ground in the spring,
and the minute ovoid pale lemon -coloured spores are
carried about in the air in millions — that, too, in the
springtime, when corn first becomes invaded, and when
the first signs of summer rust or Uredo appear upon our
cereals.
Judging by what is well known amongst other fungi,
it would be perhaps reasonable to suppose that these little
hyaline spores (particularly as they arise from specialised
resting-spores) would reproduce the rust from which they
were originally derived (and nothing else), if they came
in contact with grasses. Many botanists believe this to
be a fact ; others say they do not, but, on the contrary,
that the cycle of corn mildew is not complete with the
production of these spores. Many observers believe that
before the pro-mycelium spores can cause the rust of corn
they must be nursed by a barberry bush ; that the lemon
coloured spores invariably refuse to eifectually grow on
the leaves of grasses, but when placed on the leaves of
barberries they find themselves so thoroughly in a natural
position that they do not, on germination (like the majority
of fungus spores) gently follow the uneven surface of the
leaf cells, and so quietly enter by the stomata ; but the
mycelium from the spores, it is said, sinks into the hard
leaves of the barberry, through the cells of the epidermis
(not between them) to the body of the leaf, and there,
158 DISEASES OF FIELD & GARDEN CROPS. [CH. xxm.
having gone from a monocotyledonous plant to a dicoty-
ledonous one, another fungus is produced of an appa-
rently totally different character, named Mcidium Berber-
idis, Pers.
This is the fungus of barberry blight, described in the
next chapter. Our comments on the possible connection
of the barberry fungus with the fungus of summer mildew
of corn is discussed farther on.
Puccinia, graminis, D.O., has been recorded in Britain
upon Phalaris arundinacea, L. ; Phleum pratense, L. ; Alope-
curus pratensis, L. ; A. fulvus, Sin.; Agrostis vulgaris,
With. ; A. alba, L. ; Calamogrostis Epigejos, Roth. ; Aira
ccespitosa, L. ; Avena sativa, L. ; A.fatua, L. ; A. pratensis,
L. ; A. flavescens, L. ; A. elatior, L. ; Holcus lanatus, L. ;
Poa annua, L. ; P. nemoralis, L. ; P. pratensis, L. ; Molinia
ccerulea, Moench. ; Dactylis glomerata, L. ; Festuca gigantea,
Vill. ; F. spectabilis, Jan. ; F. tenella, Willd. ; Bromus mollis,
L. ; B. tectorum, L. ; Lolium perenne, L. ; Triticum vulgare,
Vill. ; T. repens, L. ', T. caninum, Huds. ; Elymus arena-
riuSj L. ; E. glaucifolius, L. ; Hordeum vulgare, L. ; H.
sylvaticum, Huds. ; H. murinum, L. ; H. distichum, L. ;
ovata, L. ; Secale cereale, Walld.
CHAPTER XXIV.
BARBEERY BLIGHT.
^Ecidium Berberidis, Pers.
THERE is perhaps no family of plants more free from
fungi than the Berberidacece, and in this fact the family
greatly differs from the Graminece, on various members of
which the fungi of spring and summer mildew of corn are
so prevalent. It is also noteworthy that the Boraginacece,
upon some members of the order, as already described,
the supposed second condition of spring mildew of corn —
dEcidium asperifolii, Pers., — occurs, are also free from
the attacks of fungi to an extraordinary degree. Grasses
are all badly infested with fungus parasites and epiphytes.
The only important fungus peculiar to the barberry is
the one named ^Ecidium Berberidis, Pers. The generic
name ^cidium has been explained; the specific name
Berberidis explains itself.
jficidium Berberidis, Pers., is frequent on the common
barberry, Berberis vulgaris, L. It also grows rarely
on the more ornamental species of Berberis and on the
Mahonias of our gardens.
At Fig. 82 is illustrated, natural size, a few leaves
attached to a small fragment of a branch of the common
barberry. The parasitic ^Ecidium almost invariably grows
on the under surface of the leaves, as there shown, although
it may be detected rarely on both sides, and indeed on
every part of the plant. The ^cidium growths are seen
at AAA. The A^cidium clusters, of which there are ten
in the illustration, are groups of little sulphur-coloured
spots embedded in dark red, swollen, or hypertrophied
160 DISEASES OF FIELD & GARDEN CROPS. [CH.
patches, on the leaves. If we examine the upper surface
of the leaves we shall see reddish patches similar with
those below, but these upper patches are more or less
covered with little black dots technically termed spermo-
gones. These spermogones, of which there are four groups
in Fig. 82, are illustrated at BBB. Although the spermo-
gones usually grow on the upper surface of the leaf,
sometimes they may be seen on the lower surface. They
Fig. 82.
Barberry leaves invaded by JEcvlium Berberidis, Pers.
Natural size.
sometimes grow on the same patches with the
cups on the lower surface ; at other times they appear by
themselves without ^Ecidia. As each spermogone dot is
smaller than the point of a pin they are easily overlooked,
especially when they grow in very small companies, or as
single specimens. The spermogones usually appear before
the dEcidium cups.
If we look at the black dots with a strong lens we shall
still, owing to their excessive smallness, only see them as
XXIV.]
BARBERRY BLIGHT.
161
black dots ; but if we look at the JEcidiwm clusters with a
similar lens, \ve shall see companies of beautiful sulphury
yellow cups, bursting open through the lower epidermis
of the leaf, and each cup filled with yellow powder re-
sembling small pollen-grains.
To understand the nature of the dEcidium cups and the
black spermogone dots, we must cut a section through the
barberry leaf, and this section must be so made that it
FIG. 83.
Section through a Barberry leaf, showing the cups of jEcidium. Berberidis,
Pera. , below, and the Spermogones above. Enlarged 50 diameters.
will pass through the centres of the cups and the spermo-
gones. Such a section is illustrated at Fig. 83, enlarged
50 diameters. A represents the lower surface of the leaf,
and B the upper. At C one of the little ^cidia is seen
buried (a small chamber full of spores) in the tissues of
the leaf, and at DD two of the JScidium cups are seen
quite mature and open, the epidermis of the barberry leaf
M
162 DISEASES OF FIELD & GARDEN CROPS. [CH.
being rent. These cups may be compared with, the
dScidium cups of the allied JE. asperifolii, Pers., illustrated
to the same scale in Fig. 71. At EEE three of the
little black sperm ogones are seen in section. In the
middle of the leaf, at FF, amongst the green cells, may be
seen a longitudinally cut mass of vascular tissue or spiral
vessels belonging to one of the veins. The abnormal
thickness of the leaf caused by the presence of the fungus
is shown at GH. An embedded unripe spermogonium is
shown at J, and an organ of transpiration at K.
From a large number of observations made with the
view to trace the origin of the ^cidium cups and the
spermogonia in the barberry, we believe it takes place
in the following manner : — If sections are repeatedly
taken through affected barberry leaves it will be seen that
numerous minute granules and extremely small disjointed
fragments of mycelium may be seen in the intercellular
spaces. The fragments and granules vanish by degrees into
the finest dust at one end of the series, and appear as short
threads at the other and growing end. As the disease
advances the granules extend in growth, and appear as
fine yellowish-orange tubular mycelial threads furnished
with septa. This mycelium has a natural tendency to
grow towards both surfaces of the leaf, the growth becom-
ing more profuse as the surfaces are neared. When the
mycelium has reached the epidermal cells, under and
upper, it forms minute compact knots ; the upper knots at
length become the spermogonia, and the lower the dEcidium
cups.
At Fig. 84 is represented a section through an JEcidium
cup, enlarged 150 diameters. The lower epidermis of the
leaf is shown at AB, ruptured by the fungus from within
at CD. The fine septate, almost granular, mycelium,
from which the fungus springs, is shown at the top of the
illustration at D, and creeping amongst the intercellular
spaces at E. The outer coat, or peridium, of the cup,
consisting of a single stratum of transparent polyhedral
XXIV.]
BARBERRY BLIGHT.
163
investing cells, is shown at FF, and the sulphur-yellow
spores hanging in chains from the open cup at G. As the
spores composing these chains drop away into the air, others
are produced by a continued growth from the bed of fine
mycelium or spawn at D. The shading indicates the
FIG. 84.
Section through a cup of JEcidium Berberidis, Pers.
Enlarged 150 diameters.
crimson corrosion of the cells of the barberry leaf caused
by the presence of the embedded JSridium cup, and the
constituent cells of the leaf are shown at HH.
The growth of the spermogonia is not quite the same
164 DISEASES OF FIELD & GARDEN CROPS. [CH.
with that of the dScidium cups. The mycelium gradually
approaches the upper epidermal cells by the intercellu-
lar spaces. When it is very near the surface it forms
minute knots similar with the knots of the AScidium
cups, but the growth differs in an important particular ;
for whereas the JEcidium cup grows from the base, as at
D, Fig. 84, the spermogonium grows by a constant pro-
trusion of new threads from the circumference to the
centre. A spermogonium, even when almost mature, is
like a brownish circular ball, with a loose enclosing mesh
of mycelial threads, and from this bark-like mesh innumer-
able rays of mycelium reach to the centre, the central spot
being darker than the rays. The name, spermogonium,
means a flask or case containing spermatia, and spermatia
are extremely minute bodies with a function supposed to
be similar with that of pollen.
A ripe spermogonium is illustrated at Fig. 85, enlarged
300 diameters. It must be particularly noted here that this
spermogonium is magnified to twice the scale of its accom-
panying jEcidium cup, Fig. 84. This is necessary in
order to show the much finer details of all its parts. The
first thing to be noticed by the reader is that the constitu-
ent cells of the leaf, as at AA, although magnified twice
as many times as the similar cells at HH, Fig. 84, are
not nearly so large as the latter. The explanation of this
is, that the cells belonging to the upper part of the leaf
where the spermogonia grow are very much smaller in size
and much more closely compacted together than the cells
belonging to the lower and looser portion of the leaf.
The upper epidermal cells of a barberry leaf are shown
at BC, Fig. 85, and burst apart at DE. The spermogonium
has no true wall or bark, but its entire outer surface is a
woven coating of extremely attenuated, brownish, some-
times almost granular mycelium ; this mycelium can be
traced into the adjoining intercellular spaces of the leaf,
as at FG. The illustration shows the mycelium belong-
ing to the outer surface of the immersed spermogonium
xxiv.] BARBERRY BLIGHT. 165
growing in almost straight lines towards the centre. When
maturity is reached the spermogonium opens at the top,
and the contents burst through the epidermis of the bar-
berry leaf, as shown. At the time of bursting, the little
dark central mass turns up towards the burst point, and
the growth of the spermogone threads is continued through
the orifice, as shown at HH. The septate spermogone
threads, after they have reached the air, break up into
x-300
FIG. 85.
Section through a Spermogonium of Mcidium Berberidis, Pers.
Enlarged 300 diameters.
extremely small granules, and these granules are the
spermatia of botanists. They are supposed to be 1
grains belonging to a male organism, roughly answering
to the pollen of flowering plants. Most of the illustrations
of JEcidium spermogonia hitherto published are incorrect.
After the ^cidium cups and spermogonia are once
formed, most of the mycelium amongst the cells of the leaf,
166 DISEASES OF FIELD & GARDEN CROPS. [CH.
from which the cups and spermogonia arose, breaks up
into the finest conceivable dust, or dissolves away, and so
is lost to sight.
A single dEcidium spore is enlarged to 1000 diameters
at Fig. 86. To the spore are attached four of the so-called
- 1000-
FIG. 86.
Germinating spore of sEcidium Berberidis, Pers.
Enlarged 1000 diameters.
spermatia from the spermogones, just as one commonly
sees pollen-grains agglutinated on to a stigma.
The spores germinate rather sluggishly on damp sur-
faces, but when seen in a state of germination a mycelial
thread is protruded in a convolute fashion from one of
about six privileged or weak points on the surface of the
spore. Into the mycelial thread the vital material from
the spore is poured.
The tardy germination of the spores seems to indicate
that they are of the nature of resting-spores, and therefore
able, under suitable conditions, of resting for a prolonged
time. It is generally believed that the spores arise from
male and female elements, and these facts indicate to some
observers that the cycle of the ^Ecidium is complete in
the production of these spores, or that no other spores are
likely to exist unless simple conidia or bud spores, as
opposed to resting or sexually-produced spores.
Of late considerable attention has been directed to the
fact of the occurrence of ^cidium Berberidis, Pers., on the
more ornamental species of barberry of our gardens and
shrubberies, and especially its growth upon Mahonia
Aquifolium, Lindl. The rare occurrence of the parasite
xxiv.] BARBERRY BLIGHT. 167
upon other barberries than the wild barberry of Britain
has long been well known. A short paper from the pen
of Mr. Charles B. Plowright, M.R.C.S., has been published
in the Proceedings of the Royal Society, No. 228, 1883.
In this paper Mr. Plowright considers he has proved by
experiment that the jflridium upon Mahonia is one con-
dition of the summer mildew of corn, Puccinia graminis,
Pers. Mr. Plowright kindly furnished us with numerous
examples of Mahonia berries on which the ^Ecidium was
growing, not only upon the exterior of the berries, but
sometimes upon the seeds exposed in berries burst by the
FIG. 87.
Berries of Mahonia Aquifolium, Lindl., invaded by dZcidium Berberidis,
Pers. Enlarged 5 diameters.
growth of the fungus. Doubtlessly the JEcidium cups grew
on the seeds after the berries were burst. Two MaJwnia
berries badly infested with the dScidium are illustrated at
Fig. 87, enlarged five diameters. The presence of the
fungus, as is usual, causes hypertrophied or greatly swollen
places on the affected part. We shall refer to Mr. Plow-
right's communication further on ; in the meantime we
point out that the parasite sometimes occurs on the leaves
and petioles, as well as the berries of Mahonias and the
other ornamental species of garden barberries.
168 DISEASES OF FIELD & GARDEN CROPS. [CH.XXIV.
Here we come to a halt similar to the one we came to
at the conclusion of the description of Puccinia graminis,
Pers. Some botanists believe that ^Ecidiam Berberidis,
Pers., is probably complete in itself, others state that the
jO/cidium spores will not enter the organs of transpiration of
barberry leaves, but will only enter the stomata of grasses,
and tli at then they produce, not an JEcidium,, but a Uredo,
viz. Uredo linearis, Pers., the first stage of the fungus of
the summer mildew of corn, — Puccinia graminis, Pers. It
will be remembered that a similar phenomenon was said
to hold good with the fungus of spring mildew of corn
and the fungus of borage blight.
We will now impartially review the evidence brought
forward on both sides of this disputed question, withhold-
ing nothing — as far as our knowledge goes — for or against
the alleged connection of Puccinia and j*Ecidium.
CHAPTER XXV.
THE POSSIBLE CONNECTION OF THE FUNGUS OF CORN
MILDEW AND THE FUNGUS OF BARBERRY BLIGHT.
IN the following review of the evidence for and against
the connection of the fungi found under the genera
Puccinia and jEcidium, it will be understood that the
remarks made in reference to the fungus of the summer
rust and mildew of corn, with its supposed ^cidium
condition on barberry bushes, applies also to the spring
rust and mildew and its supposed ^cidium on members
of the Borage family. In fact, the following notes apply
to every instance where there is a supposed connection
between Puccinia and its allied genera on the one hand,
with ^Ecidium and its allied genera on the other.
First we will give a brief account of the popular belief
which is said to have existed amongst rustics in old times,
then we will give the evidence brought forward by some
men of science as supposed proofs of an actual connection
of corn mildews with the blights of barberry and borage ;
and, lastly, we will state the reasons why other men of
science in some instances reject the evidence of the sup-
posed connection, and in other cases consider the connec-
tion as unproven. As each of these branches of the
subject are reviewed we will state how our mind has been
impressed by the statements. The readers of this chapter
may then form their own opinion as to the value of the
facts as well as of the deductions which have been either
reasonably or unreasonably drawn from them.
The first we hear in regard to the connection of corn
170 DISEASES OF FIELD & GARDEN CROPS. [en.
mildew and the blight of barberry bushes is a popular
belief, said to have been common amongst rustics in old
times, that barberry bushes blighted corn. Now, although
popular beliefs peculiar to rustics of the last century need
not be altogether disregarded, yet we are inclined to put
a low estimate upon them. The rustics of the last century
did not always fix on the barberry bush as a supposed cause
of corn mildew, for in some districts old hawthorn bushes
were believed to be the cause; and to this day the labourers
of some parts of the eastern counties, as in the Hardingham
district, believe hawthorn bushes to be the cause, or, if not
the sole cause, to be at least equally pernicious to corn with
the barberry bush itself. The barberry is not everywhere
considered by rustics to be capable of causing corn mildew.
In some districts the barberry is said to be the cause of
Bunt, a disease of corn described further on in this work.
This belief was at one time very prevalent on the Conti-
nent, and is described by Phillipar in his TraitS Organo-
graphique et Pliysiologico-Agricole sur la carie, le Charbon,
VErgot, la Rouille, et autres Maladies du wdme genre qui
ravagent les Cdre'ales : Versailles, 1837. The division of
opinion amongst rustics appears to us to militate against
the acceptance of one particular view and the rejection of
the others. Eustics of the last century were very super-
stitious, and the farm labourer whose family was destroyed
by ergot in the last century (referred to in this work
under " Ergot ") would not believe that bad wheat was the
cause of the limbs of his family rotting off, but insisted
on the cause being witchcraft. No one at the present day
would consider the exploded idea of witchcraft supported
by this old belief of rustics, and we are inclined to give
no better credence to the ideas of rustics as to the connec-
tion of corn mildew and bunt with barberry and thorn
bushes. The farm labourers possibly noticed that the
colour of the fungus of rust of corn and that of the fungus
of barberry blight was the same, and this may have led
them to connect the two. When the barberry is in flower
xxv.j CORN MILDEW AND BARBERRY BLIGHT. 171
it gives out an unpleasant odour ; and as the fungus of bunt
also gives out an offensive scent, it is quite possible that the
similar strong odour belonging to both parasites at one time
led rustics to connect barberry blight and bunt together.
In past times rustics not only believed in witchcraft
and magic, but they, as we all know, had strange beliefs
regarding the influence of the moon on the weather, and
other old beliefs which now prevail to a less extent. No
one would now think of bringing forward these absurd
old beliefs in support of the more exact astronomical and
meteorological knowledge of recent times.
In the agricultural journals of the last hundred years
many curious observations may be found from the pens
of sharp observers, who really thought the connection of
corn mildew and barberry blight to be proved. Other
observers, however, and equally sharp ones, have brought
forward evidence of an entirely opposite character ; so
that nothing has been proved from old experience either
for or against the connection. Mr. C. B. Plowright, in a
valuable paper published in the Gardener^ Chronicle for
19th August 1882, has given all the popular evidence
that he could collect in favour of the barberry being in-
jurious to corn ; but this gentleman has not referred with
equal fulness to the popular evidence that told the other
way. Prof. J. S. Henslow, writing in the Journal of the
Royal Agricultural Society in 1841, vol. ii. p. 13, said
that practical men were by no means unanimous in de-
nouncing the barberry. An experienced cultivator of
Hamburgh is referred to in that paper, who, after observa-
tions made for thirty-one years, expressly contradicted the
commonly-received opinion. Experiments are recorded
by Professor Henslow which were made at Copenhagen.
Wheat was there planted and surrounded by barberry
bushes without obtaining any mildew. A similar experi-
ment was made by Jussieu in the garden of Trianon with
a like result. Mr. Knight also only obtained a negative
result with experiments of the same class. Mr. Henslow
172 DISEASES OF FIELD & GARDEN CROPS. [en.
states that lie only knew of a solitary instance of barberry
bushes growing near corn, and there the corn was worse
mildewed than elsewhere, not because of the juxtaposition
of the barberries, but because the bushes were at a corner
of the field where the soil was decidedly the worst, and
where the corn was sheltered by lofty trees. Other
observers, and equally reliable, have given evidence of a
positively opposite character, — one agriculturalist going so
far as to state that he had never even seen corn growing
near a barberry bush without its being injured more or
less. Phillipar, in the work already quoted, says, in
reference to corn mildew and barberry blight, that he was
never able to meet with sufficiently conclusive evidence
for a conviction of the barberry. On the contrary, he
states that he has seen many instances in which hedges
were filled with barberry bushes, without the corn, which
Was near them, having sustained the slightest injury.
Nothing can be made of the old agricultural evidence,
for what is stated by one side is soon after flatly contra-
dicted by the other.
Sir Joseph Banks was one of the first to suggest that
the yellow fungus of the barberry might be another form
of the rust fungus of corn, but this was only a guess on
Sir Joseph's part, founded on a popular belief. A very
early reference to the supposed injurious effect of the
barberry on corn may be found in Krunit/'s Encyclo-
pedia, 1774.
Mr. "Win. Carruthers, F.R.S., in his paper on wheat
mildew published in the Journal of the Royal Agricultural
Society, ser. 2, vol. xviii., part, ii., p. 495, brings forward
an instance (quoted by Professor Henslow) in favour of the
connection of corn mildew and barberry blight ; but he
omits the opposing evidence as printed in the same paper.
In the particular instance quoted by Mr. Carruthers the
cornfield was bounded in one part by a " young and
healthy quickset hedge ; " here it is presumed there was
little or no mildew ; but in another part of the field there
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 173
was an "old hedge with several barberry and other
bushes and some elm trees." The account continues :
"It is true the current of air was somewhat impeded
by the trees." In Mr. Carruthers' paper this important
sentence is not given, although, in our opinion, the old
hedge, with its variety of bushes and elm trees impeding
the air, is quite sufficient to greatly favour a growth of
mildew, and by no means favours the idea that the fungus
of barberry blight and corn mildew are genetically con-
nected.
We acknowledge that an opinion expressed by Sir
Joseph Banks or Professor J. S. Henslow still carries
more weight than the belief of rustics in the last century ;
but it must be remembered that in the time of Sir Joseph
Banks and Professor Henslow very little indeed was known
of the anatomy or physiology of fungi, — so little, indeed,
that any opinion the two gentlemen above Darned may
have expressed on this subject can only be held in slight
esteem now. Men of science of the present day do not
generally try to support their views by quoting what other
observers thought one or two hundred years ago, particu-
larly when those observers were not specialists. Old
observers were doubtless right in many of their ideas,
but no support is given to modern views by quoting the
opinions of old authors who were but poorly acquainted
with their subject. This course is never taken with geo-
logy or zoology — then why with the most difficult and
least understood section of botany ?
We consider, then, that this part of the subject has
been weakened by the attempts of some modern writers
to support their opinions by. quoting the views of rustics
in the last century, and by printing the old ideas of Sir
Joseph Banks and Professor J. S. Henslow about fungi
with which they were confessedly but very imperfectly
acquainted. This part of the subject is indeed, in our
opinion, hardly worth consideration. Mr. C. B. Plow-
right, however, has gone further, and republished a Pro-
174 DISEASES OF FIELD & GARDEN CROPS. [CH.
vince Law of Massachusetts of 1738-61, in which it is said
that the " blasting of wheat and other English grain is
often occasioned by barberry bushes." In the intro-
ductory remarks written by Mr. Plowright to this Pro-
vince Law, he correctly says of corn mildew that germs
of the disease "will have a greater chance of gaining
admission into the interior of the wheat plants in those
parts of the field where the influence of currents of air is
least felt." The Massachusetts law enacted that all bar-
berry bushes should be extirpated, as they have now
virtually been extirpated in Britain. As far as is known,
this " extirpation " of barberries, even when enforced by
law, has not had the least tendency to lessen attacks of
corn mildew. Surely the quotation of an obsolete and,
as considered by many persons, a stupid old law, will not
convince disbelievers in the connection of corn mildew
and barberry blight that they are wrong. We might
quote other old laws, such as the one regarding witchcraft,
and refer to Matthew Hopkins, the professional and official
witch-finder, who in the years 1644, 1645, and 1646, as
recorded in Godwin's Lives of the Necromancers, pp. 434,
435, caused sixteen innocent persons to be hanged at
Yarmouth in Norfolk, fifteen at Chelmsford, and sixty at
various places in the county of Suffolk.
In approaching modern times we come to much more
exact and searching evidence, and this modern work is
really the only part of the subject worthy of serious
attention. We have been informed by Mr. J. L. Jensen,
of Copenhagen, through Mr. C. B. Plowright, that the first
person who instituted scientific experiments with the
fungus of corn and the fungus of barberry blight was a
Danish schoolmaster named Schoeler, who lived at Ham-
mel, in Denmark, and who, seventy years ago, placed the
fungus of barberry blight on rye, and produced the rust
fungus. Bonninghausen in 1819 experimented with the
spores of the fungus of barberry blight by applying them
to rye. In five or six days the rust fungus is said to
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 175
have appeared on the rye. The probability of the occur-
rence of an alternation of generations on fungi was sug-
gested by the Kev. M. J. Berkeley in the Journal of the
Royal Horticultural Society for 1848, where he wrote, in
describing the "Bunt" fungus of wheat, "It is quite
possible in plants, as well as in the lower animals, there
may be an alternation of generations." In 1865 Professor
A. de Bary, of Strasbourg, published an essay- — Monats-
hericht der Koniglichen Preuss. AJcademie der Wissenschaften
zu Berlin, Jan. 1865 — in which he stated that he had arti-
ficially produced the rust of wheat by placing the spores
of the barberry fungus on corn. One would have natur-
ally thought that Professor de Bary was led to make this
experiment by his knowledge of the popular belief as to
the supposed connection of the two fungi; but we are
told by Mr. Plowright in the Gardeners' Chronicle for 30th
July 1883 that this was not the case, but that Professor
de Bary had previously experimented with a Puccinia,
named P. tragopogonis, Corda, and that the little pro-my-
celium spores produced by the germinating teleutospores
of the Puccinia, when sown on a healthy host-plant, did
not produce a Uredo, but an ^Ecidium named JE. tragopo-
gonis, Pers., and that when the dEcidium spores were
planted on another host-plant, they in turn produced a
Uredo. We have in this country an abundance of
sEcidium tragopogonis, Pers., but Puccinia tragopogonis,
Corda, is unknown. Professor de Bary, then (says Mr.
Plowright), selected the barberry " because experience had
taught the practical farmer that it was prejudicial to the
wheat crop." This admission appears to be identical with
the one suggested by us, that the experiments were under-
taken in consequence of a popular belief amongst a certain
number of farmers and their labourers. Professor de Bary
appears to have been unacquainted with the admirable
paper by Professor J. S. Henslow in voL ii. of the Journal
of the Royal Agricultural Society for 1 84 1 , where the identity
of rust or Uredo and mildew or Pucciniawas first pointed out.
176 DISEASES OF FIELD & GARDEN CROPS. [en.
Since Professor De Bary's paper was published, other
good observers have made experiments, some with a result
pointing in one direction, some with a result pointing in
the opposite one, and others with a negative result. Many
botanists hold that the case is proved, and that the con-
nection of the two fungi is certain ; others, as Dr. M. C.
Cooke, one of the foremost fungologists of this country,
still hold the connection of the two parasites as unproven.
The latest exponent of the connection of Uredo with its
Puccinia and j*Ecidium is our friend Mr. C. B. Plowright,
M.R.C.S. This gentleman has made two sets of experi-
ments. In the first his results were negative, or, as he
himself says, they seemed to show that the barberry fungus
had very little to do with corn mildew. In the first series
of experiments in which spores of ^Ecidium Berberidis,
Pers., were placed on wheat plants, 76 per cent became
infected with rust, and amongst the wheat plants which
were kept as checks on the infected ones, no less than 70
per cent became spontaneously diseased with rust. After
his later experiments Mr. Plowright altered his first-
expressed opinion, and now he strongly advocates the
connection of the two parasites. We consider the pub-
lished change of opinion favourable to Mr. Plowright as
an observer, for it is not every one who has sufficient
independence of mind to so frankly and quickly publish
such a radical change of thought. Mr. Plowright is to be
trusted to a far greater extent than men who have advo-
cated erroneous theories, and then quietly ignored their
former teachings without acknowledging their error. Mr.
Plowright has published a series of engravings, mostly
from nature, illustrative of the subject before us. All
the illustrations are original but one, and that one is the
crucial one of the spores of the fungus of corn mildew
germinating on and sinking through the cuticle of the
barberry leaf. In this illustration (Gardeners' Chronicle,
19th August 1882, p. 233) a large open organ of trans-
piration is shown close to three germinating pro-mycelium
xxv.J CORN" MILDEW AND BARBERRY BLIGHT. 177
spores of the mildew fungus : the threads do not enter
the open stomate, but are seen boring through the cells
of the barberry leaf. This special drawing, the only
one of real interest in the paper, is not original, but a
copy from a published book. We consider it a weak
point, therefore, in Mr. Plowright's case that in his
elucidation of the only critical part of his subject he has
fallen back upon a many times copied and recopied book
illustration. How valuable a new illustration of this won-
derful phenomenon would have been ; but at present, as
far as we know, no one has ever ventured on a second
original representation. We do write in this way with a
view to throw doubt on the accuracy of the drawing.
Every one who has a microscope can easily see Uredo
spores, Puccinia spores, and JEcidium spores germinate,
and many are the original published illustrations. How
is it, we may ask, then, that so few can see pro-mycelium
spores piercing the epidermal cells of a barberry leaf, or,
if they can so see them, do not venture on new illustrations ?
It was specially necessary that Mr. Plowright should, if
possible, have published an original of this phenomenon.
As Mr. Plowright is now the chief teacher of the con-
nection of corn mildew and barberry blight in this country,
it will be interesting to record his words in reference to
infecting barberries with spores from the fungus of corn
mildew. He writes : " On 14th April, 17th April, and 9th
May, respectively, I infected one of these (barberry bushes)
with spores from the pro-mycelium of Puccinia graminis
from wheat and twitch, and kept the three remaining
barberries as control plants. In due course the jEcidium
appeared upon the infected plants, the control plants
remaining free from ufflcidium, and they continued so for two
months, when they were cut down, the experiment being
then ended." Further on Mr. Plowright writes : " On 1 5th
and 17th April T placed upon nine wheat seedlings some of
the same pro-mycelium spores which were used for infecting
the barberries, and upon 7th May one of the wheat plants
178 DISEASES OF FIELD & GARDEN CROPS. [CH.
had rust or Uredo upon it. As these plants were, however,
exposed to the air for fourteen days, an element of doubt
is admitted, although an equal number of check plants grown
in the open air in the same garden remained free from rust.
(The italics are ours.) That they did not contract the
parasite from the barberries in my garden is clear from
the fact that there were no AUcidium spores there until
many days later." Surely the evidence in. this test case
shows, if it shows anything, that Mr. Plowright really and
truly produced the Uredo from the Puccinia spores. He
admits there were no ^cidium spores ; he experimented
with germinating Puccinia spores only, on wheat, and
Uredo was the result. Bonninghausen gave the time of
the appearance of the disease after infection as five or six
days ; Mr. Plowright states twenty or twenty-two days.
The gentlemen who advocate the connection of wheat
mildews with barberries and borages bring forward the
following fact, which, it must be acknowledged, has con-
siderable weight. On repeating experiments first made on
the Continent with Puccinia and Mddium, spores one or two
species of fungi have appeared (presumably as the result of
the experiments) which had hitherto been unrecorded as
British. Undoubtedly, this fact is of importance in the
consideration of the subject ; still it must be borne in mind
that whilst fungologists are few fungi are almost without
number. No season passes but large numbers of species new
to Britain are added to our flora. In fact it seems to be
true in regard to fungi that one has only to look for certain
species in the right places and at the right time, and they
are certain to be found. Not only are numerous small
leaf fungi annually lighted on, but large species, sometimes
a foot high, as Morcliella Smithiana, Cke., by ourselves, and
Lactarius controversus, P., by our friend Dr. M'Cullough of
Abergavenny. No one supposes these large fungi did not
exist here before ; they were not seen and recorded, simply
because no one had looked for them. Prof. Elias Fries,
the illustrious Swedish botanist, wrote us, a short time
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 179
before his death, to say that he believed a large number of
his Swedish firwood fungi would be found in the firwoods
of Scotland if sought for ; but they are still but little
sought for, and therefore not in many instances found.
There has long been an extremely common fungus in this
country named ^Scidium Tussilaginis, Pers., but till lately
its supposed alternate form, Puccinia Poarum, NieL, had not
been recorded, although the host plant of this fungus is
extremely common. Mr. Plowright in his experiments
(Grevillea, voL ii. p. 56) claims to have produced this
Puccinia artificially on Poa eleven days after infection from
the germinating dfieidiwn spores ; and on close search being
made, examples of Poa annua, L., were found growing natur-
ally and bearing the Puccinia. These facts as to the artificial
production and the ultimate discovery of the naturally-
grown Puccinia have been brought forward as evidence in
favour of the connection of ^Ecidium and Puccinia. We
are inclined to give but little weight to this part of the
evidence, for the j^Ecidium still remains extremely com-
mon and the Puccinia extremely rare, just as in the common
Uredo Rubigo-vera, D.C., and the excessively rare ^Ecidium
asperifolii, Pers., described in this work under Spring Rust.
The believers in the connection of corn mildew and
barberry blight consider their views supported, and indeed
proved, by their experiments. The results of the experi-
ments leave no room they say for doubt ; still a great diffi-
culty rests with, all the experiments in regard to the lapse
of time which takes place between the application of the
infecting spores and the appearance of the fungus which is
supposed as a consequence to follow ; this period is some-
times more than twenty days. In the case of jEcidium
bellidis, D.C., as reported in the Journal of the Linnean
Society, vol. xx. p. 512, the time ranged from twenty-
four days to over two months. When a germ tube enters
the tissues of a plant through one of its organs of transpir-
ation, no one can follow it farther. Every microscopist
knows that a leaf is an opaque object, and no amount of
180 DISEASES OF FIELD & GARDEN CROPS. [CH.
artificial light will show what has become of a germ tube
from a fungus spore when it has once travelled down
amongst the constituent cells of the leaf. If a germ tube
of any given fungus is known to have entered a leaf, and
a fungus of a totally different nature appears in eight days
or two months afterwards upon the surface of the invaded
leaf, where is the clear proof that the foreign germ tube
really caused the production of the new fungus-growth ?
It is certainly not impossible that one may have arisen
from the other, but the proofs of a phenomenon so won-
derful should be unimpeachable, — proofs such as no one
could possibly doubt or question.
The believers in the connection of corn mildew with
barberries always recommend the destruction of barberries
as a preventive of corn mildew, and Mr. Carruthers, in
the paper already adverted to, writes : " The farmer should
not permit the barberry to have a place in his hedges or
in plantations on his farm." At one time barberries were
abundant in Britain, now they are very rare in a wild
state, and as a rule only to be seen in single isolated
examples. As the former extirpation of the plant has not
lessened the mildew of corn in the slightest degree, why
then should the remaining few barberry bushes be
destroyed, especially when the case of Australia and
New Zealand, where, with a total absence of native bar-
berries, corn mildew is worse than in Europe, is remem-
bered ? In our own country corn mildew is notoriously
at its worst in the fen districts, where the barberry is
absent in a wild state. The advocates of the connection
of the two fungi acknowledge that mildew is perennial in
corn ; being so, the ^Ecidium condition on barberries,
even if admitted as a condition of corn mildew, cannot
be a necessary condition. With the destructive fungus of
spring mildew of corn and its supposed jtEcidium on
borages, the case is still more striking. There is no
need to tell farmers not to allow any members of the
Borage family to have a place in their gardens ; it does
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 181
not matter what number or variety of borages they grow,
for however large the number may be, they are almost
sure to have no dEcidium upon them. dEcidium asperi-
folii, Pers., is so rare in Britain that during a thirty years'
study of fungi we have never once met with it. On
inquiring, as to the prevalence of this fungus in Britain,
of our friend Dr. M. C. Cooke, he replied that he had
carefully sought for it, but had only found it about two or
three times in twenty years. Mr. C. B. Plowright, who
wished to experiment with it, could not get it here at all,
and the illustrations in this work, Fig. 70 and Fig. 71,
were taken from a dead foreign example, simply because
no British or living specimen could be secured. Some
persons may think from this, that spring rust of corn
derives but scant benefit from a possible connection with
a fungus of such extraordinary rarity as ^Ecidium asperi-
folii, Pers. It is clear that the rust can keep in existence
for an indefinite time without any aid from the JEcidium.
Puccinia JRubigo-vera, D.C., and P. graminis, Pers., are
very close allies, and both grow upon the Graminece or
grasses ; one would have expected, therefore, that their
alternate ^Ecidium forms would both grow on one plant
or set of allied plants. We have seen, however, that
such is said not to be the case, as the Berberidacece and
Boraginacece are widely separated.
Notwithstanding these objections we are not inclined to
attach undue importance to the absence of dEcidium Ber-
beridis, Pers., in Australia, or to the extreme rarity of
^cidium asperifolii, Pers., here. The spores of these fungi
may possibly be more common in some other country, and
be carried through the air from one place to another.
Clouds of aphides and ladybirds have sometimes been seen
darkening the air and travelling towards Britain across
the sea from France, and spores may travel in the same
way or be carried by insects, birds, or man. We do not,
however, consider such a transport of spores, especially
across the ocean to Australia, probable, though not per-
182 DISEASES OF FIELD & GARDEN CROPS. [OH.
haps impossible ; and there is no proof that anything of
the sort has ever occurred.
We have little doubt that the mycelium of both the
mildew fungi and the spawn of both the ^cidia are peren-
nial, and that both can live on from year to year for an
indefinite period without aid from each other. Mr. Berke-
ley, in vol. i. of the Journal of the Royal Horticultural Society,
writes at p. 25 : " The mycelium of cereal fungi is known
to exist from the earliest period in corn ;" and further on
he remarks that " a diseased stock can scarcely be expected
to produce a perfectly healthy offspring," and " it is cer-
tain that the germs of cryptogamic plants may be present
in tissues, and yet remain more or less inert." Referring
to dEcidium, he says, in reference to the parasite of the
anemone (Introduction to Cryptogamic Botany, p. 323), " the
leaves, which will be eventually covered with the fungus,
show that they are impregnated with its mycelium as
soon as they make their appearance." If these facts are
admitted, and ^cidium quadrifidum, D.C., of the anemone
is acknowledged to be perennial, another point arises. How
can any observer tell that the plants he is experimenting
with have not the germs of disease already in their tissues ?
We, as well as many other observers, have shown that seeds
apparently sound will often, on germination, show disease
in their seed leaves ; such plants are saturated with the
germs of disease from their earliest period of growth.
An instance was adverted to by us in the Gardeners'
Chronicle for 26th January 1884, p. 120, where a well-
known nurseryman in a large way of business had im-
ported Dianthus seeds direct from Japan. These seeds
were carefully grown under glass, and, immediately they
were up in the seed-pans, they were all attacked and
destroyed by Puccinia lychnidearum, Link. On making a
microscopical examination of a series of these seeds we
detected mycelium inside the integument which surrounds
the embryo or infant plant and within the coat of the seed.
Dr. M. 0. Cooke has published a case where seeds
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 183
gathered from undiseased celery plants produced healthy
plants ; whereas seeds taken from celery plants diseased
with Puccinia Apii, Corda, produced seedlings every one
badly infested with the Puccinia. The plants were grown
in rows side by side in the same garden, and the clean
plants remained healthy all the season ; whereas the
diseased ones were destroyed by the hereditary disease
derived from the parent plants and presumably conveyed
from the parent to the offspring in the seeds.
The Kev. M. J. Berkeley has published in the Gardener^
Chronicle, 28th October 1848, p. 716, an instance of plants
of Pyracantha raised from seeds imported from Russia
being all killed by a species of Fusicladium ; whilst old
plants of Pyracantha growing at the same place remained
perfectly free from disease. The same gentleman records
an instance of a plant of Achillea Ptarmica, L., being given
to him by M. Desmazieres. When presented it was appar-
ently quite free from disease, but the donor knew that the
disease plasma of Labrella ptarmica, Desm., was in its
tissues. The Achillea was planted in March, and in the
following autumn the Labrella duly appeared, although
the fungus up to that time had not been seen in Britain.
It is common to find hollyhock seedlings showing the
Puccinia on their seed leaves. This we have traced to
the presence of pustules of the disease outside the seeds
or carpels, as illustrated by us in the Gardeners* Chronicle
for 1st July 1882, p. 23. Similar pustules occur on the
carpels or seeds of wild mallows.
Many similar instances might be given ; they all prove
that Puccinia on mildew is hereditary, — that it exists in a
finely-attenuated state in seeds taken from diseased plants,
and can be transmitted in a long interminable line from
generation to generation. No doubt it is possible that
living spores or mycelium may sometimes be present out-
side the seeds, but many fungi are able to reach the seeds,
as the fungus of bunt in corn, Tilletia Caries, Tul., and
Thecaphora within the carpels of convolvulus, etc.
184 DISEASES OF FIELD & GARDEN CROPS. [CH.
We believe we have seen instances of the spawn of
the ^Ecidium being perennial in the barberry. The
mycelium which gives rise to rust and the mycelium
from which ^Ecidium arises hardly appear to us to be of
the same nature. Attention has lately been redirected to
the occasional growth of j&Scidiwn Berberidis, Pers., on gar-
den barberries, especially on Mahonia Aquifolium, Lind.
It has been suggested that there may be garden barberries
in Australia, and that these plants may be infested with
the ^Ecidium said to belong to corn mildew. This may
possibly be correct, although no evidence has at present
been forthcoming in that direction. If garden mahonias
are found to bear JEcidium in Australia we imagine the
number of diseased garden mahonias on that large Conti-
nent will be in about the same proportion to the vast corn-
fields there found, as the borage dEcidium is to the corn-
fields of Britain, or as the fungus of Poo, annua, L., is to
the dEcidium of coltsfoot and butter-bur.
Mr. Charles B. Plowright was kind enough to send us
a good collection of Mahonia berries invaded by jtEcidium,
as illustrated in Fig. 87. Occasionally the disease ap-
peared upon the leaves and stalks, and from our examin-
ation of them we were inclined to think the mycelium of
the fungus traversed the entire plant, and especially the
berry with its seed. We successively planted a number
of seeds at overlapping intervals through last summer,
but not one germinated. This suggested to us that the
plasma of the fungus had reached the embryos of the seeds
and killed them. On inquiring of Mr. B. S. Williams,
the well-known nurseryman of Upper Holloway, he
informed us that Mahonia seeds germinate freely when
the seed has been matured, is good, and properly planted.
Mr. Williams said Mahonia seeds took from two to three
months to germinate, and he kindly undertook to test
twelve ripe berries for us, each berry showing the
^-Ecidium disease. The seeds were sown carefully in two
pots, and after the proper time had elapsed no single seed
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 185
showed the least sign of germination, but all had perished.
These facts, of course, do not prove that the mycelium is
perennial, but we think it proves that the spawn can reacli
the embryo of seeds, and in bad cases kill them. We also
think it suggests the possibility of seeds less badly diseased
being able to give rise to diseased seedlings, exactly as in the
case of the Dianthus seeds, which showed the Puccinia in
the seed-leaves. In the Proceedings of the Royal Society,
No. 228, 1883, Mr. Plowright has illustrated the ^Ecidio-
spores belonging to Mahonia Aquifolium, Lind., germinating
upon the epidermis of a fragment of wheat-leaf. Six
spores are shown : five are germinating — three germinal
threads are entering the stomata; but no less than seventeen
little branches of the mycelium are shown naturally and
correctly running along the little furrows belonging to
the junctions of the cells which form the leaf cuticle.
The illustration simply proves that the germ-tubes will
run anywhere where there is a depression, or run into
any little orifice. After eleven days, not five or six as
with Bonninghausen, Uredo appeared upon the plants
experimented upon.
Many cases are well known where host plants are always
so saturated with parasitic disease that it is almost impos-
sible to find the host without the parasite. Pythium
equiseti, Sdbk., is so common on Equisetum that we have
seldom found an Equisetum without it. Cress seedlings
are plagued in a similar fashion with another Pythium.
Now, who would place any reliance on experiments made
with a view to inoculate Equisetum and cress with Pythium,
when it is well-known beforehand that every plant is
probably already permeated with the virus of the parasite
in a latent state ?
Personally we do not esteem the fact of spores germi-
nating on the cuticle of a leaf as of the slightest value.
Nearly all spores will germinate in warm moist air upon
any surface, and the spawn-threads will run into any slight
depression, or any orifice, provided it is large enough.
186 DISEASES OF FIELD & GARDEN CROPS. [OH.
Germinating spores of Uredo, Puccinia, and dEcidium tra-
verse the scratches made by a lancet on our glass slides just
as naturally as if they were the furrows of a leaf-cuticle.
When the threads get to the edge of the glasses they dip
down just as naturally as if the vacant space were an
open organ of transpiration. We have grown spores on
moist linen, calico, and blotting-paper, and the germ-tubes
have penetrated between the orifices and run over the
reverse side just as if they were in a leaf. Such spores,
unless of moulds, of course do not reproduce a perfect
fungus like the one from which they originally arose ;
neither do those sown on leaves. Professor De Bary says
he could not cause jEcidium spores to grow effectually on
barberry leaves. It is only after a considerable time ha.s
passed, and the germ-threads have been lost to sight for
many days, several weeks or months, that some new fungus
of an apparently different nature at length appears.
One writer has said that botanists were prepared to
accept the idea of Puccinia and dScidium being one and
the same fungus because they were acquainted with the
changes of some insects such as are familiar in the cater-
pillar, chrysalis, and perfect butterfly in the insect world.
We confess that we do not see the resemblance at all,
No condition of the insect is ever lost to sight for ten or
twelve days or two months, and the change is gradual
throughout from one form to the other ; the chrysalis is
foreshadowed in the caterpillar, and the perfect butterfly
has all its parts in the chrysalis. No organisms belong-
ing to Puccinia are ever indicated by ^Ecidium, and no
analogues of Spermogones or ^Ecidiospores are ever met
with in Puccinia.
It has been stated by Professor De Bary that an ana-
logous case of the change of host exists in the animal
kingdom, as in the case of the entozoic worms, which pass
the first part of their existence in one animal, and the
second part in another and totally different animal. The
experiments with entozoic animals have doubtlessly proved
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 187
this fact ; but to compare the facts belonging to the
animals with those belonging to the plants is a case of
mistaken analogy, for the facts do not correspond.
In the first place, the test experiments with Entozoa
have invariably been made with host animals that were
perfectly well known to be quite free from entozoic parasites.
Measures have always been taken to make this fact certain
before the experiments were commenced. The two series of
animals have in no single instance been notoriously infested
with parasites before the experiments were entered upon.
The comparison that has been made between the
change of host in the Entozoa, and the supposed similar
change of host in the two parasitic fungi of spring and
summer mildew of corn, brings us to what we think may
prove a fatal objection to the connection of Puccinia with
^Ecidium, and one that strikes at the root of the whole
hypothesis. In the entozoic animals referred to, one form
of the parasite, say of the liver-fluke of sheep, is sexually
mature in a certain mammal. These sexual individuals,
which produce thousands of eggs, escape by the alimentary
canal of the invaded animal. The eggs are dispersed by
wind, rain, insects, feet of cattle, and other means, and so
at length find their way into pools, ponds, and streams.
The mature eggs contain ciliated embryos, which are set
free on moist surfaces or in water. Each embryo contains
a bud which at length becomes a larva. The ciliated
embryo attaches itself to a second host, such as a snail,
slug, or aquatic insect, and so gains access to the interior
of the new host. In this position it becomes a non-sexual
larva, and capable of producing a progeny, or other larvae,
within itself. These secondary larvse migrate from the
bodies of the water insects or molluscs and become free ;
they are then swallowed by mammals whilst eating or
drinking. The larvae now bore through the tissues of the
new host, and enter into a pupa stage whilst within the
mammal, and there at length become sexually mature
egg-producers, and so the life, cycle becomes complete.
188 DISEASES OF FIELD & GARDEN CROPS. [OH.
Now we will note liow the phenomena connected with
^Ecidium and Puccinia agree or disagree with the entozoic
life-history just given. We commence with the dBtidium.
Here we have a sexually mature parasite, with so-called
spermogonia and spermatia. The names indicate male
organs, with a male fertilising element analogous wTith
pollen. Mr. Plowright, in writing of the spermogonia
and spermatia, says : " Their function has not as yet
been absolutely demonstrated, but there is little doubt
that they play the part of the male element." It is
necessary to be exact at this point, for if there is a sexually
perfect state of the parasite it must of course be either in
the dScidium or the Puccinia ; and if the ^cidium breaks
down, the whole hypothesis falls to pieces as far as pub-
lished descriptions go.
We believe the ^Ecidium to be sexually perfect, as
indicated by the descriptive terms in general use. The
male organs or spermogones are usually, if not invariably
produced first, and the sEcidia next ; this phenomenon
roughly agrees with the sequence of the stamens and
pistils in flowering plants. In Endophyllum and Roestelia
the spores resemble the oospores of a Peronospora. We
believe that the spores with spermatia attached, as illus-
trated in Fig. 86, agree with fertilised ovules. These
fertilised ovules, if they agree in habit with the ova of
Entozoa, should produce a simple larval form, which should
reach the interior of some other plant and there live para-
sitically as a larva. The spore of the sEcidium is supposed
to so reach the leaves of corn, and to travel to the interior
by the stomata. In this position we presume it resembles
the non- sexual larva inside the snail. In the entozoic
animals in question the larva never reaches any higher
stage in its host, the snail : it attains its pupa and perfectly
sexual state when it again reaches the mammal. Here
the comparison appears to break down ; neither the
Uredo or Puccinia can be larval. A larva in fungi can
only be some simple conidioid form like the Oidium of
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 189
grass mildew — some simple, budding, non-persistent, non-
sexual form.
The Uredo and the Puccinia which follow the jfflcidmm
are the reverse of simple, the Puccinia being specially
complex and persistent. The Uredo is surely the pupa
state of the Puccinia, and Mr. Plowright himself, in
describing the teleutospores of the latter, correctly, as we
think, terms them resting -spores. Eesting- spores they
certainly are, for many of them rest for nearly a year before
germination takes place. They are as truly analogues of
ova or eggs as are ^Ecidiospores. If Puccinia graminis,
Pers., is the larval state of ^Ecidium Berberidis, Pers., it is
at least curious that the larvae should be so extremely and
continuously prolific in Australia, whilst the sexually mature
form is restricted to the other quarters of the globe.
If the teleutospores of Puccinia are resting-spores or
analogues of oospores, as they probably are, the spores of
dEcidium cannot be of the same nature without there
being two sexually mature egg -producing forms in the
same life cycle of the fungus. We lately addressed a
question to Dr. T. S. Cobbold, F.R.S., probably our
highest living authority on Entozoa, and asked if he
knew of any instance in the animal kingdom of a parasite
passing two stages of its existence in two different ani-
mals, and arising from two different forms of eggs. Dr.
Cobbold replied at once : " I give a distinct negative to
your question, without prejudice to the proven fact of
dimorphism amongst parasites ; and of course also without
the slightest reliance on the authority of Meguin, whose
erroneous views imply the belief that two sorts of eggs
may belong to parasitic conditions of one species." It
appears from this answer that the erroneous views of
Meguin may be comparable with the views held by many
botanists in regard to Puccinia and dStidiwn.
The change of host plants in fungi has been technically
termed hetercecia and metcecia, from heteros, diverse or
variable, and meta, a change.
190 DISEASES OF FIELD & GARDEN CROPS. [CH.
The accompanying illustration, Fig. 88, is copied from
Tulasne's paper in the Ann. des Sc. Nat., 4 ser., vol. ii.,
1854, pi. 9, and represents at A the germination of a
spore of Uromyces appendiculatus, Lev., and at E the ger-
mination of a spore of JEcidium Euphorbice-sylvaticce, D. C.
PIG. 88.
Spores of Uromyces appendiculatiis, Lev., and JEcidium Euphorbice-sylvaticce,
D.C., germinating and producing pro-mycelium, pro-mycelium spores,
and sporidioles of identical character. After Tulasne.
The illustration is copied to show that, according to
Tulasne, not only is Puccinia capable of producing pro-
mycelium and pro-mycelium spores, as shown in Figs. 69
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 191
and 8 1 in this work, but that Uromyces and even ^Ecidium
itself are both potentially capable of giving rise to precisely
the same growths. The pro -mycelium and pro -mycelium
spores of Uromyces are shown at B and C, and the same
growths belonging to the dScidium at F and G. The
phenomena are also exactly the same in regard to the
habit of pro-mycelium spores ; one belonging to Puccinia
graminis, Pers., as illustrated by Tulasne, is shown germi-
nating and producing a sporidiole (as Tulasne terms it) at
D, and a pro-mycelium spore of ^Ecidium is producing a
sporidiole of precisely the same class at H.
It may be well here to glance at the development of the
teleutospores in Puccinia. We will take P. Rubigo-vera,
D.C., as an example. The teleutospores, which are preceded
by simple Uredo spores, first appear as short stalks capped
by a small cell, as at A, Fig. 89, enlarged 500 diameters.
At first the young teleutospore somewhat resembles a
Uredo spore as illustrated. Although this young teleuto-
spore looks like a Uredo spore in size and shape, it is in
reality quite different in nature. Botanists familiar with
these bodies can instantly recognise the two forms. Uredo
is probably a pupa state of Puccinia ; we do not say Puc-
cinia needs a pupa state, but that a state analogous to a
pupa stage often occurs, and is certainly present in both
the species of Puccinia which cause mildew of corn. In
an early stage of growth the young teleutospore has but
one cell- wall, and the material within is a watery fluid.
As growth progresses the spore is seen as at B, with a new
growth springing from the base within the original cell ;
this growth goes on till the spore resembles C ; the
internal mass has now formed another wall round itself
within the original wall, and in the course of a few days
the new inner cell will nearly occupy the whole of the
space within the first formed cell. At this time a third
growth appears at the base, as at D. This growth often
pushes the first inner cell aside, as shown, giving the one-
sided appearance often so common in Puccinia spores.
192 DISEASES OF FIELD & GARDEN CROPS. [CH.
As the new growth enlarges, the appearance of the teleuto-
spore resembles E, with the tipper segment dark in
colour, and the lower one light. F and G show other
conditions of this stage of growth, the lower cell pushing
the upper one aside in an irregular manner ; sometimes
the upper cell becomes large and brown, whilst the lower
one remains small and almost colourless, as at H. A per-
U H U J e^r 11 K
FIG. 89.
Development of teleutospores in Puccinia RuUgo-vera, B.C.
Enlarged 500 diameters.
fectly formed teleutospore is shown at J, and a ripe
example breaking into two portions at K. The two cells
of the teleutospore do not arise from a differentiation of
the cell contents, and the ultimate formation of a septum.
There is no septum in the outer wall of a teleutospore till
the two internal cells are mature.
We are uncertain of the exact meaning that should be
attached to the two contained cells in Puccinia spores ; but
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 193
we are inclined to look upon the upper and first-produced
dark cell as female, and the lower, often smaller, and
more transparent cell as possibly sometimes male. It is
known that the lower cell is from the first often smaller
and much more transparent than the upper one. It is
the upper one that usually produces pro -mycelium.
When both cells produce pro -mycelium, as they fre-
quently do, both are female. We believe it to be possible
that impregnation from a male element is not necessary
for every generation, but that fertile female spores may be
produced for several generations without impregnation
from a male organism. A comparable case of involved
sexuality occurs amongst molluscs. Many land snails are
monoscious — that is, each individual is male and female
in itself, and capable of fertilising itself. The character
varies in different genera. In the Valvatidce the indi-
viduals which are at first male ultimately become female.
A similar phenomenon may possibly hold good in Puccinia
and its allied genera. A comparable phenomenon is
common and well known in plant-lice or Aphides. Mr.
G. B. Buckton, F.R.S., writes that in some instances
" males occur only at such remote intervals that their
action seems to exist at a minimum."
We do not say that the involved changes advanced by
some botanists are impossible in the life-history of a single
fungus, but we have shown that they are unnecessary, as
Puccinia Rubigo-vera, D.C., can apparently reproduce
itself for an indefinite period of time in Europe without
an dEcidium condition ; Puccinia graminis, Pers., can do the
same in Australia ; the allied Podisoma Juniperi-Sabince,
Fr., does so in America. Chrysomyxa Ledi does so in
Greenland, and Puccinia obscura, Sch., in America, where
the daisy, on which its supposed alternate form is said to
grow, is not found in a wild state.
To us the pro-mycelium, pro-mycelium spores, and
sporidioles, potential in Puccinia and ^Ecidium alike, tell
strongly against the idea of the genetic connection of the
0
194 DISEASES OF FIELD & GARDEN CROPS. [en.
two fungi. One may reasonably ask which is the sexually
mature condition in Puccinia and dEcidium. If there is
any meaning in the words spermogone and spermatia,
jflcidium must be the sexually mature condition ; yet the
believers in the connection make Puccinia the perfect con-
dition, for in describing these fungi the fruit of the ^Ecidium
is invariably termed "the dEcidiospore of the Puccinia"
This puts Puccinia in the foremost position, and Sachs, in
his Handbook, Ed. i., p. 247, prints "Puccinia, graminis"
under an illustration which includes Uredo linearis, Pers.,
Puccinia graminis, Pers., and ^Ecidium Berberidis, Pers.
When the teleutospores germinate they produce the so-
called pro-mycelium spores or larval spores, as illustrated
at C and G, Fig. 88. These latter spores, when they in
turn change by germination, should give rise to the Uredo
spores or pupa state of the Puccinia. If this view is cor-
rect, we have every life-stage perfect.
1. Puccinia, the perfect resting-spore stage.
2. Pro-mycelium spores, the larva stage.
3. Uredo, the pupa stage.
What need is there for another set of spores as found in
^cidium, with their resulting pro-mycelium spores and
sporidioles of precisely the same class ?
At this stage of the inquiry the question presents itself
to us — If the ^Ecidium and Puccinia are not physiolo-
gically related to each other, how is it that Uredo,
Puccinia, and ^Ecidium, sometimes grow together on the
same host ? Our reply is : — It may possibly be a mere
state of consortium — another of the many familiar instances
of two diverse organisms being found in company. The
consortism, if such it be, would not be more remarkable
than the consortism of Peronospora parasitica, Pers., with
Cystopus candidm, Lev., on the cabbage tribe ; of Perono-
spora nivea, Ung., and Protomyces macrosporus, Ung., on
umbelliferous plants ; of Peronospora infestans, Mont., and
Fusisporium Solani, Mart., on potatoes ; of Saprolegnia
ferax, Kutz., and Empusa muscce, Cohn ; and many other
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 195
cases might be added. Another instance is met with in
the Sclerotium, which produces Peziza tuber osa, Bull ; this
is generally found in company with the black, hori-
zontal rootstock of Anemone nemorosa, L. At one time a
relationship was suspected. Plant-lice and fungi often
consort curiously together. Passarini has said that the
Aphis, named Rhopdlosiphum dianthi, Sch., gives rise to a
kind of mould on greenhouse plants which the French
name Fumagine. This is the dark-coloured fungus named
Fumago by botanists. A fungus which is extremely
common on evergreens in Britain, named Capnodium
Footii, B. and Desm., is almost invariably accompanied by
the lichen, named Strigula Babingtonii, B. A coccus is also
almost constantly present with the fungus and the lichen ;
perhaps the former, by piercing the leaves or by leaving
some secretion from its body, prepares the way for the
fungus and its companion lichen. In the same way as the
jackal is sometimes termed " the lion's provider," so the
coccus may be the provider for the Capnodium or Strigula.
When the fungus and the lichen have once fixed them-
selves on the coccus-invaded plant, the host soon dispenses
with the service of the coccus. It may possibly yet be
shown that the germinating ^cidio-spore is the provider
for the Uredo, or the germinating Puccinia-spore the
provider for the sEcidium.
Some rustics believe that mushrooms spring from salt,
because "experience has taught the practical farmer"
that a dressing of salt over a non-productive pasture will
generally cause a good crop of mushrooms to appear. The
result in this instance, however unvarying, does not prove
genetic relationship.
In some instances the occurrence of Puccinia and JEci-
dium must of necessity be a mere case of consortism, as
in the familiar example of the parasites of Allium ursinum,
L., and A. oleraceum, L. In this instance we have an
sEcidium, a Uredo, and on the Continent (but not in
Britain) a Puccinia, named P. allii, Rud., all on the
196 DISEASES OF FIELD & GARDEN CROPS. [CH.
same host plant; but the botanists, who physiologically
connect Puccinia and ^Ecidium, say the Uredo, Puccinia,
and ^Ecidium in this instance are not related, and that the
Puccinia found on Allium ursinum, L., and A. oleraceum,
L., both named P. allii, are also distinct from each
other. They say the dEcidium produces, not the Puc-
cinia on its own leaves, but the Puccinia named P.
sessilis, Sch., peculiar to Phalaris arundinacea, L. The
^cidium is abundant in Britain, but the Puccinia) which
should be equally abundant, or nearly so, if one parasite
gives rise to the other, is here extremely rare or unknown.
The JEcidium state of P. allii, Eud., is also said to be
unknown, and as the Puccinia which commonly accom-
panies it is said to be foreign to it, as no doubt it is, the
case must be a clear one of simple consortism. ^Ecidium
Berberidis, Pers., is not the only dEcidium found on the
Berberidacece. There is a second large -spored ^cidium
which grows on Berberis glauca, D.C., in Chili, in company
on the same disease spots with a Puccinia named P.
Berberidis, Mont. In Europe the same ^Ecidium possibly
grows on Berberis vulgaris, L. This fact seems to tell
against the connection of the barberry Alcidium with the
Puccinia of corn ; but in an effort to overcome the difficulty
it has been said that the spores of the Chili JEcidium are
larger than those of the true dEcidium Berberidis, Pers.,
and chiefly on this account the Chilian parasite has been
elevated to the position of a new species. Personally, we
do not estimate the size of the spores in ^cidium as
worthy of marking specific distinction. They are extremely
variable in size in undoubted examples of ^. Berberidis,
Pers. A third early gro wing ^Ecidium also occurs on Berberis
ilicifolia, Forst. This is named dEcidium magelhanicum, B.,
but its germ tubes are said not to enter the leaves of grasses.
If Puccinia and JEd&ium are physiologically related,
we have proofs that the two fungi not only arise from two
different kinds of eggs which are probably both sexually
produced, but the resulting fungi often cross consort with
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 197
each other, without being related, in the most complicated
and bewildering manner.
In the case of Puccinia violarum, Link., where the
Uredo, as well as ^Ecidium violce, Schtmi., all occur on the
same host, the three forms have been accepted as differ-
ent conditions of one parasite ; but when Mr. Vize detected
a second species of dEcidium on white garden violas, and
named it 2s. depauperans, a difficulty arose, for there were
then two ^cidia and only one Puccinia.
We dissected and illustrated the original examples as
found by Mr. Vize in the Gardeners' Chronicle for 1 6th Sep-
tember 1876, p. 361. Mr. W. B. Grove, M.A., has re-
cently published a note regarding the second viola ^Ecidium
in the Journal of Botany, vol. xxi. p. 274. In this paper
the author states that he has detected the Uredo as well
as the Puccinia condition of ^Ecidium depauperans, Vize, on
violas ; and he describes the Puccinia as a new species under
the name of P. cegra ; the dEcidium appears at the end
of May and the Puccinia in August. Mr. Grove gives no
proof that the Puccinia is connected with the ^Ecidium ;
he believes they are connected because the three forms he
describes grow on the same plant. We have just shown
that in several instances it is acknowledged that jtEcidium
is not always assumed to be genetically connected with a
Puccinia when the two forms grow in company on the
same plant. Mr. Grove appears not to have experimented
with pro-mycelium spores.
The disparity in the number of species found under
j^Ecidium and Puccinia is great ; but the disparity is of
no moment if it is acknowledged that one form can go on
reproducing itself for an indefinite time without aid from
the other.
Besides the Puccinia violarum, Lk., and P. cegra, Gr.,
there is a P. violce, D.C., and a P. violce, Schum.
In conclusion, one more point must be adverted to. We
have shown that plants invaded by Puccinia and ^Ecidium
carry an hereditary disease by which they are saturated,
198 DISEASES OF FIELD & GARDEN CROPS. [CH.
and that the disease is capable of reaching the seeds and
reappearing in the youngest seedlings. Now, if plants
thus suffering from hereditary disease, and having the
latent germs of disease in every part of their organisation
are experimented upon in an unnatural way, have spores
of fungi placed near their organs of transpiration, whose
germ-threads can pierce the epidermis or enter and choke
the stomata and so reach their intercellular spaces, is it
not likely that this inociilating process may start into
activity the latent germs of disease ?
We have facts, as we think, quite comparable with this in
the animal kingdom. Persons are subject to different forms
of complaints, according to their constitution. Cereals are
notoriously constitutionally subject to mildew ; barberries
are notoriously constitutionally subject to blight.
Suppose we take an instance of a person constitutionally
subject to phthisis (consumption) ; give that person a cold
and phthisis appears ; but the same cold will give rise to
rheumatic fever with a second constitution, and scrofula
with a third, according to the tendency of the indi-
viduals to these disorders. Gout, for another example,
is said to be a jealous complaint, and, with those liable to
it, will always come and look in if any accident or ailment
should arise. Again, the same irritating article of food
will with one person produce neuralgia, with another the
vesicular skin disease named shingles, with a third indi-
gestion, with a fourth diarrhrea, with a fifth local inflam-
mation. Vaccination in the human subject, which is
comparable with spore inoculation in plants, unquestion-
ably puts latent ailments into action. Children badly
nourished will get eruptions and boils ; if of scrofulous
habits, abscesses or sores ; if gouty or of delicate constitution,
vesicular eruptions or eczema will often appear. The
shock of an accident will also often set latent ailments into
action ; a blow will set the latent germs of cancer into
activity. Under these circumstances we think artificial
cultivation of corn in pots under bell-glasses, with fungus
xxv.j CORN MILDEW AND BARBERRY BLIGHT. 199
spores which burst and enter by germ tubes amongst the
tissues of the plants experimented upon, may possibly set
the latent germs of the fungus of corn mildew into activity.
In the case of the bunt of corn and of the potato disease
the presence of the fungus is clearly evident to experienced
observers long before the slightest trace of mycelial threads
can be detected by the microscope. We cannot help
thinking that more and better evidence of the supposed
genetic relationship between corn mildew and barberry
blight is much wanted. New illustrations are also wanted,
from new and unbiassed observers, of the germinating
pro-mycelium spores of Puccinia piercing the cuticle of
the barberry. We regret that we are unable to produce
an original illustration of this process of growth. A
single illustration of this phenomenon has been copied,
recopied, and copied again, sometimes with, but more often
without, acknowledgment, that one now almost feels angry
at the mere sight of the by far too familiar engraving.
All low lying lands suffer most from mildew, and it is
said that elevated lands are next most seriously affected,
the intermediate positions being generally most free. This
fact is generally explained by the presence of mists in the
low lands, and clouds on the hilltops, the mists and clouds
being especially favourable to the development of Puccinia.
Mildew is commonly seen at its worst in placeswhere bushes
and trees abound, as these objects impede free currents of
air and aid fungus growth. Parasitic fungi which refuse
to grow in open gardens will often germinate and produce
disease at once if placed upon plants under bell-glasses.
The glass aids in keeping the air damp and motionless.
We have ourselves observed corn mildew to develop with
great rapidity after rain in August, and we have sometimes
noticed the late sown wheat to be most affected. When
the ears are badly attacked the grain is not only greatly
impoverished and reduced to "skeleton grain," but it is
hardly possible to separate the seed from the husks.
Mildew is said to be more frequent after crops of clover
200 DISEASES OF FIELD & GARDEN CROPS. [CH.
than after other crops. We think the fact of straw from
stables being so frequently thrown over old clover fields a
sufficient explanation of this fact. Wheat after clover is
certainly a favourite alternation of crops with many
farmers, — perhaps because the old decaying clover roots
act as good manure for the corn. When clover precedes
corn it should be heavily folded with sheep, and straw
from stables should not be used as manure.
It is now generally accepted as a fact amongst practical
men that after dressing the land with farmyard manure
and nitrate of soda, mildew often puts in a strong appear-
ance ; but after mineral manures, bone superphosphate,
and bone meal drilled with the seed, rust and mildew are
much less apparent. There can be no doubt that farm-
yard manure has a tendency to produce a gross soft growth
in corn which is suitable for fungi, and that mineral
manures, on the contrary, have a tendency to produce a
firm stiff growth unsuited for rust and mildew. As corn
generally does so well in dry limestone and chalky dis-
tricts, a hint might be derived from this fact as to the
desirability, where possible, of manuring land with chalk.
We have seen this done with success in North Herts and
South Bedfordshire, where chalk is easily obtainable.
It is probable that the resting -spores of the fungus
of corn mildew seldom hibernate through two seasons ;
therefore, in instances where stable manure must be used,
it should if possible be used in the crop preceding the corn
or the crop following it rather than for the corn itself.
An alternation of crops is in every way desirable.
Beans, peas, turnips, potatoes, clover, and other farm pro-
duce should be taken alternately with corn.
There is but one way of getting rid of corn mildew,
and that is certainly not by cutting down barberry bushes
and pulling up borage plants. Corn mildew is a heredit-
ary disease, and therefore no seed corn should be gathered
from mildewed plants. If the hereditary nature of the
disease is disputed, it cannot be disputed that certain
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 201
examples of corn have a strong and inherited predisposition
for mildew ; therefore predisposed examples should be
struck out and no seed gathered from them. Especial care
should be taken in the rigorous selection of seed from white
wheats, which are notoriously more subject to mildew than
red, probably because the latter are naturally more robust.
If seed merchants would guarantee that the seed corn they
sell is taken solely from corn free from mildew, in the
course of years the attacks and consequent losses from this
pest would be considerably lessened. Mildew is every
year so common in our fields simply, as we think, because
the disease is planted with the grain. Old corn stubble
should not be left too long in the fields. Some corn
growers say that a top dressing of salt has a tendency to
lessen or prevent mildew.
Mildewed straw is bad when used as food for stock in
chaff, and the inferior grain is hardly fit for pigs. The
straw is more commonly used as litter in stables. In this
position the spores of the Puccinia remain uninjured, for
neither warmth, frost, wet, or dryness materially affect
the vitality of the resting -spores of the fungus of corn
mildew. They are so small that no amount of treading
from horses, herds, or flocks injures them. The warmth
and dampness of the stable floor in every way suits them,
and they are frequently taken from this position, full of
life, and at once thrown on to the fields in the saturated
straw. If the spores are consumed with food by animals,
their passage through the alimentary canal does not injure
them. The disease is probably, as we think, propagated
by the mildewed straw being used as manure, and by the
germinating resting-spores of the fungus of corn mildew
infecting the first young leaves of the corn.
Mildewed straw should be destroyed, because the Puc-
cinia, with its myriads of resting-spores, is in this material.
We have shown that these resting-spores germinate in the
spring and early summer at the exact time when rust,
which is the early state of mildew, first appears. Whether
202 DISEASES OF FIELD & GARDEN CROPS. [CH.
the resting-spores attack barberry bushes, or whether they
do not, is of no great importance, for there are generally
no barberry bushes to attack. The mildewed straw should,
as far as practicable, be destroyed, and the hedges kept
clear of rusted and mildewed grasses.
In taking the position here advanced in regard to the
fungi of corn mildew and barberry blight, it must not be
assumed that we tinder-estimate or disrespect the valuable
published opinions of other observers. Those published
opinions are, some of them, counter to ours, and we know
the risk we run in appearing to question them. Still,
the conclusions here given have been arrived at after
many years' study, with living examples before us, and
if we are not right in our opinion, we think we have
advanced a sufficient number of facts to show that new
and better evidence is much needed before the connection
of corn mildew and the blight of barberries can be gener-
ally accepted as proved. Physiology and pathology have
taught us much, but there is infinitely more to learn.
No account of the supposed connection of corn mildew
with barberry blight would be complete without a notice of
the essay, by Professor A. S. Oersted, published in Copen-
hagen, in the Botanische Zeitung, in 1865. This essay is
intended to show that a common fungus of the Savin,
named Podisoma Juniperi-Sabince, Fr., is one condition of
an equally common fungus of pear leaves named Rcestelia
cancellata, Reb. The Podisoma is a close ally of Puccinia
or mildew, and the Rcestelia of sEcidium or blight. Pro-
fessor Oersted says that he had learned that gardeners were
of opinion that the pear fungus was never seen except
after the appearance of the fungus on Savin. In contrast
with this statement, if we turn to the Mycologia Scotica we
find that the fungus of Savin is recorded from Scotland,
but that the supposed secondary state belonging to pear
leaves is not a Scottish fungus. Scotland is famous for
its excellent gardeners, and it appears hardly possible,
therefore, that any gardener can have seen in Scotland
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 203
the pear fungus following the one on Savin. Savin, like
the barberry, is absent and under a general ban in many
country places, owing to the improper use to which the
fetid volatile oil from the leaves has often been put as
an emmenagogue ; but pear leaves with the Rcestelia are
everywhere common. The Rev. M. J. Berkeley has
recorded in Hooker's British Flora, vol. v. part ii. p. 5,
that when young pear trees are planted near old ones
suffering under the Rcestelia, the young trees have been
observed to become much injured by the fungus ; and
Mr. Knight sowed pear seeds in soil infested with
Rwstelia, and the very youngest leaves of the seedlings
showed the disease.
Professor A. S. Oersted carried out the usual successful
infecting experiment, but in regard to it he very properly
remarked : "It may very easily happen that the above
experiment may be repeated many times without success,
for those who are occupied in this kind of work know that
a certain amount of good fortune is necessary for success."
The correctness of Professor Oersted's experiments and
views were confirmed by Professor De Bary of Strasbourg
in the same year, Bot. Zeit., p. 222, 1865, and this confir-
mation tended greatly towards the general acceptance of
Professor Oersted's view. Two other species of European
Rcestelice allied to R. cancellata, Reb., were also connected
with two allied European species of Podisoma, related to
P. Juniperi-Sabinw, Fr.
In the Anniversary Memoirs of the Boston Society of
Natural History, Professor W. G. Farlow of Harvard Uni-
versity, a gentleman who studied with Professor De Bary,
and therefore had good opportunities of seeing original
experiments carried out with germinating fungus spores,
has published a paper termed the Gymnosporangia, or Cedar
Apples of the United States : Boston, 1880. In this paper
Professor W. G. Farlow reviews the whole evidence for
the connection of Roestelia with Podisoma (or, as he terms
it, Gymnosporangium) in the light of the large number of
204 DISEASES OF FIELD & GARDEN CROPS. [CH.
additional species belonging to both genera as found in
the United States. The author says : " There is nothing
to confirm the views of Oersted as to the connection of
particular species."
In reference to Podisoma Juniperi - Sabince, Fr., and
Rcestdia cancellata, Reb., Professor Farlow points out
that the former is " very common in Massachusetts,
whereas its supposed JEddium, — R. cancellata, Reb., is not
known with certainty to occur at all."
Professor Farlow experimented in 1876, 1877, and
1878, with the spores of Podisoma Juniper i-Sabince, Fr.,
with the following results : —
Nine leaves were taken, three of Cratcegus (hawthorn),
three of Amelanchier (medlar), and three of apple. Sper-
mogonia appeared on the three Cratcegus leaves only. All
nine leaves were such as the Rcestelice grow upon naturally.
An experiment was then made with a young plant of
Cratcegus. Result — nothing.
On five leaves of Cratcegus. Result — nothing.
The first experiment was repeated with three leaves of
Pyrus added, out of the twelve leaves spermogonia appeared
on the three of Cratcegus only, although Pyrus was the
plant pointed out by Oersted.
Ten leaves, six Cratcegus, three pear and one apple.
Result — nothing.
Professor Farlow also records that not only did he get
spermogonia on the leaves of Cratcegus tomentosa, L., after
the application of the spores from Podisoma Juniperi-Sabince,
Fr. ; but he got a like result from the application of the
spores of Gymnosporangium biseptum, Ellis, and G. macro-
pus, Lk., both fungi being peculiar to America, one ex-
tremely rare, and both unknown in Europe. On Pyrus,
the genus especially pointed out by Oersted, there was
invariably no result. Sometimes the spermogonia appeared
suspiciously early, as in four days, — Oersted gave nine or
ten, — and sometimes the uninfected control plants also
exhibited spermogonia. Mr. Plowright mentions eighteen
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 205
days as requisite for a closely-allied species, Gardeners'
Chronicle, 28th October 1882.
The experiments were varied in many ways, and
if Professor Farlow' s full reports of the carefully conducted
experiments are examined, it will be seen that heteroecism,
as regards the American species of Podisoma and Rcestelia,
completely breaks down at every point. Out of nineteen
experiments no less than fourteen were without result ;
when results followed they were contradictory, and
sometimes, as Professor Farlow remarks, " desperate."
In 1879 Professor Farlow was absent, but in 1880 he
returned and made further experiments : these were
invariably without result.
We have shown in this work that Puccinia (mildew)
and dEcidium (blight) are potentially perennial, heredit-
ary, and always either in an active or passive state
in the juices of the plants invaded. Professor Farlow
adverts to the same fact as regards the American fungi,
and suggests that the appearance of the spermogonia
was in consequence of the presence beforehand, in the
leaves, of the mycelium of some Rcestelia which was made
to develop by the moist condition in which it was placed.
" I am strongly inclined," writes Professor Farlow, " to
favour this view." He further states that, unless he is
mistaken, he has seen the Roestelia state earlier in the
season than the Podisoma • and so, instead of following, as
stated by Oersted, it has preceded, the Podisoma. He
concludes by saying : " Another important fact is to ascer-
tain how many of our Rwstelice are perennial. This at
least appears to be the case with R. aurantiaca, Pk. If it
should be shown that several of our Rcestelice are perennial,
a fact true of our Gymnosporangia (Podisoma), and to grow
in regions remote from Juniperus and Cupressus, then one
could not help feeling that any connection between the
two genera was probably accidental rather than genetic."
The amount of confusion that exists in books as to the
host plants, and second conditions of the so-called heterce-
206 DISEASES OF FIELD & GARDEN CROPS. [CH.
cismal or metcecious fungi is now almost beyond conception.
The literature is bewildering and contradictory in the
highest degree. The following instance is one of the
simplest; many others are involved almost beyond the
power of unravelling.
In 1872 we observed an ^cidium invading quinces in
Mr. Alfred Smee's garden at Hackbridge. At that time
no such plant on quinces had been noticed as British ; but
the fungus was published by us in Mr. Smee's My Garden
under the name of ;E. cydonice, Lenz. We afterwards
learned that this fungus was considered to be a mere form
of Rcestelia cornuta, Tul., a parasite of mountain ash, and
proved (?) by experiments with spores to be a second con-
dition of Gymnosporangium Juniperi, Lk. Professor W.
G. Farlow mentions a distinct Rcestelia frequent on quinces
in the United States, under the name of R. aurantiaca,
Pk., and this is probably our plant. Professor Farlow
considers it a true species ; but other botanists look upon
it as a variety, — not, however, of theR. cornuta, Tul., just
mentioned, but of the totally different It. lacerata, Tul.,
whose alternate condition is said to exist in Podisoma Juni-
peri, Fr., a second parasite of Juniperus communis, L. In
the United States, strange to say, R. aurantiaca, Pk., and
R. lacerata, Tul., grow in company on the same host
plants.
If the statements just given as to the quince fungus
are correct, we have two confessedly very different species
of fungi, both frequent on Juniper, and both able to
invade quinces, and produce specifically different fungi on
the leaves and fruit, — the characters of the two quince
fungi being in turn so much disputed by botanists, that
their names are sometimes transposed, and one made to
do duty for the other ; each Rcestelia being supposed to
answer to the characters peculiar to the other one.
Note. — On page 175 we have stated that the initial experi-
ment of producing an ^cidium from germinating Puccinia
xxv.] CORN MILDEW AND BARBERRY BLIGHT. 207
spores was made by Professor De Bary with Puccinia tragopo-
gonis, Corda, the assumed result being ^cidium tragopogonis,
Pers. The latter is abundant in this country, but the former,
illustrated from a continental example in the left-hand figure of
Fig. 90, has never yet been found in Britain. A second species
of Puccinia, however, named P. sparsa, Ck. , is found both in
Britain and on the Continent on Tragopogon. This has rough
or slightly echinulate spores, and is illustrated in' the right-
hand figure of Fig. 90. Dr. M. C. Cooke maintains the dis-
X-SOO
FIG. 00.
Teleutospores of Puccinia tragopogonis, Corda, and P. sparsa, Ck.
Enlarged 500 diameters.
tinctness of the two fungi, and P. sparsa, Ck., is retained as a
species in the Mycologia Scotica. Many botanists, both British
and Continental ; look on the two parasites as quite distinct,
but the advanced advocates of hetercecism now say they are the
same, and term both P. tragopogi, Pers., although Persoon
never gave this name.
We have not heard whether ^Ecidium tragopogonis, Pers.,
follows the germinating spores of Puccinia sparsa, Ck., on
goats-beard, — if it does, and the two species of Puccinia are
distinct, the case is similar with the one mentioned by Pro-
fessor W. G. Farlow, where spermogones appeared on pear
leaves after the application of the germinating spores of two
species of Gymnosporangium and one of Podisoma.
CHAPTER XXVI.
NEW DISEASES OF WHEAT, BARLEY, AND RYE -GRASS,
CAUSED BY
Fusisporium culmorum, hordei, and Lolii, W.Sm.
THERE can be no doubt that the injury caused to food
plants by various species of Fusisporium is much greater
than is generally supposed. We have already stated that
a form of the potato disease is caused by one or more pests
belonging to this genus. Perhaps one reason why some
species of Fusisporium have been overlooked is because
they are almost invisible without careful attention, com-
bined with a knowledge of their habits of growth. Some
species only resemble to the unaided eye a small gelatin-
ous patch, and when this patch is exactly the same colour
as the matrix on which it grows (which is often the case),
the difficulty of detecting the fungus is increased.
A description of Fusisporium was given under Fusis-
porium Solani, Mart., Chapter V. , one of two species found
on potatoes. We will now briefly advert to three species
which at present have found no place in the text-books of
this country.
There is a Fusisporium found on wheat generally tinted
with cream-colour or yellow, and possibly varying in col-
our from white to pink. It attacks the ears, chiefly per-
haps of those plants which have been more or less invaded
by corn mildew or other cereal fungi. This Fusisporium
forms a pale yellow-orange gelatinous stratum over the
ears or some portions of the ears. It glues the spikelets
together and stops the growth of the grain. Although
this pest has apparently been hitherto unrecognised by
OH. xxvi.] NEW DISEASES OF WHEAT, ETC.
209
botanists, we have heard of it from agriculturists at various
places. The upper portion of an infected ear, as sent to
us by Mr. Chas. B. Plowright from "West Lynn, Norfolk,
is illustrated, natural size, at Fig. 91. The dwindled por-
tion at the apex from A to B is the
part overrun by the Fusisporium.
The fungus, owing to its orange
colour, gives the ear a spurious ap-
pearance of ripeness. When the
plant is magnified 400 diameters, it
is seen, as at A, Fig. 92, where the
crescent - shaped fusiform septate
spores, so characteristic of the fungus,
are illustrated. One of the spores
at B is seen breaking up into four
portions. After a short rest each
portion will burst and produce new
mycelium. A single spore is farther
enlarged to 1000 diameters at C.
The long cells at D belong to one of
the outer glumes of the wheat spike-
let. This plant may be named
Fusisporium culmorum, W.Sm., and
described as follows : — Mycelium
effused, gelatinous, yellow or orange,
sparingly septate, torulose ; spores
large, fusiform, 3 '5 septate, orange. On wheat, fixing
the pales, glumes, and spikelets together.
Another Fusisporium, belonging to barley, has recently
attracted attention, and this was described and illustrated
with two plates in the Jour. Roy. Micro. Soc. for June
1883, p. 321, under, as we think, the incorrect name, as
furnished by Dr. Chr. Hansen of Copenhagen, of Fusarium
graminearum, Schwb. ; this is the F. graminum of Corda.
Unfortunately no scale of magnification was given with
the plates. The author of the article, Mr. Chas. Geo.
.Avs, AV.IS good enough to send us some of the infected
P
FIG. 91.— Upper part of
an infected ear of wheat
invaded by Fusisporium
ciiZmorwra, W.Sm. Nat-
ural size.
210 DISEASES OF FIELD & GARDEN CROPS. [on.
grains of barley, from which our illustrations were made.
At Fig. 93 five of the " red corns," as maltsters term them,
o
•X-400
FIG. 92.
Fusisporium, culmorutn, "W.Sm. Enlarged 400 diameters.
Spore enlarged 1000 diameters.
are illustrated twice the natural size. These misshapen
" red corns " are to a great extent covered with fungus
FIG. 93.
Red corns of barley, with growths of Fusisporium Jiordei, W.Sm.
Twice the size of nature.
spawn and spores, ranging in colour from pale orange to
bright scarlet or deep cinnabar -crimson. The fungus
itself, illustrated to the same scale as Fusisporium culmorum,
xxvi.] NEW DISEASES OF WHEAT, ETC.
211
W.Sm., is shown at Fig. 94. It will be noticed that
many of the spores have broken up, as in the last, and
formed little globular spores of a second series, as at AA.
These are destined to rest for a short period. Other of
the spores, as at BB, are germinating whilst still attached
to their supporting threads, — a common phenomenon in the
genus Fusisporium. A single spore is enlarged to 1000
diameters at C. The cells at D belong to the flowering
glume. Mr. Matthews states that the spawn of the fungus
•X-.400
x-iooo
FIG. 94.
Fusisporium hordei, W.Sm. Enlarged 400 diameters.
Spore enlarged 1000 diameters.
will run over paste made of crushed germinating barley,
and throw up red patches from half an inch to three-
quarters of an inch high. The fungus is said chiefly to
invade barley of poor quality and ill-conditioned crops
and ears, seldom or never appearing on good sound barley.
The germinal end of the grain is distinctly the part most
seriously attacked, perhaps because it is the softest, being
the spot whence the plumule and radicle of the young
plant is destined to emerge. It is clear that no badly
212 DISEASES OF FIELD & GARDEN CROPS. [cir.
affected grains can possibly germinate.
In both wheat and barley the fungi
when present give the grain a peculiar
and disagreeable taste. Mr. Matthews
states that when the Fusisporiurn is
introduced into sterilised beer-wort it
gives rise to sluggish ferments, pro-
ducing alcohol and carbonic acid gas.
This species may be named Fusisporium
hordei, W.Sm., and described as fol-
lows: — Mycelium rose-coloured or
crimson ; torulose, effused, forming
a thick gelatinous stratum, septate ;
spores fusiform, acuminate at both
ends, 1'3 septate, rose-coloured, crim-
son, or cinnabar-red. On barley, form-
ing red gelatinous patches.
The third illustration at Fig. 95
shows, natural size, an orange-coloured
Fusisporium on the common and valu-
able perennial rye-grass, Lolium perenne,
L. The spike illustrated is ergotised,
and the example illustrated was sent by
Mr. Chas. B. Plowright from Norfolk.
The Fusisporium is shown at A, B, C,
and D. Lolium perenne, L., is unusu-
ally subject to ergot, and it is remark-
able that in the spike illustrated the
Fusisporium was invading the ergot as
well as the different parts of the
spikelet and seed. Young ergots are
shown at E, F, and G ; others are
Hum perenne, L., in- •? . •*• .
vaded by Ergot and curious species is enlarged 400 dia-
Loiu, meters at Fig. 96, a single spore being
enlarged to 1000 diameters at A ; the
illustration at B shows the mycelium running over the
Fusisporium
W.Sm.
xxvi.J JVEW DISEASES OF WHEAT, ETC. 213
cells of the ergot. This species may be named Fusisporiutn
Lolii, W.Sm., and described as follows: — Mycelium
orange, torulose, effused, forming an orange gelatinous
X-400- c-^ -X-IOOO
FIG. 96.
F'usisporium Lolii, W.Sm., growing on Ergot. Enlarged 400 diameters.
Spore enlarged 1000 diameters.
stratum ; spores fusiform, acuminate at both ends, 1*4
septate, orange. On rye -grass and its ergot. It is quite
possible that some species of Fusisporium may be parasitic
in habit. One species, F. mucophytum, W.Sm., grows on
sound edible mushrooms ; and F. obtusum, Ck., is described
as parasitic on old fungi found under Diatrype, and a
similar species with the last occurs in North America.
CHAPTER XXVII.
ERGOT.
Glaviceps purpurea, Tul.
THE terrible effects of ergotised grass on animals that have
partaken of it are so serious and so well known that a
clear knowledge of the nature of ergot should be possessed
by all persons interested in agriculture.
Ergot has a powerful and immediate effect, and especially
so when quite fresh, in exciting muscular contraction in
certain parts of animals, notably the uterus. The same
contracting power of ergot is no doubt the primary cause
of the well-known gangrenous diseases always popularly
associated with this substance. Ergot, by contracting the
muscles, stops the flow of blood to the extremities, and
these extremities, unsupplied with fresh blood, sometimes
rot and drop off.
The ergot produced by rye, Secale cereale, L., is one of
the largest, best known, and probably the most potent,
and this is the substance invariably used in medicine.
Ergot is commonly termed ergot of rye, but the fungus
growth is very common on other cereals and 011 many grasses.
Amongst others it has been recorded in Britain as parasitic
upon the following plants : — Mat grass, Nardus stricta, L. ;
catstail-grass, Phleum pratense, L. ; foxtail grass, Alopecurus
pratensis, L. ; reed canary-grass, Phalaris arundinacea, L. ;
vernal grass, Anthoxanthum odoratum, L. ; waved hair-
grass, Aira flexuosa, L. ; turfy hair-grass, Aim ccespitosa, L. ;
oat-grass, Arrhenatherum avenaceum, Beauv. ; meadow soft-
grass, Holcus lanatus, L. ; creeping soft-grass, Holcus moil-is,
L. ; cocksfoot grass, Dactylis glomerata, L. ; smooth meadow-
CH. XXVII.]
ERGOT.
215
grass, Poa ann wa,L.; floating sweet-
grass, Glycerin fluitans, R.Br. ; tall
fescue - grass, Festuca elatior, L. ;
meadow fescue-grass, Festuca pra-
tensis, Huds. ; wheat, Triticum
sativum, L. ; wheat-grass, T. repens,
L. ; rye-grass, Lolium pefenne, L. ;
darnel -grass, Lolium temulentum,
L. ; lyme- grass, Elymus arenarius,
L. ; rye, Secale cereale, Welld. ;
barley, Hordeum distichum, L., on
species of Agrostis, and no doubt
on many other grasses, including
rice, Oryza sativa, L. A large North
American species of lyme-grass,
sometimes seen in our gardens
under the name of Elymus gigan-
teus, Vahl., produces a very large
ergot.
In describing ergot we will take
a spike of ergotised rye, and after
examining one of the ergots which
it has produced, we will follow up
the life-history of the fungus and
show how it produces other ergots
like itself. We will then glance at
the effects of ergot on man and
other animals, and see how the
attacks of ergot on grasses may be
lessened or prevented.
If we take a spike of ergotised
rye — Secale cereale, L., as illus-
trated, natural size, at F4g. 97 —
we see one or more of the rye
seeds replaced by blackish hornlike
* FIG. 97.— Spike of ergotised Rye, Secale
cereale, L. Natural size.
216 DISEASES OF FIELD & GARDEN CROPS.
growths, as at A, B, C, D, twice or three times as long and
stout as the normal rye seeds. In old times the ergot was
supposed to be an unusually large and diseased rye grain.
If it grew on wrheat it was considered a somewhat large grain
of wheat, just as the galls known under ear-cockle were
at one time supposed to be unusually small and cockled
If we remove an ergot from a spike we shall see
X 5 X
FIG. 98.
Ergots of Rye, and section. Twice the size of nature.
Lodicule of Eye. Enlarged 5 diameters.
that its production has not materially injured the spike of
grain, and that its growth has been confined to the spike-
let from which it was taken. On looking at ergots with
a lens we see them, enlarged to twice their natural size,
as at Fig. 98. We notice that they are longitudinally
corrugated and minutely granular, often slightly split both
transversely and longitudinally, the cracks often showing
xxvn.] ERGOT. 217
a reddish margin, and exposing the whitish interior sub-
stance of the ergot. A transverse section through an ergot
is shown at A. Sometimes the minute scales or lodicules
remain attached to the ergot, as at B. These latter organs
are farther enlarged to five diameters at C, so that they
may be compared with the smaller lodicules of wheat
enlarged to the same scale at Fig. 42. See also Fig. 43.
There is a faint sickly odour of camphor attached to fresh
ergot, and if we hold it in a flame it immediately takes
fire and burns like the kernel of a nut, constantly giving off
little jets of flame, and dispelling a not unpleasing odour.
The ergot burns thus freely because it contains a brownish-
yellow, viscid, aromatic, slightly acrid, oil. Its taste when
raw is slightly bitter and nauseous. If we now cut an
ergot in two, either transversely or longitudinally, and
then remove an extremely thin transparent fragment from
the exposed surface and magnify 400 diameters, we shall
see the structure as at
Fig. 99. We now ob-
serve a densely -com-
pacted mass of cells
with thick pale-brown
walls, many of them
made polyhedral by
the pressure of adjoin-
ing cells. A few cells
are elongated and ap-
pear to wind between
the globular and polygonal cells in a sinuous fashion. If
the ergot is sliced in any part, the same appearance
presents itself — densely -packed thick -walled cells filled
with a viscid oily liquid.
After this examination we clearly see that we are not
dealing with a perfect fungus, but with a Sclerotium not
dissimilar from the one found on potatoes, and illustrated
to the same scales with ergot in Figs. 4 and 5. In
colour, size, and general appearance, the Sclerotia of
218 DISEASES OF FIELD & GARDEN CHOPS. [CH.
potatoes and the ergot of rye are so much alike that if a
large number were mixed together it might be no easy
task to distinguish in every instance one from the other.
Ergot is described in old botanical books as Sclerotium
clavus, D.C., or Spermadia clavus, Fr.
If we keep ergots all through the winter on moist
sand on a garden bed, or indoors, they will germinate in
the early summer precisely in the same way as the
Sclerotia of potatoes already described ; and although the
ergots will not produce a Peziza as the potato Sclerotia
did, yet they will give rise to a fungus of equal if not of
greater interest.
If ergots are laid in a clean, moist, shady place in a
garden, they will germinate naturally in June ; or if a
search is made where dead ergotised rye or other ergotised
grasses have lain, the germinated ergots will sometimes
be found without difficulty.
Just as different grasses vary a little in their time of
flowering, so ergots vary in their time of germination.
An early flowering grass is invaded by an early germi-
nating ergot, and a late grass by a late ergot. Grasses
and ergots alike flower and germinate at a somewhat
different period in the south of England and the north of
Scotland. Surrounding circumstances have modified the
habits of both grass and fungus. The range of time in
the flowering of grasses and germinating of ergots is in-
cluded in about three months.
If we now take a germinated ergot such as either of the
two illustrated, twice the natural size, at Fig. 98, and
examine it, we shall see that it has produced several club-
shaped growths, curiously answering in appearance to,
although considerably smaller in size than, the Peziza
with the slender tortuous stem produced by the potato
Sclerotium, as illustrated, natural size, at Fig. 6. Like the
potato Peziza the growths from ergot have a somewhat
long tortuous stem and a cap, as illustrated the natural
size, at Fig. 100, and here the similarity ends. Each little
XXVII.]
ERGOT.
219
white-stemmed fungus which, grows from a white downy
base out of ergot in the summer
is furnished with a small spherical
head of a beautiful pale purplish
colour. This growth, enlarged five
diameters at A, Fig. 101, was at
one time supposed to be a parasite
on ergot, but it is now known to
be the perfect condition of the Fia.ioo.-Ergots^erminating
ergot itself. The Claviceps derives
its nourishment from the ergot,
and after the Claviceps has appeared
the ergot collapses and perishes ater the manner of the
seed tuber of a potato plant. Tulasne has named this
perfect state of ergot Claviceps purpurea. Claviceps of
and producing Claviceps
purpurea, Tul. Natural
size.
2.0
FIG. 101.
Claviceps purpurea, Enlarged 5 diameters ; and section through head or
stroina. Enlarged 20 diameters.
course refers to the clublike head, and purpurea to its
beautiful purplish colour. The popular name " Ergot "
is French for the spur of certain birds.
Our attention must now be directed to one of the pale
220 DISEASES OF FIELD & GARDEN CROPS. [CH.
purple globose heads. Externally they are irregularly
dotted over with little prominences, as at Fig. 101, A,
enlarged five diameters ; and in this they greatly resemble
the clubs belonging to Torrubia found growing upon a
buried truffle (see Fig. 22). On cutting longitudinally
through the head of the Claviceps of ergot we find it like
the parasitic growths belonging to the truffle (and unlike
the potato Peziza illustrated
at Fig. 7). We see it, if
enlarged twenty diameters,
as at B, Fig. 101, packed all
round its outer surface with
small flasks, conceptacles, or
perithecia, with the mouths
of the flasks all opening
towards the outside. The
little projecting mouths, as
at CC, represent the minute
prominences seen on the
outside of the cap or stroma
at A. As with the former
fungi we have described, we
must now cut an extremely
thin transparent slice off
the exposed cut surface of
the head, and magnify with
the highest powers of the
microscope to make out the
nature of one of the minute
conceptacles or perithecia
and its contents.
If we magnify a single perithecium or conceptacle 200
diameters, we shall see it as at Fig. 102. We now notice,
as in former examples, that the flasks, conceptacles, or
perithecia are closely packed with fine long transparent
bladders, which spring from the base of the perithecium.
The mouth from which these bladders ultimately emerge
•X ZOO
FIG. 102. -Section through a concep-
tacle or perithecium of Claviceps
purpurea, Tul. Enlarged 200
diameters.
XXVII.]
ERGOT.
221
is at A. By dexterously moving
the cover-glass, or "by manipulating
with a needle, we may easily get
some of these contained asci or
bladders free from the flask, and
when free a single ascus will be
seen if magnified 500 diameters, as
at A, Fig. 103. "We now perceive
that they are not empty bladders,
for through their transparent walls
we can see that each contains eight
extremely fine long attenuated
bodies, which are sporidia or spores.
One of these spores from an ascus
is farther enlarged to 1000 diam-
eters at B. If reference is now
made to the ascus and sporidia
belonging to the potato Peziza, as
illustrated to the same scale at Fig.
9, the difference, especially in the
spores, will be seen ; and if the
sporidium of the Torrubia of the
truffle is turned to at H, Fig. 22, a
similar long attenuated body will
be seen. The sporidium from the
Torrubia is furnished with an enor-
mous number of joints, whereas it
will be noticed that the body now
before us is apparently in one piece.
The exceedingly small, long, needle-
like, extremely attenuated sporidia
belonging to germinated ergot are
produced in June. When we have
an ergot with the club-shaped
Claviceps upon it we have the ergot
in fruit.
On a June day, then, we have a fruiting ergot before
\
X-5DO X-000
FIG. 103. — Ascus of Clavi-
ceps purpurea, Tul. En-
larged 500 diameters.
Sporidium of ditto, en-
larged 1000 diameters.
222 DISEASES OF FIELD & GARDEN CROPS. [CH.
us, and the problem presents itself for solution — How do
these inconceivably minute hairlike bodies or sporidia
free themselves from the asci and perithecia of the
Claviceps, and cause ergot in grasses ?
If germinated ergots are kept in moist air under a bell
glass and observed against a black background in sun-
shine, the ejection of the spores like shining needles or
almost invisible glittering arrows may be clearly seen
with a lens. What power it is that causes the discharge
from the flasks no one has at present certainly explained ;
but the phenomenon is well known to occur in many
fungi. The discharge takes place after a sudden touch
or movement, or on a change of light or temperature, as
when a sunbeam suddenly falls on any ascus- bearing
fungus. The glittering hairlike spores are shot from the
mouths of the perithecia into the air radially in all
directions. The sporidia may be easily caught and
examined if strips of glass are smeared slightly with
glycerine and placed under a bell glass near the ripe
fungi. The sporidia can then be examined whilst alive
and fresh. The asci as well as the free sporidia are
often expelled, and it frequently happens that the act of
expulsion is too weak to propel the sporidia or asci into
the air, and they hang only half expelled from the mouths
of the perithecia.
If we now suppose ourselves to be in a district where
rye is common, and where rye was ergotised during the
previous autumn, we shall have the rye in flower at
the precise time when these myriads of glittering little
needle-like sporidia are sailing through the air. Such
needle-like spores as do not light upon flowering grasses
perish ; but where there are so many millions of sailing
spores some must of necessity fall upon the flowers of a
grass — let us say rye.
We must now imagine a needle-like sporidium falling
close to the pistil and stamens, and reaching the base of the
pistil of a flower of rye. Here the spore bursts or germi-
xxvii.] ERGOT. 223
nates, and in bursting forms a microscopic drop of glittering
vital material. This glittering viscid drop at the base of
the pistil speedily increases in size, and after about three
days it becomes visible to the unassisted eye. Its increase
in size is probably aided by an exudation from the rye
spike itself near the base of the pistil, and by absorption
of moisture from the atmosphere. The glittering liquid
has a great attraction for various flies, and no doubt
insects aid in the propagation of ergot. The liquid,
however, appears to kill some flies ; for our friend Mr. A.
S. Wilson has made the curious observation that he has seen
hundreds of flies standing on newly-ergotised grasses in
the stillness of death. On examination of this viscid drop
with the microscope, we see it traversed by a transparent
filamentous sweet -tasting mycelium, the beginning of
ergot. The mycelium attaches itself to the base of the
pistil, and partly covers the ovary ; it partially penetrates
the tissues of the pistil, generally leaving the upper part
exempt. The mycelium now occupies the place of the
pistil which it soon pushes from its place, and as the
fungus enlarges in size it becomes deeply furrowed and
honeycombed, and often carries the feathery styles on its
apex, and so somewhat resembles an aborted grain of rye.
As the viscid mycelium continues to grow, its base be-
comes compact and indurated, and this indurated base,
anatomically connected with the viscid matter above, is
the beginning of true ergot. A half-grown ergot is illus-
trated at Fig. 104, enlarged five diameters, with the true
ergot at the base, A, and the viscid matter at the top, B.
The apex at C is crowned with the withered styles which
have been forced from their natural position. At D a
longitudinal section through this young ergot and its
glutinous top is illustrated.
In old times the early viscid condition of ergot, coating
and pushing up the aborted seed was considered to be a
distinct fungus, and, like the Claviceps of germinating
ergot, was considered a parasite of ergot. The glutinous
2-24 DISEASES OF FIELD & GARDEN CROPS. [CH.
•x-s
FIG. 104.
Half-grown Ergot and section. Enlarged 5 diameters.
condition of ergot is generally described under the name
of Spliacelia, a word derived from the Greek, and meaning
gangrene. The same word in a secondary sense means
XXVII. ]
ERGOT.
225
mildew. We have the English word sphacelate, which
means to become affected with gangrene.
It unfortunately happens that this viscid early condition
of ergot has had other names in addition to Sphacelia.
Queckett termed it Ergotcetia abortifaciens, whilst Messrs.
Berkeley and Broome believed it to be an Oidium, and
described it as 0. abortifaciens; and in this they were
perfectly justified, for the fungus displays the characters
of a true Oidium, a condition of some fungi which has
been adverted to be-
fore in this work
under the Oidium of
the turnip, and the
Oidium or early con-
dition of the Ery-
siphe of grass mildew.
The Sphacelia is the
o
Oidium or larval
state of ergot.
We must now once
more return to the
microscope, and take
a thin, transparent
slice from the point
of junction between
the indurated ergot
below and its viscid
Oidium top. If we
magnify a thin sec-
tional slice 400 di-
ameters, we shall see
it as at Fig. 1 05 : the
X-400
FIG. 105.— Section through the Sphacelia or
Oidium of Ergot. Enlarged 400 diameters.
compacted cells of the ergot are at the bottom left-hand
corner, and the much looser mucilaginous and filamentous
growth is shown above. The upper part is the Oidium or
Sphacelia, which not only carries its own nucleated conidia,
but is often swarming with bacteria and minute infusiorial
226
DISEASES OF FIELD & GARDEN CROPS. [CH.
0
animals. We now see that the walls between the fur-
rows consist of elongated Oidium-like cells springing from
a gelatinous substratum, and each cell supports an Oidium
spore or conidium. These little spores or sporidia ger-
minate very readily in water or on any moist surface, and
by this means they not only continually increase the
Oidium growth, but they can, on being blown on to grass
flowers, cause the production of
other viscid early states of ergot.
It will be remembered that Oidium
Balsamii, Mont., reproduces itself in
the same manner on turnips ; 0.
monilioides, Lk., does so on grasses,
and 0. Tuckeri, B., on the vine. A
conidium or stylospore germinating
at A, and producing another stylo-
spore exactly like itself at B, is
shown at Fig. 106, enlarged 1000
diameters. As the true compacted
ergot below increases to its full
FIG. 106. — Spore of Spha- . ,, , J./TT-
celia or Oidium of Ergot, slze> tne less compact Oidium growth
germinating at A, and above collapses and falls away. The
producing a spore or true ergot js then left devoid of its
ZZSESL?^ viscid, watery apex, with its little
tortuous furrows and conidia or
stylospores in the condition in which we first began our
observations upon it.
The value of the Oidium growth to the ergot is obvious,
for if one needle-shaped spore from a Claviceps produces
the infant state of ergot on one rye-flower, we see that in
a few days this larval condition can produce thousands
of new Oidium spores, each spore being equally powerful
with a Claviceps spore in producing ergot. Ergots seldom
germinate and produce the Claviceps when more than a year
old ; two-year old examples have, however, been known
to sprout, but we have not heard of three-year old specimens
germinating. In concluding this subject we will mention
xxvii.] ERGOT. 227
some of the effects of this pest, and say how its general
prevalence may be reduced.
The consumption of ergot when it is ground up with
grain causes chronic dry, black, or livid gangrene, some-
times with but little fever, inflammation, or pain. The
limbs affected become insensible and cold, and, in the
progress of the disorder, dry, hard, black, and withered.
In some instances violent pain has been recorded, with
redness, insupportable heat, and delirium. A line of
separation, reaching to the bone, is formed by the disease
between the dead and living tissues. Ergot is especially
potent, when taken fresh and raw, in exciting strong uterine
contractions ; and when a small portion only is consumed,
it causes weaker contractions which wear out the injured
animal with fatigue. A correspondent of the Gardeners'
Chronicle, 1st June 1876, states that he has observed
cows lose their calves prematurely at the time ergot
appears. It is curious that, when ergot is ground with
grain and cooked in bread, abortion or premature birth is
not the usual result. It has been computed that in wet
seasons, in some parts of France, one-fourth of the rye
used for bread consists of ergot ; the poor, it appears, taking
no pains to separate the ergots, and the result of its con-
sumption is often some form of ergotism. M. Duchamel
mentions, in the Memoirs of the Royal Academy for 1748,
that out of 120 persons attacked scarcely four or five
escaped with their lives.
In the Philosophical Transactions (1763) vol. lii., part
ii., for the year 1762, p. 526, is a printed extract from a
letter from the Rev. James Bones, M.A., of Wattisham,
near Stowmarket, Suffolk, to George Baker, M.D., F.R.S.,
relating to a case of mortification of limbs in a family there.
The letter says that on Sunday, 10th January, Mary,
daughter of John and Mary Downing, sixteen years old,
felt a violent pain in her left leg, which, in an hour
or two, also affected her foot, and particularly her toes.
On the next day her toes were much swollen, and black
228 DISEASES OF FIELD & GARDEN CROPS. [CH.
spots appeared on them. By degrees the whole foot
became swollen and black. The pain, which was now
chiefly in her toes, was, as she said, as if dogs were
gnawing her. The blackness and swelling advanced
upwards by slow degrees, till they reached the knee,
where the flesh broke, and a great discharge followed.
In a little time the flesh of her leg putrified and came off
at the ankle, together with the whole foot, leaving the
leg-bones bare. Her other foot and leg were affected in
a few days, and decayed, nearly by the same degrees and
in the same manner. She had then an abscess formed in
one of her thighs. In a subsequent note it is stated that
this girl, who had sat for fourteen weeks in a chair, and
for seven days without any feet, or flesh on her leg-bones,
had at length consented to have the bones taken off.
Mary, the mother, was seized within a few hours of her
daughter's first seizure with the same violent pain under
her left foot, or, as she sometimes said, in her left leg.
Her toes, foot, and leg were affected in the same manner
as her daughter's ; and in a few days her other leg and
foot suffered in like manner. The flesh of one leg had
separated, and come off at the knee, leaving the bones
bare, which she would not suffer to be taken off. The
other foot had rotted off at the ankle. Her hands and
part of her arms were, from the first attack, without sen-
sation, and her fingers contracted. In a subsequent letter
it is stated that the mother " still remains in bed with
her leg-bones bare, which she will not suffer to be taken
off."
In four or five days after the eldest daughter and the
mother were first affected, Elizabeth, aged fourteen years,
Sarah, aged ten, Eobert, aged six, and Edward, aged four,
were all taken on the same day with violent pains in the
feet and legs, chiefly in the left.
Elizabeth was seized only in one leg and foot, which,
during three weeks, she could not set on the ground, but
stood all the time on the other foot, leaning against the
xxvii.] ERGOT. 229
chimney ; after which, being taken in the same manner
in the other foot, she was obliged to lie down. One foot
mortified and came off at the ankle, and the other leg near
the knee.
Sarah was taken in one foot, which mortified and came
off at the ankle. The other leg suffered in the same
manner, and also separated at the knee.
Robert was taken in both feet. His legs separated at
the knees. . . . Edward was taken in both feet, which
separated at the ankle . . . etc.
The report then describes the death of an infant, whose
hands and feet turned black after death, and the illness
of the father, whose fingers were benumbed, contracted,
and black, the nails coming off, and two of the fingers becom-
ing ulcerated. Then follows a description of the persons
afflicted and their food, but no hint is given as to the
presence of ergot.
In the subsequent letter, before referred to, it states at
page 530 that the family had no rye, but had been " used
to buy two bushels of clog-wheat, or revets, or bearded wheat
. . . every fortnight. Of this they made their household
bread." The wheat, it appears, had been laid, and was
thrashed separately lest it should spoil the samples. It
was not mildewed or grown, but discoloured and smaller
than the other. It made bad bread and worse puddings.
A labouring man who used the bread was affected with a
numbness of both his hands for about four weeks. His
hands were continually cold, and his finger ends peeled.
One thumb was at the time of the report still without any
sensation.
A note at the end of the second letter says "there
is, in L'Histoire de VAcademie Royale des Sciences, for the
year 1710, a paper, the title of which is Sur le bled cornu
appelU Ergot (Secale corniculatum nigrum, mentioned as
a poison by Hoffmann)." Here it is said that M. Noel,
surgeon of the Hotel-Dieu at Orleans, within about a
year's time had received into the hospital more than fifty
230 DISEASES OF FIELD & GARDEN CROPS. [CH.
patients afflicted d'une gangrene seche, noire et livide, which
began at the toes and advanced more or less, being some-
times continued even to the thighs. He adds "he
observed that this disease affected the men only ; and that,
in general, the females, except some very young girls,
were free from it."
In the same paper is mentioned, as a fact well known
to the Academy, the case of a peasant who lived near
Blois. In this patient a gangrene, at its first attack,
destroyed all the toes of one foot, then those of the other,
afterwards the remaining parts of both feet ; then the flesh
of both his legs and that of his thighs rotted off success-
ively, and left nothing but bare bones.
The members of the Academy were of opinion that
the disease (of which M. Noel had sent an account) was
produced by bad nourishment, particularly by bread in
which there was a great quantity of ergot. This substance
is described by M. Fagon, first physician to the king, and
is said by him to be a " kind of monster in vegetation,
which a particular kind of rye sown in March is more apt to
produce than what is sown in the autumn, and which often
abounds in moist, cold countries and in wet seasons."
Professor J. S. Henslow, in commenting on the Mattis-
hall case, in the Journal of the Royal Agricultural Society
of England, vol. ii., 1841, says there was no evidence
that the presence of ergot was suspected in the wheat
used, and although ergot is supposed seldom to attack
wheat, yet Professor Henslow says that he had found it
in 1841 in four different fields of wheat, and gathered
more than a dozen specimens. Some of the Suffolk
farmers were sufficiently acquainted with it to satisfy
Professor Henslow that ergot was more common on wheat
than was at that time commonly suspected. Upon asking
his miller to search, he soon picked out about three dozen
ergots from two bushels of revet wheat which had been
sent to be ground at his mill, and he said that he had left
at least as many more in the sample. This wheat was
ERGOT. 231
grown in the next parish to Wattisham. A very cursory
look into a sack of gleaned wheat then at the mill also
furnished Professor Henslow with three or four more
specimens.
From our own experience we should say that ergot in
wheat is by no means uncommon, as we have generally
found it on searching. Our examples have always been
much smaller than the ergot of rye, and not much larger
than a grain of wheat.
Our friend the Kev. Canon Du Port, M.A., of Mattis-
liall, Suffolk, writing in the Transactions of the Norfolk
and Norwich Naturalists' Society, vol. iii. p. 199, says a
considerable quantity of ergot was found among the
marshland wheats in the year 1879, in which the summer
was abnormally wet and sunless.
It unfortunately happens that ergot is extremely
frequent on the common rye grass, Lolium perenne, L., a
valuable grass, never absent from pasture-land and always
present in permanent pastures. Professor Henry Tanner
states, in the Journal of the Royal Agricultural Society of
England, No. xli., 1858, that he knew of a Shropshire
breeder who lost £1200 in three years from the preval-
ence of ergot in his fields. Ergotised grass is especially
damaging to animals at the time when the uterus is nearly
ready to exclude its contents.
In some instances it is easy to sift ergot out of grain,
as the ergots are larger in size than the seeds ; but in other
instances, as in wheat, the ergots are often so similar in
size with the grain that hand picking is the only means
that can be used. As ergots are generally black, there
is no special difficulty in recognising them amongst
seeds.
It is said that ergot is most abundant in ill -drained
positions, and that good draining materially lessens it.
When the grasses of pastures are ergotised it is well to
pass a sharp scythe over the top of the grass and remove
as far as possible the spikes, racemes, or panicles ; and
232 DISEASES OF FIELD & GARDEN CROPS. [CH. xxvn.
this material, which is a common cause of abortion, should
be raked together and placed out of the reach of the flocks
and herds. In districts notoriously subject to ergot the
scythe may be used in a similar manner at the flowering
time of grasses, for as the spores of the Claviceps attack the
young flowers, it is obvious that the ergots will be many
or few according to the number of grass flowers.
CHAPTER XXVIII.
WILSON'S VARIETY OF CLAVICEPS ON ERGOT.
Claviceps purpurea, Tul., var. Wilsoni, W.Sm.
No account of ergot would be complete without a descrip-
tion of a curious form of the germinating ergot of floating
sweet grass, Glyceria fluitans, RBr., observed by our friend
Mr. A. Stephen Wilson near Aberdeen.
Glyceria Jluitans, E.Br., is common in wet and muddy
places and in stagnant pools and slow-running streams ;
and, in our opinion, the peculiar variety of Claviceps named
Wilsoni entirely owes its origin to its peculiar environ-
ment, so different as it is from the environment of wheat,
rye, and other cereals, and many grasses. Mr. A. S.
Wilson, in July last, obligingly forwarded us a consider-
able number of germinated ergots of Glyceria fluitans,
R.Br., on which the new and curious variety of Claviceps
was growing, and from these examples the following notes
and illustrations have been prepared.
Mr. Wilson detected these growths in July 1882 after
a very wet spring and early summer. In July 1883 they
were less common, and were intermixed with the normal
purple form of the Glaviceps described in Chapter XXVII.
Sometimes the purple form as well as the new white or
yellowish one was growing from the same ergot.
Four germinated ergots carrying Glaviceps purpurea,
Tul., var. Wilsoni, W.Sm., are engraved, natural size, at
Fig. 107. This illustration may be compared with Fig.
100, where the normal form is illustrated, natural size.
Two germinated ergots are enlarged five diameters at
Fig. 108, for comparison with Fig. 101, A, where the
234 DISEASES OF FIELD & GARDEN CROPS. [CH.
normal form is similarly enlarged. The variety differs
from the typical form in being whitish or yellowish
instead of pale purple in colour, and in the perithecia or
Fiu. 107.
Claviceps purpurea, Tul., var. Wilsoni, "W.Sm., growing from Ergot
Natural size.
conceptacles being almost free on an elongated clublike
growth instead of being immersed in a globular head or
stroma. Many of the growths of the variety Wilsoni,
FIG. 108.
Claviceps purpurea, Tul., var. Wilsoni, W.Sm., growing from Ergot.
Enlarged 5 diameters.
W.Sm., are hair-like, others are attenuated upwards from
a thicker base and bear no perithecia. The whole growth is
less firm than the type, and instantly reminds one of an ab-
XXVIII.]
CLAVICEPS ON ERGOT.
235
normal variety. The Claviceps grows from the interior of
the ergot, and bursts through cracks on the surface. Some-
times the crack or opening is very small, and through this
small opening the Claviceps emerges
and speedily produces a matted base
of mycelium upon the surface of the
ergot, before the club is produced.
In some instances the base of spawn
is so thick that the Claviceps super-
ficially resembles a parasite upon
ergot rather than a true fruiting
condition of ergot itself. Some-
times this effused mycelium spreads
over the ergot, and several clubs
arise from one stratum of mycelium,
which may have emerged from one
minute hole or crack in the black
ergot or Sclerotium. The pales of
the grass flower are shown attached
to the ergots in Fig. 108.
The upper part of a club is illus-
trated at Fig. 109, enlarged twenty
diameters, for comparison with the
normal club of Claviceps purpurea,
Tul., illustrated to the same scale
at Fig. 101, B. It will now be
noticed that the perithecia or con-
ceptacles are, on the average, the
same in size and character in both
fungi, the only difference being
that the club is drawn up in the FlG m_Upper part of
variety Wilsoni, W.Sm., and its soft
substance is drawn away from the
perithecia, which are left almost
free instead of being embedded as in the typical C. pur-
purea, Tul. A phenomenon of this class is one of the
commonest in the vegetable kingdom, and is especially
Claviceps piirpurea, Tul.,
var. Wilsoni, W.Sm. En-
larged 20 diameters.
236 DISEASES OF FIELD & GARDEN CROPS. [CH.
frequent in the inflorescences of flowering plants. The
growth is comparable with an umbel lengthening into a
raceme or corymb. Such abnormal growths are especially
common in wallflowers (Oheiranthus) and candytuft (Iberis).
A single conceptacle is shown, enlarged 200 diameters,
at Fig. 110, for comparison with
the normal conceptacle illustrated
to the same scale at Fig. 102.
There is a little difference in the
form, but virtually the two
growths are the same in both
fungi. Tulasne has illustrated
the normal form of G. purpurea,
Tul., precisely like our Fig. 110.
The conceptacle is packed with
asci or bladders containing
sporidia.
A single ascus removed from a
conceptacle, with its contained
sporidia, is illustrated at Fig. Ill,
A, enlarged 500 diameters; and
a single sporidium, enlarged to
1000 diameters, at B, for com-
parison with the same parts en-
graved to the same scale from the
FIG. no. — Conceptacle of normal growth in Fig. 103. A
x'200
no. — Conceptacl
larged 200 diameters.
noted; but this ascus and
sporidium were taken from the
largest and most mature conceptacle we could find —
others were not half the size. Average specimens are
exactly the same in size and form in both fungi.
The ergots from which the variety Wilsoni grew were
from the bottom and shady side of a half-dry Aberdeenshire
ditch. In this position they were found naturally covered
with decaying grass. To us the ergots appeared much less
firm than the ergots from rye, and in some instances they
XXVIII. ]
CLAVICEPS ON ERGOT.
237
were diseased and covered with a
remarkable Saprolegnia, with dis-
tinctly septate mycelium, not unlike
Fig. 24 in this work. To our mind
every fact points to the conclusion
that the variety Wilsoni, W.Sm.,
owes its peculiar conformation to
the wet and shady position in which
it grows.
"We have been particular in the
description and illustration of this
variety, as our friends Messrs.
Plowright and Wilson have de-
scribed this form under another
generic name in the Gardeners1
Chronicle for 9th February 1884.
It is there described and illustrated
as a parasite of ergot under the
name of Barya aurantiaca, P. and
Wils. The authors state that they
have been unable to make the
sporidia reproduce ergot in wheat,
rye, and Poa trivialis, L. ; but this
fact need not cause special surprise,
if the whole growth is regarded, as
we regard it, as a mere monstrous
and abnormal form ; besides, the
experiments should have been made
with Glyceria fluitanS) K.Br., whence
the ergots were derived, and not sob 1 II x • 1000
with wheat, rye, and Poa trivialis,
L. Mr. Wilson originally considered
the new variety to be a mere de-
graded condition of Claviceps pur- FIG. ill. — Claviceps pur-
purea, TuL, and in this original Pu™a, Tui., var. Wilsoni,
determination we consider he was £5^™
correct. enlarged 1000 diameters.
238 DISEASES OF FIELD & GARDEN CROPS. [OH. xxvm.
Dr. M. C. Cooke, M.A., in Grevellia, vol. xii., March
1884, p. 77, has, since the above notes were written,
described Barya aurantiaca, P. and Wils., as a species
under the name of Claviceps Wilsoni, Cke. He says it
differs from all other species of Claviceps in the elongated
clavate capitulum and in the lax manner in which the
conceptacles or perithecia are immersed. "We do not,
however, esteem this form to be worthy of specific rank.
We consider also that the Claviceps microcephala, Tul.,
on reeds is not a species but a mere variety of C. pur-
purea, Tul. One is connected with the other by a con-
tinuous series of intermediate forms, and the typical form
of C. purpurea, Tul., sometimes occurs on the ergots of
reeds. If these two fungi should ultimately prove to be
distinct, hybrids undoubtedly occur, which probably
arise from the spores belonging to the two different forms
of Claviceps lighting on a single grass flower, and there
bursting and producing a hybrid Sphacelia, which is the
beginning of ergot. Or if the spores of the Oidiuni state
of Claviceps purpurea, Tul., and C. microcephala, Tul.,
were blown on to the same grass flower, the watery con-
tents of the burst spores or conidia would conjoin and
form a hybrid Sphacelia.
CHAPTER XXIX.
MILDEW OF PARSNIPS.
Peronospora nivea, Ung.
THE descriptions we have already given of Peronospora
trifoliorum, D.By., P. exigua, W.Sm., P. Schleideniana, Ung.,
and P. parasitica, Pers., more or less hold good with the
present species, which is often described imder the name
of P. umbelliferarum, Gasp. Peronospora nivea, Ung., affects
various umbelliferous plants, but is especially formidable
in its attacks on our garden parsnips. The fungus lives
within the tissues of the invaded plants, and attaches
itself to their constituent cells by minute suckers growing
from its somewhat torulose mycelium. Like other species
of Peronospora it causes putrescence in the plants it attacks,
and sets up decay in the leaves, stems, and roots. This
is very noticeable in the root of the garden parsnip. As
with the fungus of the potato disease, the mycelium often
descends the stem by the interior. In the parsnip the
large fleshy esculent tap roots become spotted and at length
putrid in a way similar with diseased potatoes.
The parasite is very common on the Cow parsnip, or
hogweed, Heracleum, Spondylium, L. ; the wild Angelica,
Angelica sylvestris, L. ; and the Goutweed or Bishop weed,
jfigopodium Podagraria, L.
The general appearance of this fungus is shown in Fig.
112, where a single fruiting-stem or conidiophore is en-
larged 400 diameters. The spores or conidia, as may
be seen, are supported on threads or spicules of extreme
tenuity. The point of junction between the parsnip leaf
and the fruiting branch of the Peronospora is shown at E.
240 DISEASES OF FIELD & GARDEN CROPS. [CH.
X-IOOO
FIG. 112. — MILDEW OF PARSNIPS.
Peronospom nivea, Ung., enlarged 400 diameters ; conidia and zoospores,
enlarged 1000 diameters ; oospore or resting-spore, enlarged
400 diameters.
xxix.] MILDEW OF PARSNIPS. 241
At this point the fungus emerges through one of the
organs of transpiration as shown in Peronospora trifoliorum,
D.By., Fig. 1; P. parasitica, Fig. 29; and P. infestans,
Mont., Fig. 127. A single conidium or spore is farther
enlarged to 1000 diameters at A. As maturity is reached
the vital material within the dull-coloured, non-lustrous,
spores, divides or differentiates itself into several polyhedric
portions, and each of these portions ultimately bursts out
of the conidium as at B. Soon after emergence these little
spores become furnished with two extremely fine and
attenuated vibrating hairs or cilia as at C ; and with these
cilia the spores are able to propel themselves and sail about
with some rapidity over any moist surface. These zoo-
spores are doubtlessly carried from plant to plant by the
wind causing damp diseased leaves to flap against each
other. After swimming about for a short time the
zoospores burst and produce mycelium, which in turn
reproduces the Peronospora after entering the host plant
by the organs of transpiration. It is remarkable that at
a time when little or nothing was known of zoospores in
the Peronosporece that the Rev. M. J. Berkeley wrote in
the English Flora, 1836, under Botrytis crustosa, Fr. (a
synonym of Peronospora nivea, Ung.), — " the spores, very
large and decidedly filled with sporidia, as in Mucor."
See the description of Mucor subtilissimus, Berk., in this
work. Oospores or resting -spores are produced within
the tissues of the host plant ; one of these, enlarged 400
diameters, is shown at I).
Often associated with Peronospora nivea, Ung., especially
in the leaves and stems of ^Egopodium Podagraria, L., a
second fungus named Prot omyces macrosporus, Ung., is found.
The name Protomyces is derived from protos, first, and
mukes, a fungus ; macrosporus^ of course, means large-
spored. The constant occurrence of the Peronospora with
the Protomyces is very remarkable, but until better evidence
is forthcoming we are inclined to look upon the phenom-
enon as a striking example of " consortism." We have
R
242 DISEASES OF FIELD & GARDEN CROPS. [OH.
never seen the Peronospora in anatomical connection with
the Protomyces.
An illustration of Protomyces macrosporus, Ung., is given
in Fig. 113, enlarged 400 diameters. These bodies, which
'are true oospores or resting-spores, have two or three mem-
branes, and when small in size, as at C, are hardly to be
distinguished from the resting-spores of Peronospora nivea,
Ung. When, however, they are of large size (and they
vary greatly), and furnished with a dissepiment across
the inner or endospore, as at B, they are unlike the usual
resting-spores belonging to Peronospora. These bodies
A B
FIG. 113.
Protomyces viacrosporus, Ung. Enlarged 400 diameters.
occur in the intercellular spaces within the leaves and
stems of umbelliferous plants, often in large companies.
They not only cause decomposed spots to appear, but they
distort and distend the affected parts in a remarkable
manner ; so much is this the case that the presence of the
fungus within is easily detected by the knotty swellings
they set up. As the spores reach maturity they cause the
tissues of the leaf to rot, and by this means they reach the
ground by falling from the decayed places. They doubt-
lessly rest on and in the ground in a hybernating state
till the following spring or summer. Mr. Berkeley says
the Protomyces is probably one form of fructification of the
Peronospora, In. this he agrees with Caspary ; but De Bary
xxix.] MILDEW OF PARSNIPS. 243
states that tlie mycelium belonging to the two fungi is
different, and that the oospores of Protomyces contain great
numbers of very minute slightly oscillating spores, which
escape from the expelled inner membrane as in Cysto-
pus, conjugate in pairs like the conidia of the Smut
fungus of corn, and produce zygospores, from zygos, a yoke,
— in reference, we presume, to their supposed pairing.
The zygospores on germination are said to be capable of
penetrating, by their germinal filaments, the epidermis of
the plant invaded exactly after the manner of Peronospora.
The mycelium is said also to be only capable of effectually
growing when within the special host plants of the Proto-
myces. In this position only are the oogonia and oospores,
which at length bear the conjugating spores, produced.
This habit is also the same with that of Peronospora. The
so-called zygospores are minute transparent oscillating
bodies without cilia ; they are about the same in size with
(or perhaps a little smaller than) the zoospores produced
by the conidia of Peronospora nivea, Ung., as illustrated at
B, C, Fig. 112. True zoospores sometimes do not produce
cilia, and it is by no means uncommon to see zoospores
conjoined by a short band. The oospores burst and dis-
charge the little conjugating zygospores in spring, again
reminding one of the habit of Peronospora. Although we
are inclined to esteem the oospores of Protomyces as differ-
ent from the oospores of Peronospora nivea, Ung., yet there
is evidently room for a different opinion. New observa-
tions are needed. Prof, de Bary's paper, Beitrage zur
Morphologie und Physiologie der Pilze, was published in
1864.
Peronospora nivea, Ung., produces resting-spores within
the stems and roots of the plants attacked (they occur in
company with the oospores of the Protomyces) ; therefore
all plants which have been attacked by these fungi should,
as far as possible, be burned. As both the Peronospora
and the Protomyces are peculiar to umbelliferous plants,
parsnips should not be grown a second year in the place
244 DISEASES OF FIELD & GARDEN CROPS. [CH. xxix.
where other umbelliferous plants have been invaded by
mildew, as the resting-spores will be on and in the ground
ready for attacking other umbelliferous plants on their
germination in spring.
The mildew of parsnips has been seen on baldmoney,
Meum Atkamanticum, Jacq. ; on burnet saxifrage, Pimpi-
nella Saxifraga, L. ; P. Anisum ; parsley, Petroselinum
sativum, HofFm. ; the garden and wild chervil, Antbriscus
Oerefolium, Hoffm. ; and Chcerophyllum sylvestre, Linn. ;
and hemlock, Conium maculatum, Linn.
The common red -rust and black mildew of celery,
another umbelliferous plant, is caused by Puccinia apii,
Corda, and its Uredo, U. apii, Wall., a fungus closely allied
to the rusts and black mildew of corn. This fungus has
been proved to be hereditary, and its plasma capable of
being transmitted in celery seeds. Celery is sometimes
infested with Puccinia heraclei, Grev., a parasite frequent
on cow-parsnip or hogvveed. The Umbelliferee are unusu-
ally subject to attacks from fungi.
CHAPTEE XXX.
BUNT OF WHEAT.
Tilletia Caries, Tul.
THE disease of wheat generally known as bunt is recog-
nised in some districts as pepper - brand, smut balls,
bladder - brand, stinking -smut, stinking -rust, and even
smut. Its ravages are almost confined to cultivated
wheat ; it rarely occurs on barley. A distinct species
occurs on wheat in the United States. The scientific
name of the fungus which causes bunt in wheat is Tilletia
Caries, Tul. Tilletia is named after Matthieu Tillet, who
wrote the Dissertation sur la cause qui corrompt et qui
noiriet les grains de bldd dans les e'pis, Bordeaux, 1755, and
a similar work published in Paris in 1755. Caries means
rottenness or decay. The meaning of the popular name,
bunt, is very obscure. We have the word bunter, which
means an offensive person (woman), and the verb bunt, to
swell ; the former may be a cognate derivative. Dr.
Murray of Mill Hill informs us that the word bunt is
used, for a fungus in the same way as touchwood, by
writers of the seventeenth century. This is perhaps the
most likely origin. The cavity or belly of a sail is called
the bunt, and the material of the sail bunting. The
bellying part of a seine-net is also called the bunt, which
name may have been transferred to the blight in reference
to its fishy smell. Bunt may be a corruption of burnt.
In the fields it is difficult without experience to dis-
tinguish bunted from sound wheat ; there is very little
indeed to indicate the presence of the hidden foe ; this
is why the disease is so dreaded by farmers. As in
246 DISEASES OF FIELD & GARDEN CROPS. [CH.
many other fungoid ailments the fungus appears to
excite an abnormal growth of chlorophyll, and the
spikes of affected plants are commonly greener than the
sound ones. Even on examining the ears it sometimes
happens that but little seems amiss ; it is not until the
glumes and pales are pushed aside that the dark diseased
seeds . become visible. Sometimes the bunted grains
burst whilst still in the ear, and the escaped spores stain
the glumes and pales a dark colour. Practised eyes
readily detect these slight black stains.
A bunted grain of wheat is illustrated at Fig. 114, A,
B
X-5
FIG. 114.— BUNT OF WHEAT.
Grains of wheat destroyed by Tilletia Caries, Tul.
Enlarged 5 diameters.
with a transverse section at B, and a longitudinal section
at C, enlarged five diameters. Reference may be made
to Fig. 42 for a normal grain of wheat, and to Fig. 45
for an example of ear-cockle, all engraved to the same
scale. The external appearance of bunted grains of wheat
is different from healthy grains. Bunted wheat seeds are
shorter and wider than healthy ones ; they are dwarfed
in height and distended in width, and generally somewhat
pointed towards the base. Instead of being pale buff in
colour, they are of a somewhat dark, dull green tint.
They are frequently cracked, as shown at A, and from
xxx.] BUNT OF WHEAT. 247
these cracks a black powder emerges. On cutting affected
seeds in two, the outer coat of the grain is found to be
weak and brittle, and its whole inner substance a mass of
black powder, which has replaced the natural inner farina-
ceous material of the grain. On crushing bunted wheat
between the fingers the black pulpy powder feels soft and
greasy, and a foetid odour resembling decaying fish is
dispelled ; hence the popular name, — stinking-rust. One
of its old botanical names is Uredo fcetida, Baeur. Bunted
grains do not occur as isolated examples in the ear ; the
rule is that every grain in an affected ear is bunted.
When these bunted grains are ground into flour their
FIG. 115.
Spores of the Bunt fungus, Tilletia Caries, Tul.
Enlarged 400 diameters.
presence is made known to the miller, not only by the
black streaks they cause in the white flour, but also by the
disgusting odour which arises at the time of crushing.
The black powder, when placed under the microscope
and magnified 400 diameters, is seen as at Fig. 115, one
mass of beautiful brownish spores, with a few fine mycelial
threads, to which some of the spores will be seen still
attached ; the supporting threads are best seen in young
examples, for as the fungus approaches maturity, the
threads break up into dust and perish. The spores are
spherical, or sometimes slightly oval, reticulated and
slightly spinulose, reminding a microscopist of pollen-
grains belonging to the Campanulacece, the colour of
course being different. They are so small that a single
248 DISEASES OF FIELD & GARDEN CROPS. [CH.
FIG. 116.— BUNT OF WHEAT.
Spores of Tilletia Caries, Tul., germinating.
Enlarged 1000 diameters.
grain of wheat is large enough to contain 4,000,000 of
them.
BUNT OF WHEAT. 249
If these spores are kept in moist air or on a wet surface
for three or four days, they will germinate, as at Fig. 116,
A, enlarged 1000 diameters. The epispore bursts, and a
thick septate tube is protruded. This tube, after it has
grown to three or four times the diameter of the spore,
forms a sort of small terminal crown, and on the minute
papillas of this crown it bears four, eight, or ten rocllike
sporidia, as shown at B. When these sporidia are fully
grown, and whilst still adherent to the apex of the germ-
tube, they coalesce, as at CO, by means of short transverse
tubes. When these conjugated bodies drop from the
supporting tube they germinate and produce secondary
sporidia of a different form, as shown at DD. At times
the supporting threads of these conidia are extremely long
and jointed throughout ; at other times there is no sup-
porting thread, but the conidium may grow from the end
of a secondary spore. Sometimes the sporidia are pro-
duced whilst the secondary spores are still attached to
their supporting stem. A spore or conidium so growing
is illustrated at G. The conidia, in turn, are capable of
germination and the production of conidia of the third
order, as at EE. Sometimes they so germinate without
the spores, DD, falling from their attachment. When
the secondary and tertiary sporidia, as at D and E, ger-
minate, they produce a septate thread of extreme tenuity,
as at F, and on this thread the bunt spores are at length
borne, as illustrated in Fig. 115. Sometimes bunt
spores do not produce, on germination, the minute crown-
like terminal cell, with its conjugating secondary spores ;
but the thick germ-tube grows for a great length, branch-
ing and rebranching, and all the time forming septa.
The vital material is chiefly confined to the terminal end
of each branch. When this mode of growth takes place,
conjugating spores are never formed.
In some instances the secondary spores become con-
joined in two places instead of one, as illustrated by Mr.
Berkeley, and confirmed in two instances by Dr. Oscar
250 DISEASES OF FIELD & GARDEN CROPS. [CH.
Brefeld ; in other cases three secondary spores may become
conjoined, and it is not uncommon to see the conjugating
band itself burst and produce a mycelial thread.
Under favourable conditions a few germinating second-
ary spores will form a dense involved mass of mycelium,
bearing a vast number of conidia of the first and second
generation ; and under the microscope these may be seen
in all stages of growth. When the mycelium for any
reason ceases to grow, a crop of conidia is at once pro-
duced. Sometimes the mycelial threads become furnished
with an enormous number of short joints or constrictions,
giving the threads a necklace-like appearance. Ultimately
this chain or necklace breaks up into separate joints, and
each joint acts as a conidium. Each conidium thus
formed in a chain is capable of producing other necklace-
like growths of conidia.
An elaborate essay on bunt and smut was published in
Paris in 1877 under the name of Aperfu SysUmatique des
Ustilaginfos, by Alexandre Fischer de Waldheim. This
author at one time advocated the idea of bunt and smut
fungi living in two forms on different plants in the sup-
posed style of corn mildew on the barberry bush, because
he failed to infect corn experimentally, and because he
had probably learned that " experience had taught the
practical farmer " that (according to Phillipar) the barberry
bush, a stinking plant, when in bloom, was in some places
the cause of bunt. No experiment is, however, easier
than the artificial and direct infection of wheat with bunt
and smut.
Wheat becomes affected with bunt by the spores of the
fungus being sown with the grain. The spores do not
germinate whilst they are dry and stored with the seed,
but in and on the damp ground after the grain has been
planted. The whole series of changes illustrated in Fig.
116 takes place on and in the ground, and when the
attenuated thread at F is produced, it readily finds its
way, aided by its inconceivable fineness, into the tissues
xxx.] BUNT OF WHEAT. 251
of the young wheat plant by entering the first-formed
organs of transpiration in the infant plant. The spores
themselves do not, of course, enter the stomata, and the
germ-tubes probably do not attack the rootlets unless the
latter are broken or injured, although it has been said
by Le Maout and Decaisne, in their General System of
Botany, that the spores can pierce the tissues of the
roots. The germ -tube, when once within the infant
plant, speedily ascends the stem. It is now by no
means difficult to trace the course of the mycelium up
the shaft of the affected plant, and an instance has been
recorded by the Rev. M. J. Berkeley in vol. ii. of the
Journal of the Royal Horticultural Society, 1847, where a
streak of bunt appeared upon the outside of the stem of a
wheat plant. Mr. Berkeley was the first to publish a
description and illustration of germinating bunt spores,
with the conjugating spores borne on the germ-thread.
This was seven years before the publication of L. E.
Tulasne's memoir in the Annales des Sciences Naturelles,
Botanique, series iv., vol. i., 1854. Mr. Berkeley,
although at first inclined to look upon these conjoined
bodies as having something to do with the reproduction of
bunt — the appearances seen in some Algce indicating this to
him — abandoned this first and correct opinion for the idea
that the conjugated bodies were parasites of bunt. He
described the growth as a Fusisporium, partly owing to
the septate spores, under the name of F. inosculans. It is
curious that Tulasne illustrated the conjoined spores of
germinating bunt as non-septate, and therefore unlike a
Fusisporium, and since 1854 this view appears to have
been generally accepted as correct ; but last year Dr. Oscar
Brefeld, in his elaborate work, Botanische Untersuclmn-
cjen uber Hefenpilze, correctly illustrate the germinating
secondary spores as furnished with septa, sometimes three
and sometimes four, precisely in the style of Fusisporium.
This observation, the correctness of which we are able to
confirm, proves the accuracy of Mr. Berkeley's observa-
252 DISEASES OF FIELD & GARDEN CROPS. [CH.
tion in 1847. The septa are best seen in the spores
which have germinated and borne conidia, and such as
have lost their vital material, as at DD, Fig. 116. For
spores of Fusisporium illustrated to the same scale as
germinating bunt, see Figs. 10, 11, 92, 94, and 96 in
this work.
It is easy to prove that bunt in wheat is propagated by
the spores of the fungus, for if wheat seeds are dusted with
the spores or watered with water containing spores, every
wheat plant will come up bunted ; whereas neighbouring
plants, if not so treated, will come up free from disease.
Bunt spores are said (perhaps on insufficient grounds)
to be more or less injurious if mingled with flour and
made into bread. We have frequently seen them in flour
and bread, together with spores of Urocystis and other
fungi. Bunt spores were, we believe, at one time sup-
posed to be the cause of cholera, because they were found
in cholera evacuations. Professor Hallier has erroneously
referred cholera to the presence of Urocystis occulta, Pre.,
a fungus common in Britain on rye, as well as to bunt, as
may be seen by his Phytopathology, and the reports of Drs.
Cunningham and Lewis in the Lancet of 2d, 9th, and 1 6th
January 1869. Fowls have been fed with bunted wheat
without any bad result.
A fungus allied to bunt, but still nearer to smut, and
named Ustilago grand is, TuL, is said to cause headache
and other bad symptoms amongst the men engaged in
cutting reeds for thatching, in consequence of their inhal-
ing the abundant spores. The same fungus is said to
cause eruptions on the face amongst the labourers of the
South of Europe.
When bunt is known to be amongst seed grain it
should be washed or steeped in some weak poisonous solu-
tion, as the minute spores from bunted grains adhere to the
healthy seeds. Water, salt, quicklime slacked with boiling
water ; sulphate of copper, a quarter of a pound to a bushel
of corn, and sulphate of soda have all been recommended.
xxx.] BUNT OF WHEAT. 253
Sulphate of soda in solution and the seeds afterwards dried
with dusted quicklime is said to be one of the best
preventive solutions. The lime combines with the soda
and forms sulphate of lime or gypsum, whilst caustic
alkali is set free. As the spores are lighter than water,
mere steeping in brine or even pure water is often effectual,
as the spores float, and are easily washed away. It is
probable that the presence of a few scattered greasy spores
are quite as, if not more, damaging than the whole bunted
grains with unbroken seed coats. Some alkaline ley
should be added if water is used, as the oil on the surface
of the spores combines with the alkali and forms a soapy
substance which is fatal to effectual spore germination.
Sufficient permanganate of potassium may be added to the
water until it becomes rose-coloured, or one per cent of
carbolic acid may be mixed with the water. It is not
proper for the seed to remain long in these solutions ;
they should be washed quickly and then allowed to dry.
When millers see bunted grains amongst the wheat they
generally pass it through a dresser with a strong exhaust,
and this draws away the foetid spores.
Bunt, Tilletia Caries, TuL, seems to be confined in this
country to wheat, Triticum vulgare, Vill. ; T. sativum,
Lam. ; and barley, Hordeum distichum, L. Alex. Fischer
de Waldheim describes no less than fourteen species of
bunt. Some of these, as T. Lolii, Awd., and T. Icevis,
Kuehn. ; the first recorded on three different species of
Loliutn, and the second on five species of Triticum,
including wheat, probably occur in this country. One
species attacks five different species of Agrostis, and several
other bunt fungi are confined to grasses.
A curious species of bunt named Tilletia bullata, Fl.,
has recently been found in ' Scotland on docks, Eumex
obtusifolius, L. On the Continent this bunt attacks
different species of Polyyonum.
CHAPTER XXXI.
SMUT OF CORN.
Ustilago carlo, TuL
THEEE are few fungi more familiar to agriculturists than
the common " smut " of corn, so common on wheat, bar-
ley, and especially oats. Its black colour, and its profusion
of sooty spores produced on impoverished ears of corn,
makes it apparent to the least observant. It appears
earlier in the season" than bunt, with which it is some-
times confounded by rustics. It is in some places called
"bunt ear," "blackball," "dust brand," and "chimney
sweeper." Its scientific name is Ustilago carbo, Tul. The
generic name is derived from Ustio, a burning, and carbo,
charcoal, in reference to the burnt and sooty appearance
of the diseased panicle or spike. Farmers look on this
fungus with less dread than the fungus of bunt, perhaps
because the last is virtually a hidden foe and may cause
unexpected sudden and serious loss, whilst smut is always
seen, and indeed, makes itself obtrusively apparent. In
some districts the loose smutted panicle of oats is ignor-
antly termed the male plant, the spores of the fungus
being esteemed as black pollen, and so more beneficial
than harmful. As there is no disgusting odour belonging
to the " smut" fungus, it does not spoil the flour to an
equal extent with the Tilletia. It is probably not
very injurious if taken in food by animals, as fowls
are not injured by eating smutted grain. It has,
however, been said that the straw of corn that has
been infested with the smut fungus is distasteful to
cattle in chaff. In bad cases as much as one -third of
CH. XXXI.]
SMUT OF CORN
255
FIG. 117.
Panicle of oats invaded by the fungus of Smut, Ustilago carbo, Tul.
Natural size.
256 DISEASES OF FIELD & GARDEN CROPS. [CH.
the crop has been destroyed by the presence of the
smut fungus.
The general appearance of a panicle of oats affected
with smut is illustrated, natural size, at Fig. 117. In the
field it is evident that (as with bunt) the disease springs
direct from the root, for it is common to see numerous
stems, and every one diseased, springing from one base. It
is equally clear that the disease traverses each stem from
the bottom upwards, as the lower spikelets of the young
FIG. us.
Part of a spikelet of oats with Smut fungus, Ustilago carbc/j Tul.
Enlarged 5 diameters.
panicle are the first to show the disease, as illustrated in
Fig. 117. Nothing is more common than to see the basal
branches of the panicle with their spikelets diseased and
the upper part of the panicle presenting the normal
healthy appearance.
Part of a spikelet only slightly affected with the smut
fungus is illustrated in Fig. 118, enlarged five diameters.
AA are the glumes ; B, outer glumelle, sometimes fur-
nished with a beard or awn ; C, inner glumelle ; D, abor-
tive flower ; E, the pistil or grain. The interior of the
SMUT OF CORN. 257
diseased grain at first presents a whitish viscous mass,
which is produced at the expense of the tissues and juices
of the invaded plant. This viscid mass at length exhibits
a structure of closely packed cells filled with a homo-
geneous mass of minute granules ; the cell walls ulti-
mately disappear, and the whole contents of the invaded
organ appear as one black dusty mass of simple smooth
spores. The disease is, however, by no means confined to
the grain, as it attacks every part of the panicle with its
spikelets, including their slender stems, with equal viru-
X 25
FIG. 119.
Fragment of glume of oats, showing burst pustules of the Smut fungus,
Ustilago carbo, Tul. Enlarged 25 diameters.
lence, till at last the glumes and other parts are left as
transparent ragged fragments thickly coated with the
masses of black spores, which have been produced at the
expense of the normal tissues of the spikelet. If a frag-
ment of an infected glume is now taken, and enlarged
twenty-five diameters, it will be seen as at Fig. 119.
This shows (horizontally) four of the fine ridges and
three minute furrows, which can be almost seen with
the unaided eye on every glume belonging to oats.
The fungus bursts through the glumes and other structures
258 DISEASES OF FIELD & GARDEN CROPS. [CH.
on both sides from within outwards, and as the spores in-
crease in number they tear the epidermis of the invaded
part to shreds, which shreds are, with the spores, speedily
carried away by the wind. The pustules illustrated at
Fig. 119 may be compared with the sori of a similar
nature belonging to Puccinia Bubigo-vera, B.C., Fig. 67,
and Puccinia graminus, Pers., Fig. 78. The spores are
brownish-black in colour, roundish, and extremely small,
as illustrated, enlarged 400 diameters, at Fig. 120. If the
spores of bunt are now re-
ferred to, as illustrated to the
same scale at Fig. 1 1 5, it will
be seen how different they
are in size. The spores of
the smut fungus are so ex-
X-4OO ' tremely small that it would
FIO. 120.— Spores of the Smut fun- take nearly 25,000,000 of
gus, Ustilago carto, Tul. Enlarged t^em fa cover a superficial
400 diameters. . , ml_ -,
square inch. They are also
quite smooth, wliilst the spores of the bunt fungus are
reticulated and spinulose. Dr. Oscar Brefeld, in his work,
Botanische Untersuchungen ilber Hefenpilze Fortsetzung der
tSchimmelpilze, Part V., states that some kinds of yeast are
nothing more than conidial or larval forms of smut
fungi. The reproduction of smut in the yeast condition
continues for an indefinite period, so long as the spores
are kept upon a suitable nutrient matrix. Upon this
nutrient matrix they always remain in the yeast, conidial,
or larval condition. If these views are correct they indicate
that smut fungi are only truly parasitic during the perfect
condition of their existence, and when in the yeast condi-
tion lead a non-parasitic life.
In Fig. 121 the spores of Ustilago carbo, Tul., are shown
in different stages of germination enlarged 1000 dia-
meters. The extreme smallness of these spores is very
marked when compared with the large spores of Peronospora
tSchleideniana, Ung., Fig. 15,0; Puccinia graminis, Pers.,
XXXI.]
SMUT OF CORN.
259
Fig. 80, and others engraved to the same scale in this work.
Germination in water commences in less than twelve hours,
as at 1 ; as germination advances buds or conidia are
formed, as at 2, A A ; farther growth is seen at 3 and 4,
FIG. 121.
Spores of the Smut fungus, Ustilago carbo, Tul. , germinating.
Enlarged 1000 diameters.
with fused or conjoined cells at B. This fusion is similar
with the conjugation seen in Tilletia. Still more advanced
growth is shown at 5 and 6. The long germ-tubes like
the one illustrated at 5, commonly fuse with other germ-
260 DISEASES OF FIELD & GARDEN CROPS. [OH.
tubes, and the two fused tubes will giv^ rise to a third
common conidium-bearing tube. Sometimes this tube is
extremely long and fine, and furnished with numerous
joints throughout its entire length. The conidia, as pro-
duced by the germ-tubes, are capable of producing other
conidia by budding, till at last large colonies of conidia
are produced in a yeast- like condition, free from the
original Ustilago spore and its septate germ tube. The
conidia, which are very unequal in size, germinate as at 7
and 8. Germination may take place at one or both ends
or one or both sides, and the germ-tubes may fuse with
each other at any point of contact. Sometimes the germ-
tubes produced by the conidia are of almost inconceivable
fineness and attenuation, but always more or less jointed
throughout their entire length ; the contained vital ma-
terial is generally most abundant at the growing end.
Sometimes, when growth has quite ceased, owing to
unfavourable conditions, it is renewed with vigour under
changed conditions, and new colonies of yeast-like conidia
are formed, either from old threads or old conidia.
In the allied fungus named Ustilago antherai'um, Fr.,
the conjugation of cells is very clearly seen, and lateral
conidia are produced, as in Tilletia, Fig. 116, D, G. This
parasite is common on the flowers of Silene, Lychnis, etc.,
in fields and hedge sides, reducing the anthers with their
pollen to black dust. The conjugation is also very dis-
tinct in the spores of Ustilago longissima, Tul. ; common
on Poa aquatica, L., and P. fluitans, Scop. No doubt the
so-called conjugation of cells is potential in all species of
Ustilago and its allied genera.
We recently observed the germination of the spores of
Ustilago carbo, Tul., under curious and natural conditions.
"We placed a large number of spores on the top of a
flower-pot filled with moist white sand to note whether
the spores would be readily filtered through the mass by
rain. They did so filter in large numbers ; but many of
the spores germinated on the top stratum of sand, and
SMUT OF CORN. 261
vast quantities of the transparent conidia were filtered
through the sand on to the plate below ; and in this posi-
tion, beneath the bottom of the flower-pot and in the plate,
they formed colonies of yeast-like conidia, and these conidia
germinated by producing threads, as we have illustrated
them in Fig. 121, 7 and 8. The spores germinate very
readily and produce yeast colonies in diluted beer and
diluted expressed juice of horse-dung.
The disease is doubtlessly propagated by the spores of
the fungus being blown over the fields and absorbed by
the earth, and by the fungus spores which adhere to the
seed at the time of sowing. The black spores germinate
in the ground, and there produce the secondary and
tertiary series of transparent spores illustrated in Fig.
121. These spores or conidia of the second and third
order at length protrude extremely fine germinal threads,
and these threads find their way into the earliest
produced stomata of the infant plant. The spawn or
mycelium then travels up the stem towards the panicle
and attacks the lowermost spikelets first. We have
never noticed the upper part of a panicle diseased whilst
the lower part has been sound. It may be commonly
noticed that every stem, from two to eight or more, on
an invaded plant, will show the disease, whilst adjoining
plants remain free. The evidence therefore seems com-
plete that the infection comes from the ground and
travels upwards. It is obvious, therefore, that smut can
only be prevented by dressing the seed, as in the case of
bunt, and the directions for one apply to the other. A
remedy against smut, much in favour in the north of
England, and one which is said to never fail, is the pre-
paration of the seed, immediately before sowing, with a
sprinkling of stale urine, the seeds being afterwards raked
in powdered quicklime till the seed is white. Sometimes
the seed is prepared with vitriol or sulphate of copper
solution, or " bluestone " dissolved in boiling water. One
pound of " bluestone " dissolved in five quarts of water
262 DISEASES OF FIELD & GARDEN CROPS. [CH. XXXT.
is sufficient for a sack of four imperial bushels. The
wheat is soaked for ten minutes, or the ten pints of solu-
tion may be poured over till all is absorbed.
Bunt, Tilletia Caries, Tul., is almost confined to two
species of Triticum ; but as smut invades many grasses
found in pastures and waysides, the disease is clearly often
nursed by the weeds. This fact points to the necessity
for clean and careful farming. All smutted grasses, as
well as the smutted panicles of oats, wheat, and barley,
should be gathered in their earliest recognisable stages
and burnt. A few common -sense hints given to the
labouring men and boys would often save the employer
from great losses.
The smut fungus Ustilago carbo, Tul., has been met
with on the following grasses : — Andropogon hirtus, L. :
Cynodon Dactylon, L. ; hair grass, Aira ccespitosa, L. ;
oats, A vena sativa, L. ; yellow oat grass, A. flavescens, L. ;
downy oat grass, A. pubescens, L.; Arrhenatherum aven-
aceum, Beauv. ; species of Melica ; fescue grass, Festuca
pratensis, Huds. ; Brachypodium ciliatum, P.B.; wheat,
Triticum vulgare, Vill.; T. turgidum, L.; barley and
barley grasses, Hordeum distichum, L.; H. murinum, L. ;
H. vulgare, L. ; rye, Secale cereale, L. ; rye grass, Lolium
perenne, L. ; darnel grass, L. temulentum, L. ; rice, Oryza
saliva, L. ; and .no doubt other plants. The different
species of millet or sorghum, which in the south of
Europe and some parts of Asia are grown for bread in
the place of the oats and barley of northern Europe, are
sometimes badly smutted. Sorghum vulgare, P., and Setaria
italica, Beauv., are the chief food plants affected in India,
Arabia, Asia Minor, Spain, and Italy.
Dr. Maxwell T. Masters, F.R.S., has recently detected
a new British smut fungus, Ustilago Kiihniana, Wolff, on
Rumex Acetosa, L. ; and a second new species, U. Candollei,
Tul., has been found on Polygonum.
CHAPTER XXXII.
TARE OR VETCH, AND PEA MOULD.
Peronospora vicice, Berk.
LIKE several other species of Peronospora described in
this work, P. vicice was first detected by the Rev. M. J.
Berkeley, and described and illustrated by him in vol. i.
of the Journal of the Royal Horticultural Society in 1846.
This parasite is frequent on the tinder surface of the
leaves of tares or vetches and field and garden peas.
Although allied to Peronospora trifoliorum, D.By., as illus-
trated at Fig. 1, and P. exigua, W.Sm., Fig. 2, it is
clearly distinct from both. Like the two plants just
described, it grows within the foliage and causes brownish
downy patches on the leaves and putrescence of the
tissues. The conidiophores of the fungus grow in
clusters, the spores are supported on long extremely
slender spicules, and are tinted with a dull gray
colour. Peronospora vicice, Berk., is illustrated, enlarged
400 diameters, at Fig. 122; a single spore or conidium
is farther enlarged to 1000 diameters at A. The spores
on germination do not burst at the apex, but the germ-
tube is generally protruded from the side. Damp close
weather greatly favours the extension of this fungus, and
dry weather retards its growth. It unfortunately happens
that late peas are sometimes quite destroyed by another
fungus named Erysiplie Martii, Lk., whose growth is
favoured by dry weather and retarded by wet. When
both fungi are present on one crop the destruction of
peas is complete.
The resting- spores are brown, and at length reticulated
264 DISEASES OF FIELD & GARDEN CROPS. [on.
X - 1000
FIG. 122. — TARE OR VETCH, AND PEA MOULD.
Peronospora viciw, Berk. Enlarged 400 diameters.
Conidium, A, 1000 diameters ; Oospore, B, 400 diameters.
by the shrinking of the outer coat as illustrated, enlarged
400 diameters, at B. All diseased and decaying tare and
xxxii.] TARE OR VETCH, AND PEA MOULD. 265
pea refuse should be gathered together and burnt, for it
is in this material that the resting-spores hibernate during
the winter, and burst into new growth in the early spring.
Resting-spores are not destroyed by passing through the
stomach of an animal as food. P. viciw, Berk., is a dis-
tinctly early species as distinguished from P. infestans,
Mont., the fungus of the potato disease, which is a dis-
tinctly late one. Being early, the late crops of peas are
seldom affected by this parasite ; they are, however, fre-
quently destroyed by the fungus next described.
We have seen P. vicice, Berk., growing within the pods
of garden peas and upon the contained seeds.
This parasite has been detected on the bush- vetch,
Vicia sepium, L. ; the common vetch, V. Sativa, L. ;
slender vetch, V. tetrasperma, Moench. ; Lathyrus macror-
rhizus, Wimm. ; and on many species of Pisum and Orobus.
, CHAPTER XXXIII.
PEA MILDEW.
Erysiphe Martii, Lk.
THE description given of Erysiphe graminis, B.C., the
blight or mildew of grass, applies generally to Erysiphe
Martii, Lk., of peas, beans, Umbelliferae, etc. The species
is named in honour of Martins, the famous botanist, and
is the same with E. Pisi, Grev.
The general appearance of the fungus, as seen upon a
r
FIG. 123.— PEA MILDEW.
Conceptacle of Erysiphe Martii, Lk., on leaf of pea.
Enlarged 100 diameters.
leaf of the pea under the microscope, is shown, enlarged
100 diameters, in Fig. 123. The dark body in the
centre of the illustration is the conceptacle of the fungus,
with its jointed tentacle-like appendages. The involved
pattern in outline shows the epidermis of the leaf with its
CH. XXXIII.]
PEA MILDEW.
267
FIG. 124.
PEA MILDEW.
siphe Martii, Lk.
Enlarged 500 dia-
meters.
numerous stomata as at A A A. The conceptacle or peri-
thecium is distinctly smaller in size than in E. graminis.
D.C., as illustrated in Fig. 57, and the cells forming the
bark of the conceptacle are larger. Each
conceptacle contains four or eight spor-
angia or spore cases, and each sporangium
carries four to eight sporidia, as shown
at Tig. 124, enlarged 500 diameters ;
this illustration may be compared with
Fig. 60. The conceptacles arise from a
dense woven mass of very fine white
mycelium, omitted in Fig. 123 in
favour of the epidermis of the leaf with
its stomata. This mycelium is woven
all over the stomata, and so one of the Sporangium or ascus,
chief vital functions of the leaf, the with spores, of .Ery-
transpiration of vapour, is arrested.
The fungus grows on both sides of the
leaves.
This destructive blight of peas invariably invades the
late varieties, and is especially virulent in dry seasons.
In small gardens the attack of the fungus may be pre-
vented by keeping the peas well supplied with water.
This treatment, however, cannot be adopted in the fields,
and watering favours the growth of the Peronospora last
described. Sometimes late peas are so badly attacked by
this fungus that they appear as if thickly dusted with
powdered chalk, and on the white surface thus formed
the innumerable black conceptacles of the Erysiplie may
be readily seen with the unaided eye. An attack of this
fungus generally stops the growth of the invaded plants,
and makes the production of pods impossible. The
mycelium is provided with the minute suckers termed
haustoria, and these haustoria pierce the epidermis of the
attacked plant and cause decay. We have seen this
fungus growing with its conceptacles inside the pods of
peas.
268 DISEASES OF FIELD & GARDEN CROPS. [CH.XXXIII.
It unfortunately happens that this parasite is not con-
fined to peas ; it sometimes grows on beans and melilot,
plants belonging to the same family with the pea. It is,
however, able to support itself on plants of St. John's
wort, Hypericum; some umbelliferous plants, and the
meadow-sweet, Spiraea Ulmaria, L. It is thus capable of
growing effectually upon plants belonging to four different
natural orders.
The only known preventive against attacks of this pest
is the destruction by fire of all invaded material. The
fungus is preserved in decaying refuse ; in this material
the little brown conceptacles remain intact during the
winter, and in the following summer they burst, and each
example discharges about fifty little living transparent
spores, as shown in Fig. 124, ready to grow on the leaves
of peas, weave a spider-weblike mycelium over the surface,
and pierce the epidermis.
CHAPTEE XXXIV.
LETTUCE MILDEW.
Peronospora ganglioniformis, Berk.
THE putrefactive fungus of lettuces was detected by the
Rev. M. J. Berkeley, and described and illustrated by him
in vol. i. of the Royal Horticultural Society, 1846. The
specific named ganglioniformis refers to the resemblance of
the fruiting-threads of the fungus to the natural enlarge-
ments termed ganglions in the course of a nerve.
Tulasne thought this parasite was a mere variety of P.
parasitica, Pers., as found on cabbages and illustrated in
this work at Fig. 29 ; but a reference to our illustration
of P. ganglioniformis, Berk., enlarged at Fig. 125 to 400
diameters, as compared with P. parasitica, Pers., enlarged
in Fig. 29 to 200 diameters, will show how distinct the
two are from each other. Professor de Bary disapproved
of Mr. Berkeley's specific name ganglioniformis, and substi-
tuted gangliformis for it, considering the latter more
correct, but no alteration was required. Had it been
necessary, the word gangliiformis, as printed by Dr. Max
Cornu, would be most correct. In both P. ganglioni-
formis, Berk., and P. parasitica, Pers., the fruiting-
stems or conidiophores and branches are flattened, and as
these flattened stems and branches twist a little -in the
process of growth, they have a spurious appearance of
swelling in a ganglionic manner. Each ultimate branchlet
of P. ganglioniformis, Berk., is beautifully dilated into a
saucer-like expansion, with a single excessively-attenuated
spicule growing from the centre of the saucer, and with
from three to five similar minute spicules growing round
270 DISEASES OF FIELD & GARDEN CROPS. [OH.
Fia. 125. — LETTUCE MILDEW.
Peronospora ganglionifoi-mis, Berk. Enlarged 400 diameters.
Tip of branchlet with spicules and conidium, 1000 diameters.
the saucer's edge. Each spicule is like one of the
xxxiv.] LETTUCE MILDEW. 271
glandular hairs of a sundew leaf, or like a snail's tentacle
with its eye-bulb. P. parasitica, Pers., has no saucer-like
expansions. One of the exquisitely fine tentacle-fringed
saucers with a spore, balanced on one tentacle, is enlarged
to 1000 diameters at A. At the top of the spore at B,
a small, almost invisible, papilla may be seen. This is the
weakest point of the spore, and at the time of germina-
tion this part dissolves away or bursts, and the contained
material pours out as a germinal thread. The spawn
threads of this species grow within and upon the leaf,
and attach themselves to the constituent cells in a manner
similar with the cabbage Peronospoar, illustrated at Fig. 29.
In whatever part of the leaf or stem the spawn grows it
there sets up putrescence. The mycelium within the leaf
is thick and coarse, and much too stout for emergence
through the organs of transpiration. To meet this diffi-
culty the spawn becomes naturally flattened as it ap-
proaches the stomata, and it emerges through the little
slit on the under surface of the leaf, with a chisel edge,
as does P. parasitica, Pers., see Fig. 30. As soon as the
spawn thread of P. ganglioniformis, Berk., reaches the air
it commonly branches, and, instead of a single fruiting-
branch depending from each stoma there may usually be
seen a small bunch of threads or conidiophores.
This parasite causes pallid patches of decomposition to
appear on lettuce leaves, and on the margin of these
patches little white knots of the destructive mould may
be readily seen with the unaided eye. In the spring the
pest generally shows itself first on the outside leaves near
the ground where the air is still and humid, and grows
inwards to the heart of the lettuce, carrying decay in its
course. In bad cases summer lettuces are quickly reduced
to putrescent masses. In the autumn the flowers and
seeds are chiefly damaged ; sometimes the harvest of seeds
is totally destroyed. Thickly sown plants are worst
affected. The fungus lives on the lettuces all the season,
from spring to October.
272 DISEASES OF FIELD & GARDEN CROPS. [CH.
It frequently happens that young lettuces grown in
frames in the spring are badly attacked, and in many
instances wholly destroyed ; the equable warmth and
humidity of a plant -frame is highly favourable to the
growth of this fungus. When frame lettuces are attacked,
a good plan for the destruction of the fungus is to give as
much air as practicable, and if possible to leave the frames
open for at least a part of one cold night. A short exposure
to cold or slightly frosty air will not materially hurt the
young lettuces, but will so cripple the vitality of the
fungus that the lettuces, on being planted out, will often
be found quite free from the parasite. With both lettuces
and onions, however, it has been observed that trans-
planted examples are often the weakest, and these weakest
plants are worst affected by their respective mildews.
The existence of Peronospora ganglioniformis, Berk., is
preserved through the winter by the means of oospores or
resting-spores. The resting-spores may sometimes be easily
found in the old rotting stems of lettuce plants which have
been destroyed by the fungus. They are generally most
abundant between the spiral vessels and the external
shells of old lettuce stems. A group of the resting-
spores, as found by the Rev. J. E. Vize, near Welshpool,
is illustrated at Fig. 126, enlarged 400 diameters. The
oospores grow in enormous conglomerated lustrous
masses : so profuse is this, their natural habit of growth,
that under the microscope the masses of oospores look
like the shining roe of some fish. As in the fungus of
the potato disease, the resting-spores not uncommonly
grow within the spiral vessels as illustrated. These
oospores hibernate in rotting lettuce refuse during the
winter, and germinate in the early spring. On germina-
tion the first produced conidia perish, unless they alight
on lettuces or other suitable plants. When garden lettuces
are not near, the fungus is nursed by several common
cruciferous weeds; of these the worst is said to be the
common groundsel, Senecio vulgaris, L. The statement is
XXXIV.]
LETTUCE MILDEW.
273
a common one in text books that P. ganglioniform'te, Berk.,
does not produce oospores in lettuces, or P. infestans,
Mont., oospores in the potato. Our experience has been
the reverse of this.
Peronospora ganglioniformis, Berk., grows on various
other living composite plants in addition to the garden
lettuce, Lactuca saliva, L., and L. altissima, M.B. Of
these the different sorts of sow-thistles, as the corn sow-
thistle, Sonchus arvensis, L.? are most often attacked ; then
•X'4-OO
FIG. 126.
Oospores or resting-spores of the fungus of Lettuce Mildew, Peronospora
ganglioniformis, Berk. Enlarged 400 diameters.
Garduus arvensis, Curt. ; Cichorium Endivia, L.; the
nipplewort, Lapsana communis, L. ; groundsel, Senecio
vulgaris, L.; and other plants, both wild and cultivated.
Weeds act as nurses of the fungus for the garden
lettuce.
For the prevention of the disease it is desirable that no
decaying lettuces or weeds belonging to the series just
mentioned be allowed to rot in the fields or kitchen
T
'274 DISEASES OF FIELD & GARDEN CROPS. [OH.XXXIV.
gardens where lettuces are grown ; such material, where
possible, should be carefully gathered together and burnt.
Old lettuce stumps left in the ground are especially
dangerous, as in these decaying stumps the resting-spores
of the mildew often exist in myriads.
CHAPTER XXXV.
POTATO DISEASE, I.
Peronospora, infestans, Mont.
ITS ACTIVE STATE.
THE question is often asked, When did the potato disease
first appear ? No one is able to answer this question.
The fungus which causes the disease is, like the potato
itself, of exotic origin. Peronospora infestans, Mont.,
grows on the wild potato plants of Peru. The strong
probabilities are, that ever since the potato plant has
existed, there has also been the putrescent fungus to prey
upon it. The family of parasites to which the potato
fungus belongs existed in geological times, long prior to
the potato plant or any of its relatives.
It is important to remember, in the consideration of
this subject, that the potato and its immediate allies are
not the only plants destroyed by the potato fungus, for
various members of the family to which the potato belongs
also fall before the parasite. Of late years, in some dis-
tricts the out-of-door cultivation of the tomato, Lycopersi-
cum esculentum, Mill., has been quite stopped by the
ravages made upon it by the potato fungus. All the
species of Lycopersicum, of which we have at least eight
forms in our gardens, are commonly attacked by the fungus
of the potato disease. Sometimes the pest may be seen
growing upon the henbane, Hyoscyamus niger, L., or com-
mon bitter-sweet, Solanum Dulacamera, L., of our hedges ;
at other times it may be observed upon various other species
of Solanum, as S. demissum, Lind., and S. cardiophyllum,
276 DISEASES OF FIELD & GARDEN CROPS. [CH.
Lind., or on the Petunias of our gardens. Sometimes the
parasite may be seen upon an entirely different natural
order of plants from that of the potato ; it may leave the
Solanacece and prey upon members of the Scrophulariacece,
as the New Holland plant named Anthocercis viscosa, R.Br.,
or, as pointed out by Professor de Bary, the Chilian
Schizanthus Grahami, Gill. We have both these plants in
our gardens. It is worthy of note that there is a second
species of Peronospora met with on Solanaceous plants,
named Peronospora Hyoscyami, P., and peculiar, or nearly
so, to the common henbane, Hyoscyamus niger, L.
Hyoscyamus is by no means common in Britain, but its
parasite has been recorded from a single locality near
Market Deeping.
Circumstances might have been, and probably were,
adverse in this country to a rapid spread of the potato
fungus soon after the first introduction of the potato. At
length the time arrived when circumstances changed, and
something — it is impossible to say what — greatly accelerated
the growth and vigour of the fungus. When the potato
and its parasite were transferred from South America to
Northern Europe, the climatic conditions were changed,
and the potatoes were grown in a new, artificial, and
unnatural manner. The constitution and habit of the
potato also became changed, — we do not say weakened,
although this may be the case, but altered. At the time
of the alteration or modification of the nature of the potato
plant, the potato fungus acquired greater potency over it.
The same phenomenon has occurred with the tomato : as the
plant has been gradually altered in habit by cultivation,
so the habit of the assailing fungus has varied. There is
no evidence to show that the Peronospora has altered in
the least in the potency or non-potency of its attacks
upon our neglected wild plants.
The first accounts of the potato disease in Europe are
very obscure, but as our business is less to detail the
history of the fungus than to describe it with its effects,
xxxv.] POTATO DISEASE, I.— ITS ACTIVE STATE. 277
we shall dismiss this part of the subject briefly. Mr.
Berkeley, writing in vol. i. p. 9 of the Journal of the
Royal Horticultural Society, stated that at that time, 1846, a
"very serious disease" had existed for more than half a
century under the name of " Curl," which committed
" immense ravages " in the north. At the present time
we know that the " Curl " was, and still is caused by the
fungus of the potato disease, Peronospora infestans, Mont.
In a communication to the French Academy, 17th Nov-
ember, 1845, M. Boussingault wrote, on the information
of M. Joachim Acosta, that the malady was well known in
rainy years at Bogota, where the Indians live almost
entirely on potatoes. There was a disease of potatoes in
1815, and a second noticed under the name of " Dry
Rot " in Germany in 1 830 ; if this " Dry Rot " of potatoes
was equally moist with the " Dry Rot " of timber, it would
exactly agree with what we know of the potato disease
now. It must be remembered here that Fusisporium
Solani, Mart., often really dries up and destroys potato
tubers. The year 1830 was not a year of daily news-
papers, of sharp scientific observers, and students of the
microscope. It may therefore be reasonably concluded
that if potatoes were sufficiently diseased with " Rot " in
1830 to warrant a published account of the disease, they
most probably were diseased to a less extent for several
previous years, and probably before the year 1815 just
mentioned. Many articles appeared in the newspapers
and agricultural periodicals of 1833 regarding the "Rot"
of potatoes in the northern counties of England (see
Journal of the Royal Horticultural Society, vol. iii. p. 22).
In 1840 the disease was widely spread in Germany and
France, and in 1841 it again attracted great attention in
Belgium. A sharp observer, Dr. Morren, at that time
advised that the putrid stems should be immediately re-
moved from diseased potato plants — a piece of advice which,
under proper conditions of the growth of the potato tuber,
might be followed with good results at the present day.
278 DISEASES OF FIELD & GARDEN CROPS. [OH.
In the same year it was recorded in Vik, in Norway, by
Mr. Westrem, the director of the Agricultural School.
During the next year it had greatly extended itself and
was recorded from Sogndal, as well as from Denmark in
both years. In 1843 the disease was very destructive in
Western Jutland, and in 1844 the potato disease was
epidemical in St. Helena and Canada. From the pub-
lished accounts of the periods mentioned, it may be seen
that the potato murrain was then exactly as we see it
now. It appeared at a similar period of the year, and
during the typical moist warm weather so favourable to
the growth of the fungus. The dark disease blotches were
on the leaves and the tubers were murrain-stained and
rotten. The offensive odour so familiar to us now was
then specially noticed. The next year, 1845, was the
ever memorable year of the great outburst of the potato
disease over Western Europe, from Norway to Bordeaux,
and the northern parts of the United States. In 1845
the fungus of the potato murrain acquired its greatest
possible power for destruction. It was first noticed in the
south of England in the middle of August, and in a fort-
night it had spread over every part of the British Isles.
So apparently sudden and destructive was this attack,
that in the month of September it was hardly possible to
procure potatoes unstricken by the murrain. From 1845
till now we have never been free from the assailing
fungus ; sometimes the attacks are extremely virulent — at
other times slight ; sometimes the fungus is common on
various field and garden plants allied to the potato — at
other times very little of the fungus is to be seen. Mr.
Duncan Stuart has stated in the North British Agriculturist
for 3d October 1883, that the fungus of the potato disease
has never yet appeared in the Island of Eum, fifteen
miles from the mainland, on the west coast of Scotland,
and seven miles to the south of Skye. An exhaustive
account, and the best ever written of the rise and spread
of the potato disease in Europe, is given in the first volume
xxxv.] POTATO DISEASE, I. —ITS ACTIVE STATE. 279
of the Journal of the Royal Horticultural Society, from the
pen of the Rev. M. J. Berkeley. This paper also gives a
complete description of the potato fungus, — a description
so complete and admirable that, even now, very few new
facts can be added to it. Since 1846, when that account
was published, many fresh observers have written on the
potato fungus, and some of Mr. Berkeley's original obser-
vations have been amplified, enlarged, and curiously con-
firmed.
The potato disease is seldom seen in Europe before
July or August, although its appearance has been noted
on rare occasions in May and June. Mr. Jensen has
stated that the fungus of the potato disease cannot exist
in any country where the mean temperature exceeds 77°
Fahr. for any length of time during the period when the
fungus generally perfects itself, and that in a temperature
of 34° it cannot produce either mycelium or spores. It is
generally first distinctly seen in the midland and southern
counties of England between the 20th and 31st of July.
It generally appears during close humid weather, when
there are mists in the fields in morning and evening,
and the days are hot, damp, and possibly stormy. Many
other fungi suddenly perfect themselves under exactly
similar meteoric conditions. It will be pointed out later on
why it is, as we think, that the potato fungus appears with
apparent suddenness under these conditions and at this
particular time of the year. In the meantime it may be
noted that the fungus generally makes itself manifest to
the less experienced observer as a fine white bloom on the
leaves, accompanied by dark putrid spots. The bloom is
sometimes more profuse on the lowermost leaves of potato
plants, not because the fungus has travelled up the stem
from the seed tuber, but because the air is more moist
and stagnant near the ground. The bloom, with its accom-
panying black disease blotches, soon travels to the stems,
and when at length the tubers are reached the exhausted
seed tuber (the weakest part of the plant) is commonly
280 DISEASES OF FIELD & GARDEN CROPS. [CH.
traversed in every part by the spawn of the fungus.
During warm, humid conditions of the weather the black
decomposed spots are sometimes present for several days on
the leaves before the fungus is seen. These blotches indi-
cate that the putrefactive spawn of the fungus is within
the leaves, awaiting favourable conditions for its complete
development as a white bloom outside. The phenomena
just mentioned are accompanied by a peculiar and very
offensive odour well known to every person who has
walked through a field of potatoes suffering from disease.
The odour is caused by the putrescence set up in the
tissues of the host plant by the contact of the mycelium of
the potato fungus. Although the attack of disease in
potato plants is apparently sudden, and made on
apparently sound plants, yet all known facts point to
the probability of the existence of the fungus in a nascent
state during at least several weeks prior to its general
recognition. The belief in the extreme suddenness of
fungoid growths is, in many instances, a mere popular
delusion. The common field mushroom is supposed by
rustics to grow in a single night ; but it is well known
to careful observers that the infant mushroom exists
just beneath the surface of the soil in a growing state for
several weeks before it suddenly bursts through the earth
and expands its cap or pileus. We have ourselves seen
fields of potatoes which were apparently undiseased one
day, prostrate on the ground the next, and the haulms
blown away by the wind on the third day. This apparent
suddenness of the attack in the early autumn appears to be
well known in America ; for Professor W. G. Farlow of
Harvard University writes in reference to the " Potato
Rot." (Bulletin of the Bussey Institution, part iv. p. 319):
" At times its advent is so sudden that, within a few
hours, the potato fields change from green to brown and
black, and the plants which, in the morning, gave pro-
mise of an abundant crop, before night present a mass of
decaying vegetation, in which are involved not only the
xxxv.] POTATO DISEASE, I.— ITS ACTIVE STATE. 281
leaves and stems, but also tlie tubers." The following
interesting and instructive sentence occurs in an excellent
essay written by Dr. W. Peard, LL.B., on "Certain Enemies
of our Roots," and published in the Journal of the Bath and
West of England Society and Southern Counties Association,
vol. iv., third series, p. 14 : — "At that time (Aug. 1845)
we were spending some weeks at Bally shannon, and close
to our cottage was a magnificent field of potatoes, about
twenty acres in extent, through which we passed regularly
every morning and evening. One day, during the last
week in August, as we brushed through the dark-green
foliage, earthy disagreeable odours, before unknown to us,
rose from the plants. On the following morning the
entire crop looked as if it had been exposed during the
night to the action of steam. Stems and leaves were
soft, pulpy, and blackened ; in six-and-thirty hours a few
sickly stems and discoloured leaves were all that remained.
The crop had ceased to exist."
Mr. J. G. Baker, F.R.S., of Kew, is of opinion that the
potato plant, Solanum tuber osum, L., in its present " tuber-
bearing state is in a disorganised, unhealthy condition, a
fitting subject for the attacks of fungi and aphides ;" and
he quotes Mr. T. A. Knight to the effect that the formation
of the tubers more or less deprives the potato plant of its
requisite amount of nutriment. He considers that the
potato is grown in a necessarily unnatural way in masses
in our fields, instead of in isolated examples as in Nature ;
and that the fact of the almost total absence of flowers and
fruit in many cultivated varieties shows that the plant
is in a disorganised state. Mr. Baker, from an examina-
tion of a large number of examples, has come to the con-
clusion that all the garden varieties have originated from
S. tuberosum, L. Out of 700 or 900 species of Solanum
it appears that only six produce tubers or potatoes at all ;
the rest " maintain their hold on the world as most plants
do, by their flowers, fruits, and seeds."
Other observers hold an opinion at variance with tlie
282 DISEASES OF FIELD & GARDEN CROPS. [OH.
one advanced by Mr. Baker, and say it is impossible to
over-cultivate any plant ; that seedling potatoes are as
badly affected as those grown from cut tubers ; and that
animals and plants placed in an artificial position by man
only need an extra amount of care corresponding with
their new position. Race horses are said to live as long
as cart horses, domestic as long as wild animals, and the
delicate children of towns as long as the more roughly
nurtured children of country villages. It is acknowledged
that extra care is required, but it is maintained that the
constitution is not impaired.
In the potato disease, as in every other disease, both of
animals and plants, it is necessary that the ailment should
be completely understood before any serious attempt can
be made towards the prevention or palliation of the
attacks. We will, therefore, closely examine the struc-
ture and mode of growth of the potato fungus.
For an exact examination of Peronospora infestans,
Mont., a very minute and extremely thin and transparent
slice must be cut from a diseased leaf at a spot where
the white bloom caused by the presence of the fungus is
visible underneath. A good plan is to cut a diseased
leaf in two through a disease spot, and then with a sharp
lancet cut an extremely thin slice off from one of the ex-
posed cut surfaces. If the slice last cut is somewhat
longitudinally wedge-shaped, it will often best show the
structure of the leaf and the contained fungus at the
thinner end of the section. Such slicing requires great
care and experience, and the art is only acquired after
many failures. To those, therefore, who are unequal to
the task we advise the purchase of slides ready prepared
by the Rev. J. E. Vize. The atom to be examined
should be placed on a glass slide in a drop of glycerine
(this is preferable to water, as the latter often dries too
quickly), and then covered with a clean thin cover-glass.
The magnification given by an ordinary lens is useless for
the observation of the minute fungus now before us, so we
xxxv.] POTATO DISEASE, I.— ITS ACTIVE STATE. 283
must at once place it under the higher powers of the micro-
scope. If the slicing through a disease spot is successful, we
shall probably see the atom when magnified 100 diameters,
as at Fig. 127. The thickness of the lamina of the leaf
is shown at A, B ; the under side of the leaf is represented
at A, from which surface the fungus almost invariably
springs. The fungus, therefore, really grows downwards.
The true upper surface is shown at B. This reversal
of the leaf in the illustration is merely, as in other in-
stances in this book, to show more clearly the treelike
branching growth of the fungus. If we confine our
attention for the present to the section of the leaf, we
shall note that it is made up of minute bladder-like cells,
loosely packed together ; and that the cells at top and
bottom, representing the lower and upper cuticle of the
leaf, are devoid of the shading, which is meant to indicate
the green colouring matter or chlorophyll within. An
opening into the interior of the leaf will be seen at C ;
this is one of the stomata or organs of transpiration,
sometimes referred to as " breathing pores." The stomata
are like the gates to a camp or to an entrenched position ;
they are the weak points through which an enemy may
enter, and when once these gates are passed, the whole
interior of the plant is at the mercy of the invader. At
D may be seen a hair built up of four transparent cells,
the two lower being traversed by a mycelial thread of the
potato fungus. On the upper part of this hair, attached
to the outside at E, may be seen one of the small branches
of the fungus ; this branch has burst and thrown out a
mycelial thread from its side. Every fragment of the
potato fungus is capable of growth, and of ultimately
reproducing the parent fungus. The cells immediately
under the true upper cuticle of the leaf at F are termed
pallisade cells ; and their disposition in the manner
illustrated serves to give the necessary firmness to the
exposed upper surface of the leaf.
If we now look within the fragment of the leaf we
284 DISEASES OF FIELD & GARDEN CROPS. [en.
X-loo
FIG. 127.
Section through a fragment of a potato leaf, with the potato fungus, Perono-
spora infestans, Mont., growing within its substance, and emerging
through the epidermis. Enlarged 100 diameters.
xxxv.] POTATO DISEASE, I.-ITS ACTIVE STATE. 285
see transparent threads running between the small
spherical leaf -cells ; these are the spawn -threads or
mycelium of the fungus. It should be especially noticed
that wherever the spawn touches the cells it discolours
them (as indicated by the darker shading), and causes
putrescence by contact. If we again look at the pallisade
cells near G, we observe that a spawn-thread has pushed
itself between them and between the cells of the upper
cuticle, and is emerging into the air. If We trace the
spawn-threads to the organ of transpiration at H, we
notice that a thread in its passage from the body of the
leaf has blocked up a so-called mouth. This choking
prevents the transpiration of vapour, and hastens pu-
trescence. Two other threads have pushed themselves
between the leaf -cells at G and A. When the larger of
the emerged threads is traced upwards to K, a treelike
growth is noticed; and this branching form is the fruit-
ing condition of the fungus of the potato disease called
Peronospora infestans, Mont. The name Peronospora has
been explained ; infestans needs no explanation. The
whole fungus is perfectly transparent, like colourless glass,
and extremely fine, thin, and attenuated in all its parts.
Some book illustrations give a very erroneous idea of the
fungus, owing to the use of thick lines and an unnatural
amount of dark shading. We notice as a rule that the
fruiting-stems or conidiophores, as at K, have comparatively
few joints or septa ; sometimes, however, old examples, as
at L, are full of joints. If we now look at the branches
MM, we observe that each is surmounted by a transparent
spore, technically termed (as in other species of Perono-
spora] a conidium as at NN; and to these bodies we
shall more specially refer further on. It must also be
noticed that all the branches are more or less constricted
or jointed in a peculiar manner, as at 00 ; and that each
joint has at one time carried a conidium, the lower conidia
having been pushed off as the branches have continued
their growth, as at PP. Sometimes :„ weakly ini-
286 DISEASES OF FIELD & GARDEN CROPS. [on.
poverished thread, if grown in dry air, will quickly
become strong and robust in growth if transferred to
warm moist air, as in the thread illustrated at Q. In
many species of Peronospora the branches which carry the
conidia only produce one conidium, and do not continue
growing and producing new conidia. Owing to the mode
of spore production in the potato fungus, Professor De Bary
has recently suggested that the parasite should be placed
in a new genus by itself under the name of Phytophthora.
Other botanists, however, as the Kev. M. J. Berkeley
and ourselves, would prefer reducing rather than increas-
ing the genera of the Peronosporece ; and so include in
Peronospora not only Phytophthora but Ovularia and even
Ramularia.
At Fig. 128 we have engraved a fragment of the
potato fungus to 400 diameters, so that the parts may
be compared with the other species of Peronospora en-
graved to the same scale in this work. A considerable
difference will be observed if Fig. 2 and Fig. 16 are
referred to. The spores or conidia are shown at A,
Fig. 128, and the peculiar constrictions in the jointed
branches are better seen. If ripe conidia are placed
in water it will be noted that a differentiation of the
contained protoplasm takes place, as shown at BB ; and
that the interior mass of each conidium becomes divided
into from five to nine or more portions, each contained
portion being furnished with one or two lustrous vacuoles.
These differentiated portions speedily emerge from the
top of the conidium when placed on any moist surface
as at C ; and each portion now free, becomes quickly fur-
nished with two extremely fine hairlike cilia, tails, or
vibrating hairs, as at D. These secondary spores or
zoospores are able to sail about in the slightest film of
moisture. After a brief time the little motile zoospores
or animal -like spores rest and take a globular form, as
at E, and the vibrating hairs dissolve away or drop into
the finest dust. After a short rest the now quiescent
xxxv.] POTATO DISEASE, I. -ITS ACTIVE STATE. 287
FIG. 128.
Fragment of the upper part of a conidiophore of the potato fungus, Perono-
spora infcstans, Mont., with conidia and zoospores germinating.
Enlarged 400 diameters.
288
DISEASES OF FIELD & GARDEN CROPS. [on.
zoospores burst and produce a thread of spawn, as shown
at F ; this germinal thread is capable of carrying on the
existence of the potato fungus. Zoospores were first dis-
covered in the genus Cystopus, allied by some authors, with
Peronospora (see chapter xvi.), by Prevost in 1807; and
Mr. Berkeley described and illustrated them in the potato
fungus, though he did not see the vibrating hairs, in
1846, Journal of the Royal Horticultural Society, vol. i. pi.
4, fig. 18. Sometimes the conidium, which, when it
bears zoospores, is really a sort of spore-case, sporangium,
or zoosporangium, does not differentiate within, but bursts
X-1000
FIG. 129.
Conidium or zoosporangium and zoospore of the potato fungus, Peronospora
infestans, Mont. Enlarged 1000 diameters.
and protrudes a small mass of protoplasm or vital material,
as at G, Fig. 128. This mass speedily elongates into a
mycelial thread capable (like the thread from the zoospore)
of carrying on the life of the potato fungus. A ripe
conidium or zoosporangium and zoospore of the potato
fungus are farther enlarged to 1000 diameters at Fig. 129,
for comparison with other reproductive bodies illustrated
to the same scale in this work. A considerable difference
in the size of zoospores will be noted if Figs. 33 and 39
are referred to. The soft papilla or bursting point of
the zoosporangium is shown at A, Fig. 129; and
the minute footstalk by which the zoosporangium was
xxxv.] POTATO DISEASE, L— ITS ACTIVE STATE. 289
originally attached to the stem of the parent fungus at
B. It must be specially noted that water or moist air
is essential for the existence of the fungus, for nearly
every part speedily perishes in dry air, heat, or frost.
When the conidia burst and set free the minute zoospores,
the latter sail over the damp surfaces of leaves, and even
float into the organs of transpiration. A zoospore swimming
in an intercellular space is shown at R, Fig. 127. One
has only to imagine a large field of potatoes, with all the
leaves moist and swaying backwards and forwards with
the wind, to perceive that such a field, say on a warm
misty morning or evening, would form a sort of continuous
lake of moisture on which the zoospores could float from
one plant to another. The conidia, with the contained
zoospores, are also carried through the air in millions by
the wind ; they are so lightly attached to their sup-
porting stems and so extremely small and light, that the
faintest breath of air wafts them away. Insects and
other creatures also carry the conidia from place to place.
The flies which alight on potato plants carry off hundreds
of conidia on their bodies. If a bird drops in a field of
diseased potatoes, the fluttering of its wings will disperse
millions of the conidia of the fungus of the potato murrain
into the air. The same phenomenon occurs when a dog
or other animal runs amongst diseased potato plants.
When the conidia or zoospores burst and germinate, the
threads which emerge are corrosive or putrefactive. To
such an extent is this the case that the spawn is said to
be capable of piercing or boring through the cuticle of
the leaf from within or without, regardless of the natural
openings or stomata, and even of piercing the bark of the
stem or the tuber itself.
The fungus of the potato disease generally attacks the
leaves first, and, as the leaves produce successive crops of
fungus growth, the disease quickly spreads to the leaf-
stalks, from the leaf-stalks to the chief stems, and from
the stems to the tuber. Sometimes a week or two elapses
290 DISEASES OF FIELD & GARDEN CROPS. [CH.
before the tubers are reached by the putrefactive spawn
of the fungus, but in other instances the attack is so
sudden and so highly destructive that the whole of the
potato plants above ground in a large field will be
destroyed in a day or two. The disease doubtlessly
starts at first from a few centres only ; there it remains
for a brief time more or less unobserved. The fungus,
however, possesses such wonderful powers of spore pro-
duction and rapid growth, especially when the air is
moist and the temperature ranges from 60° to 70° Fahr.,
that in a few days one fungus growth will become ten
thousand. This growth goes on in a constantly increas-
ing ratio until at length the great flood of disease seems
to almost suddenly cover the potato fields. When the
attack is not violent it is obvious that a good plan is to
remove all the tainted potato stems and foliage before the
spawn reaches the tubers ; but, on the other hand, if the
stems are removed before the tubers are ripe, injury must
accrue to the crop, as the starch which is subsequently
stored up in the tubers is formed in the leaves. Still, it
is better to have a poor or partial crop than none at all.
Some cultivators advise the growing of more early ripening
potatoes, as such varieties now commonly escape the
murrain ; but there can be little doubt that if a change
could be brought about in the general habit of the potato
plant it would be followed by an exactly corresponding
change in the habit of the parasite. Late ripening varie-
ties are, moreover, wanted, and their cultivation cannot
be dispensed with.
When the fungus spawn reaches the tuber it decom-
poses the cells and corrodes the starch. In bad cases the
tubers are soon reduced to a mass of putrefaction. In
mild cases the spawn of the fungus hibernates and be-
comes perennial, as was first pointed out by the Rev.
M. J. Berkeley in vol. i. of the Journal of the Royal Horti-
cultural Society for 1846. Mr. Berkeley writes (p. 26)
in reference to the fungus growing from ripe, harvested,
xxxv.] POTATO DISEASE, I.— ITS ACTIVE STATE. 291
apparently sound, but really diseased tubers : " It
should seem certain, then, that the mycelium or elements
of the fungus must have pre-existed in the tuber, and,
as it uniformly springs from the decayed spots, that it
has itself caused the decay. But here a difficulty arises
from the great obscurity, or, as some say, the total absence
of mycelium in an early stage of the disease. I have
satisfied myself, however, of its existence in some cases,
but not uniformly." Again, at p. 28, he writes : " On
examining the cuticular cells of a young tuber, with a
view to ascertain the changes which occur in the process
of greening, I found evident traces of mycelium within
them." In the same paper Mr. Berkeley refers to the
perennial mycelium of corn mildew. In the Outlines of
British Fungology, p. 42, Mr. Berkeley writes : " Spawn
. . . may exist for years without producing fruit . . .
whether it runs through soil or decaying substances, or
amongst living tissues, whether without or within their
walls." Professor de Bary was no doubt unaware of
these published observations when he wrote for the
Royal Agricultural Society, vol. xx. p. 265, 1876, that he
was perhaps the first to point out the presence of perennial
mycelium in the potato in 1863. No vegetable growth
is more common and well known than perennial my-
celium ; indeed, with very few exceptions, the spawn
belonging to all fungi must at times be perennial. Al-
though perennial mycelium has a far stronger hold on
life than have simple spores or conidia, it must not be
imagined that perennial mycelium always survives after
an unusual amount of heat, cold, or moisture. In the
same way as mushroom spawn often dies in the " bricks *
of the nurseryman, so the perennial mycelium perishes in
the invaded tuber of the potato. In examining diseased
potatoes in the winter and spring it is common to find
the mycelium dead, and if such diseased potatoes are
planted a perfectly sound crop will be the result. In the
Gardeners' Chronicle for 24th January 1874, we have re-
292 DISEASES OF FIELD & GARDEN CROPS. [CH.
corded a number of curious experiments made with diseased
potatoes. In those experiments it is shown that not only
will diseased tubers sometimes produce perfectly sound
plants, but that slices of diseased tubers, inserted within
the substance of sound ones at the time of planting, will,
in some instances, have no ill effect, but that the roots of
the sound plant may grow in the putrid remains of the
diseased one and still remain untainted. If the perennial
mycelium is alive, the result is generally, but not invari-
ably, different. The common mushroom spawn, as sold
by nurserymen in " bricks," the white fungus threads we
everywhere see when the earth is upturned, and the
white felt so commonly noticed amongst fallen leaves, is
perennial spawn. The spores of fungi are so extremely
delicate that a slight variation of heat, dryness, or
moisture often destroys them at once ; but when spores
have once germinated and produced spawn, this spawn
does not so readily perish, but may rest for a long time
in a hibernating state. This is proved in the case of
the familiar fairy-rings of our lawns and pastures,
which sometimes are not seen for many years, as the
subterranean spawn is awaiting suitable conditions of
warmth and moisture to cause it to produce the perfect
fungi.
Here we must not forget that the virus of the disease
may exist in some form which has not yet been detected
by our microscopes. Before the higher powers of the
microscope were used no one suspected the presence of
motile zoospores with vibrating cilia ; and if we could use
still higher powers, it is not unreasonable to imagine that
some other condition of the parasite, at present quite
unknown to and unsuspected by us, might be brought to
light. The fungus may exist in inconceivably fine dust-
like particles, or in the condition of a mucous fluid.
Because we are acquainted with a certain number of
curious facts regarding fungi, it does not follow that we
know all. We have long suspected that the virus of this
xxxv.] POTATO DISEASE, I.— ITS ACTIVE STATE. 293
and many other fungi may exist in inconceivably small
and perhaps Amoeboid particles.
The resting of the mycelium in a state of hibernation
through the winter may, perhaps, sometimes account for
the reappearance of the disease the next season ; for it has
been known, since Mr. Berkeley pointed it out in 1846,
that a broken or cut surface of a diseased potato will, if
the mycelium is alive, give rise to the potato fungus at
any time of the year on the cut potato being exposed to
an atmosphere suitably warm and moist. It is obvious
that, if the potato disease is annually reproduced by dis-
eased tubers containing perennial mycelium, the disease
must invariably begin in the seed-tuber and ascend the
stem ; but it is known by experience that in the vast
majority of instances this is not the case, but that the
disease first invades the leaves.
Flowering plants have three familiar modes of increase.
One is by suckers, runners, or underground stems ; these
runners are roughly comparable with perennial mycelium.
A second is by buds or bulbils, at times very common in
the axils of the leaves of some lilies ; these may be com-
pared with the conidia or bud spores of fungi; and a third
is by the reproductive organs, or stamens and pistil.
^Reproductive organs of a like nature, as far as sex is con-
cerned, are known in fungi, and they are potential to and
extremely common, well marked, and easily seen in the
genus Peronospora, to which the potato fungus belongs.
The organs belonging to Peronospora infestans, Mont.,
as far as we have at present described them, have been
distinctly asexual, or without sex; no male and female
organs answering to the stamens and pistils of flowering
plants have yet been referred to.
Mr. Berkeley as long ago as 1846 described and illus-
trated, from materials furnished to him by Dr. Montagne,
what he believed to be an oogonium, — an organism which
may be compared with an ovule or unimpregnated egg, and
its oospore, or resting-spore condition, which is more or less
294 DISEASES OF FIELD & GARDEN CROPS. [CH. xxxv.
comparable with the impregnated ovum, or fertilised seed ;
but it unfortunately happened that from 1846 to 1875 no
one saw the bodies again as originally described by Messrs.
Montagne and Berkeley. This failure may have arisen from
bad searching or from searching at the wrong time, or,
as we believe, in wrong material, through imperfect know-
ledge ; it may partly have arisen from the fact of the
oogonia being exactly the same in size with the cells of the
potato leaf. Whatever the reason may have been, we at
length saw these bodies again in 1875 within the leaves
of badly diseased potatoes sent to us from the garden of
the Royal Horticultural Society at Chiswick. The sexual
organs are illustrated at S, T, U, and V, Fig. 127. As
we were, at the time of the discovery, familiar with Mr.
Berkeley's writings and views, we instantly perceived, on
looking at these growths for the first time, that we had
before us the bodies first detected by Dr. Rayer, Chief
Physician of the Hopital de la Charite" at Paris ; described
by Dr. Monatgne, and referred by Mr. Berkeley to Perono-
spora infestans, Mont.
CHAPTEE XXXVI.
POTATO DISEASE, II.
Peronospora infestans, Mont.
ITS PASSIVE STATE.
WE will now closely examine the bodies found in spent
potatoes by Dr. Rayer, illustrated by Dr. Montagne in
1845, and described by the Eev. M. J. Berkeley in vol.
i. of the Journal of the Royal Horticultural Society, 1846.
Dr. Montagne termed the bodies discovered by Dr.
Rayer in the intercellular passages of potatoes, Artotrogus
hydnosporus. The generic name is derived from the
Greek artos, bread ; and trogo, eating or consuming, and
bears reference to the power of the fungus in consuming
the nutritious material of the tuber ; the name shows
that Dr. Montagne suspected the true nature of the fungus.
Hydnosporus indicates that the spores resemble the fungus
named Hydnum, which has its fruiting surface covered
with spines or prickles ; Hydnum is from the Greek hud-
non, a word used by Theophrastus to denote a truffle.
The specific name is somewhat misleading, as it was only
meant to refer to the mature spores, as is proved by the
writing on Dr. Montagne's original drawing, see page
84 ; and the description of Mr. Berkeley's plate, Journal
of the Royal Horticultural Society, 1846, p. 34. In
infancy Artotrogus is smooth spored. No phenomenon
is better known in fungi, as in Cystopus and the Gas-
teromycetes, than a smooth oogonium or spore becoming
warted or spinulose with age ; and Mr. Berkeley, in the
volume above quoted, from Dr. Montagne's examples,
296 DISEASES OF FIELD & GARDEN CROPS. [CH.
figures both the smooth and echiimlate form on the same
plate. Plain and echinulate spores are produced on the
same plant in some Saprolegniece ; and Dr. Max Cornu
maintains that the Saprolegnia asterophora of De Bary is
merely the warted form of Achlya racemosum, Hildb.
By the courtesy of the Eev. M. J. Berkeley, we have had an
opportunity of carefully examining the original examples
found by Dr. Eayer, and described by Dr. Montagne ; and
we have no hesitation in stating that they are in every way
the same with the bodies found by us on the mycelium of
Peronospora infestans, Mont., in 1875. It is useless to re-
produce Dr. Montagne's illustration from vol. i. of the
Journal of the Royal Horticultural Society, or to engrave
his preparations, as they agree precisely with the illus-
trations made from fresh specimens, and engraved in the
following pages. Dr. Montagne's examples represent
fertilised semi-mature oospores, most of the specimens
have a smooth external surface, but some of the more
mature specimens are spimilose ; and for this reason Dr.
Montagne doubtlessly selected the specific name hydno-
sporus.
Dr. Eayer, then, was the first person who in 1844 or
1845 detected resting -spores in the genus Peronospora;
and the Eev. M. J. Berkeley was the first person who
pointed out the fact of Peronospora being an oospore-bearing
fungus. Mr. 0. Edmund Broome next found the Artotrogus
of Peronospora parasitica, Pers. ; and Mr. Berkeley again
pointed out its true nature in the Gardeners' Chronicle for
1854, p. 724. In 1845 Tulasne made great advances
in our knowledge of the oospores of Peronospora ; and his
observations were laid before the French Academy in
1854, and published in Comptes Rendus for 26th June of
that year. In 1855 Dr. Caspary published still further
advances in the Monthly Transactions of the Royal Academy
of Berlin ; he there illustrates the oogonia or sporangia, as
he terms them, of Peronospora Hepaticce, Gasp., and P. densa,
Gasp. Dr. Caspary's observations were, he says, made
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 297
before he was aware of Tulasne's discoveries. Professor
De Bary of Strasbourg, in 1863 (Ann. des Sc. Nat., ser.
iv., vol. xx.), made still further progress in detecting the
oospores or resting-spores of Peronospora.
Oogonia or immature resting-spores are illustrated,
growing from the mycelium of the potato fungus within a
potato leaf, at S, T, U, and V, in Fig. 127. At S an
oogonium is seen intercalated within a thread of the
potato fungus ; at T and U the oogonia are terminal,
each with a second smaller body, termed an antheridium,
attached ; and at V another intercalated example is shown.
FIG. 130.
The sexual organs, or oogonia and antheridia of Peronospora infestans,
Mont. Enlarged 400 diameters.
These bodies belong to the Artotrogus hydnosporus of Mon-
tagne. As the magnification of 100 diameters is insuffi-
cient to distinctly show the nature of these oogonia, we
have enlarged them to 400 diameters in Fig. 130. In each
of the four examples A represents an oogonium or cell in
which a female reproductive body, termed an oosphere,
is formed ; and each of the two bodies at BB is termed
an antheridium, or cell which contains the male repro-
ductive material. The oogonium may be roughly com
pared with a pistil in flowering plants, and the antheridium
298 DISEASES OF FIELD & GARDEN CROPS. [en.
compared with the anther and its pollen. In the process
of growth the antheridia naturally come into contact
with the oogonia, just as an anther may touch a stigma.
When this contact takes place the antheridium projects a
fine tube into the walls of the oogonium till the oosphere
within is touched and pierced. Some of the vital mate-
rial from the antheridium then very slowly passes through
the tube, mingles with the protoplasm within the oosphere
and fertilisation, and the
formation of an oospore is
the result. Just as in
flowering plants an ovule
becomes a seed after fertil-
isation, so the oosphere
becomes an oospore or rest-
ing-spore after the contact
of the antheridium with
its contents.
An antheridium in con-
tact with an oogonium is
farther enlarged to 1000
diameters in Fig. 131, to
more clearly show the beak
or fecundating tube in the
act of piercing the oogonium
and its contained oosphere.
By careful watching under
the microscope, the granu-
lar protoplasm from the
antheridium may be seen
to pass very gradually into
the oosphere. The oogonium originates by the con-
tained protoplasm in the mycelium congregating in certain
positions, generally at the ends of, or within the mycelial
branches, the end of the branch becomes distended with
vital material, and a bladder is formed, which is speedily cut
off by a septum or joint, as at A, Fig. 131. In many in-
FIG. 131. — Peronospora infestatis,
Mont. Oogonium and antheridium.
Enlarged 1000 diameters.
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 299
stances (but not in all) the antheridium springs from the
same thread as the oogonium, and close to the oogonium
itself ; here the protoplasm again forms a smaller bladder-
like expansion, which soon becomes (as in the oogonium)
separated from the supporting thread by a septum, as at
B, Fig. 131.
After fertilisation the oogonia readily fall from their
supporting threads, just as simple spores drop from their
supports, or as seeds drop out of seed-vessels. There is
no further need of connection with the parent stem, so.
the resting-spores, like seeds, are set free.
It follows from this fact, that unless the oogonia are
sought for at the precisely right time, they will not be
seen upon the supporting threads, which soon perish. A
notable instance of this phenomenon occurs with the
Peronospora of the lettuce. In this species myriads of
oogonia are formed in dense conglomerated masses, Fig.
126 ; but they are no sooner formed than they are cut off
from their supporting mycelium by septa, and then the
mycelium perishes. Nothing is more common than to
see large numbers of oogonia and no mycelium. The
case is similar with free seeds or seed-vessels where the
old tender flower remains have perished.
Every competent botanist who has sought for these
potato oogonia since 1875 has found them. Mr. C. B.
Plowright found both the smooth and spiny form in the
summer of 1876, as described and illustrated in the
Gardeners' Chronicle for 29th July 1876. At the present
time both oogonia and ripe oospores, not only of the
potato fungus, but of various other species of Peronospora,
may be purchased from the Kev. J. E. Vize.
In 1875, when our observations were made, great
attention was directed towards the potato disease, as it
not only occurred with great virulence, but it appeared
unusually early in the season. In that year the leaves
of the potatoes were the first organs attacked, and old
observers said that no such curled and distorted leaves had
300 DISEASES OF FIELD & GARDEN CROPS. [OH.
been seen for many years. We were, therefore, naturally
desirous of making a close examination of them. Some
of the earliest examples were sent on to us, and as we
were well acquainted with the potato fungus and its habit
of growth, we placed a series of infected leaves — one over
the other, like the leaves of a book — in a saucer. To keep
the leaves constantly and naturally moist, we placed a
very little water in the saucer, and this water just touched
the points of the leaves. The saucer was next slightly
tilted, so that the water might remain at one spot, and
the leaves slowly and naturally absorb it. The saucer
with the leaves was then placed under a bell-glass, the
bell-glass was covered with a cloth, and the whole kept in
a warm room. The potato leaves were then, as we thought,
in a favourable position for the full development of the
Peronospora, with no chance of a sudden check from too
much dryness or cold. At that time we had no idea
whatever in the direction of the artificial production of
oospores or resting -spores ; but on examining the potato
leaves as they gradually fell into decay, we were surprised
to see numerous examples of what we believed to be
Artotrogus attached to the Peronospora mycelium. At
first the examples were few in number, but at length they
were abundant. Soon after our results were published,
Mr. C. Edmund Broome, M.A., F.L.S., of Batheaston,
repeated the experiments, and obtained results precisely
the same with ours. Ultimately Mr. Broome went over
the ground a second time, and again obtained like results ;
at length, many other observers repeated the experiments,
and always with the same issue.
A very successful plan for procuring resting -spores,
and one which we have not known to fail, was last year
suggested to us by our friend Mr. A. Stephen Wilson. A
number of leaves must be taken from potato plants invaded
by Peronospora infestans, Mont.; these leaves must be
slightly moistened and placed one over the other near the
top, inside a bell-glass ; the bell-glass must then be put
xxxvi.] POTATO DISEASE, IL— PASSIVE STATE. 301
mouth downwards on any flat surface, and a saucer of
water placed underneath to keep the air humid by evapor-
ation. The result invariably is, that the Peronospora
mycelium within the potato leaves gives rise to an enor-
mous number of oospores or resting-spores ; and as the
leaves gradually decay, the decayed material swarms with
the Artotrogus hydnosporus of Montagne, — the resting
condition of the potato fungus.
During the early autumn of 1875 another important
fact in regard to the potato fungus came to light. On
making a rigid examination of every part of diseased potato
plants for oospores, we found them in great abundance
in the old exhausted seed tubers. In every other part of
the potato plant the oogonia were rare ; but in the old
sets the oogonia sometimes swarmed in myriads. The
explanation of this fact may be that the tubers were in
this instance planted with the perennial mycelium of the
potato fungus in their tissues. As this mycelium on
starting into growth could not produce conidia, being
underground, it spent itself in the tuber by a vast pro-
duction of oospores. There is no more certain position
for lighting on large colonies of resting-spores than in the
old exhausted seed tubers belonging to potato plants
destroyed by the Peronospora, or in the old diseased and
damaged tubers that are left in the field to rot, or are
incorporated in dung heaps as manure.
It is curious that at the very time when we were
making the above observations, Dr. Sadebeck of Berlin
found a parasite, named by him Pythium Equiseti, first
upon Equisetum arvense, L., and afterwards (as he at first
thought) upon living potato plants near Coblenz.
Whether the Pythium upon the Equisetum was the same
as the parasite upon the potatoes is uncertain, as Professor
Sadebeck could not transfer the parasite from one plant to
the other, neither could we do so on repeating his experi-
ments. Our impression is that the parasites are distinct.
P. Equiseti, Sdbk., was described and illustrated by us,
302 DISEASES OF FIELD & GARDEN CROPS. [CH.
from nature as British in the Gardeners' Chronicle for 20th
May 1876. It is extremely common in this country. Dr.
Sadebeck found his second fungus in the possession of the
living potato plant, for he wrote — (Untersucliungen iiber
Pythium Equiseti. Beitrdge zur Biologie der Pflanzen.
Breslau, 1875):— "In the first days of July 1875 I saw
at Metternich, not far from Coblenz, a potato field which
to all appearance was affected with the murrain ; a closer
examination, however, showed that the signs of the disease
were traceable almost entirely to Pythium Equiseti. The
anticipated Peronospora was not found on any of the
plants examined ; on the contrary, the Pythium was dis-
covered in a great number of plants and in all parts of
the plants." In a criticism published in the Journal of
Botany for March 1876, it was stated, in reference to this
part of the subject, that it had " lately been attempted to
connect this fungus (Pythium Equiseti, Sdbk.) with the
oospores of Peronospora infestans."
In the same year Mr. James Renny, a member of the
Scientific Committee of the Royal Horticultural Society,
was also studying a Pythium which he believed to be new,
and which was provisionally named by him P. incerlum.
This was exhibited at the Royal Horticultural Society
and Linnean Society, and according to the Journal of
Botany, 1876, p. 156, Mr. Renny considered his P.
incertum to be the same with the oospores found by us.
P. incertum was engraved by us in the Gardeners' Chronicle
for 1st July 1876, and we need hardly say is totally
different from Artotrogus.
Dr. Max Cornu, who at this time had our preparations
before him, said they reminded him of P. proliferum, De
Bary, another different fungus ; for an illustration of this
see Gardener's Chronicle, 1st July 1876. He also thought
they looked like the Myzocytium of Schenk.
At the time when these investigations were going on,
Professor de Bary himself was, by a commission received
from the Royal Agricultural Society of England, making
xxxvi.] POTATO DISEASE, I L— PASSIVE STATE. 303
observations upon the potato fungus for that Society. It
appears that Professor de Bary also lighted on what he
considered to be a fourth new species of Pythium, and
first seen by him in potatoes in 1874 and 1875. This
fungus the professor named P. vexans, and an original
description, with an illustration, is given in the Journal
of the Royal Agricultural Society for 1876, vol. xii. p. 252.
In 1881, in the Beitrage zur Morphologie und Physiologie
der Pilze, Professor de Bary has compared Artotrogus with
two other species of Pythium, both new, and named by
him P. micracanthum and P. megalacanthum. He says
the former may perhaps be Artotrogus.
It does not specially concern us here what these six
species of Pythium are, or whether they are new or dis-
tinct from each other or not. P. proliferum, D.By., is
probably distinct, but we can see no difference between
P. incertum, Ky., P. Equiseti, Sdbk., and P. vexans, D.By.
They are simply referred to here because some writers
have at times confused the potato oogonia seen by us
with one or other of these six organisms.
As P. vexans, D.By., appears to us to be the same
with P. incertum, Ky., and P. Equiseti, Sdbk., we here
reproduce at Fig. 132 the original illustration altered to
400 diameters for comparison with the Peronospora
oogonia given to the same scale in Figs. 130, 134, 135,
and 136. It will be noticed that the Pythium is smaller
in all its parts, and that the oogonia AAA are invariably
non-echinulate. The mycelium is thick and septate in
the Peronospora, and non- septate and very thin in the
Pythium.
The great point of difference is this: the Pythium
oogonia will, as soon as formed, and within twenty-four
hours germinate in water. In germination, a tube is some-
times produced, as at B, Fig. 132. In other instances when
the oospores of P. vexans, D.By., are kept for a few days
so as to ensure their complete maturity, they germinate by
ejecting a small transparent bladder, as at C ; the proto-
304 DISEASES OF FIELD & GARDEN CROPS. [CH.
plasm from the oogonium now pours into the bladder and
becomes quickly differentiated into six or eight little
zoospores ; the bladder then dissolves, the zoospores swim
away, and the short-life cycle of the Pythium is completed.
The phenomenon of speedy germination is foreign to
Artotrogus and the oogonia of Peronospora infestans, Mont.
The latter bodies do not remain transparent or germinate
at once ; on the contrary, they hibernate for at least ten
months, and during this long period of rest they increase
in size, become warted or echinulate, and attain a rich
palish-brown colour.
We will now leave the potato fungus as seen in a
X 400
FIG. 132.
Pythium vexans, D.By. Enlarged 400 diameters.
living potato leaf and take a fragment of a dead leaf, one
that has been destroyed by the Peronospora, such as may
be seen in fields and gardens in September, or, if preserved
with care, such as may be kept on a garden-bed till the
following June. A fragment of such a potato leaf is
illustrated in Fig. 133, enlarged, like Fig. 127, to 100
diameters. The upper surface of the leaf is shown at A,
the lower surface with two stomata at BB, and a small
hair belonging to the leaf is seen at C. Nearly all the
mycelium of the potato fungus has vanished ; a fragment
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 305
only, in a hibernating, septate state, is seen at D. The
transparent oogonia of the summer have now become
brownish ripe oospores or winter resting-spores of a larger
size. Six resting-spores are shown in the illustration, —
X- 100
FIG. 133.
Section through a fragment of old potato leaf, with resting-spores or
oospores, of Peronospora infestans, Mont., in situ.
Enlarged 100 diameters.
two in the transparent leaf hair, three in the intercellular
spaces of the leaf, and one inside a spiral vessel, in which
position it is extremely common to find them.
The perfectly mature resting-spores are best seen in the
X
306
DISEASES OF FIELD & GARDEN CROPS. [OH.
remains of old rotten tubers left in the fields from the
previous year, and commonly seen on the ground and
about dung-heaps and hedge-sides in March and April.
A section through a fragment of decayed tuber is shown,
FIG. 134.
Section through fragment of diseased tuber of potato, with starch granules
and oospores of Peronospora infestans, Mont., in situ.
Enlarged 400 diameters.
enlarged 400 diameters, at Fig. 134. One large oospore
is seen in an intercellular space, another in a cell amongst
the granules of damaged starch, and a third within the
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 307
coils of a spiral vessel. In colour the oospores are of a
beautiful palish-brown tint, like brown sherry ; sometimes
they are darker. The protoplasm within is at maturity no
longer seen as a loose, transparent, finely granular mass ; it
has become compact and slightly convolute, as illustrated
— ready under favourable conditions to burst the walls of
the oospore, and, by producing a germ-tube, reproduce,
after nearly a year's rest, the fungus of the potato disease.
It may be observed here how totally different this con-
dition of the fungus is from Pythium vexans, D.By.,
engraved to the same scale in Fig. 132. An original
microscopic slide of P. vexans, D.By., is preserved in the
Department of Botany, British Museum, South Kensington;
X- 4OO
FIG. 135.
Peronospora infestans, Mont.
Oospores of large size on slides A and B in the British Museum.
Enlarged 400 diameters.
it may there be compared with true potato oospores
furnished by the Rev. J. E. Vize and ourselves. Some-
times oospores of the potato fungus attain large dimen-
sions, as in Fig. 135. The right-hand example is on
slide A, the left-hand on slide B, in the British Museum.
On an examination of a large number of resting-spores
it will be found that the convolute mass of protoplasm
within, though generally in one coil, may at times be in
two or even three distinct portions, which on germination,
will produce one, two, or three germ -tubes, as shown
at A, B, C, Fig. 136, enlarged 400 diameters ; in other
instances the interior mass becomes differentiated into zoo-
308
DISEASES OF FIELD & GARDEN CROPS. [CH.
spores which escape, as at DD, and speedily come to rest
and germinate, as at E ; the germinal threads from oospores
and zoospores alike, when placed either on the foliage
or tubers of potatoes and kept uniformly moist and warm,
Peronospora infestans, Mont.
Oospores a year old germinating in summer.
Enlarged 400 diameters.
soon give rise to the fungus of the potato disease, and
cause discoloured patches of decomposition as the growth
proceeds. This condition of the fungus was described
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 309
and illustrated by us in the Gardeners' Chronicle for 8th
July 1876. Germinating resting-spores in a more advanced
state of growth are illustrated at F and G. On ger-
mination, the walls of the oospore break up into many
pieces, sometimes into fine dust. The first sign of ger-
mination is generally shown by the walls of the oospore
breaking into two hemispheres, or into three or more
pieces. In some instances more than one oospore exists
within the oogonium, and all oospores may at times pro-
duce zoospores, as in Cystopus. In the Gardener^ Chronicle
for 8th July 1876 a full description, with numerous
illustrations of germinating oospores, will be found.
Our experiments were at first objected to on the ground
that all the species of Peronospora were, it was said, so
sensitive to decay that they invariably perished with the
death of the supporting plant. This statement is now
known to be erroneous, and the resting-spores of the
Peronosporece are at this time always sought for, and almost
invariably found, in material which has been more or less
destroyed by the mycelium of the invading parasite.
This decayed material is obviously the only material in
which ripe oospores can be expected to occur. Every
part of the fungus, except the oospores, generally perishes
with the supporting plant ; the oospores or resting-spores
are left alive Upon or in the ground where potato material
has decayed, and in this position the oospores germinate
in June and produce the first conidia of the season. Such
of the conidia as are blown from the ground or from
decaying potato refuse on to potato plants, or certain allied
plants, produce disease ; such as fall in unsuitable positions
perish. The progress of the disease is, therefore, necessarily
at first extremely slow: it only progresses with rapidity
after the living potato plants are thoroughly invaded.
We have secured potato oospores direct from the ground
by observing water filtered through earth on which
diseased potato material has been allowed to decay.
The best time for seeing the reproductive organs of
310 DISEASES OF FIELD & GARDEN CROPS. [OH.
Peronospora infestans is in August and September. A
potato plant should be selected that has been destroyed
or reduced to putridity by the disease. This plant should
be taken up with a fork, and the exhausted seed tuber
from which the plant has arisen carefully sought out.
This seed tuber, or what is left of it, may be frequently
found reduced to a sort of transparent jelly, and this
jelly-like mass will in many cases be found swarming
with the living oogonia and antheridia of the potato fungus.
The fungus has attacked the leaves and proceeded down-
wards by the stems into the seed tuber from which the
plant originally arose, and there, having run its course, it
has produced resting-spores for the invasion of the follow-
ing year's crop of potatoes. It is much less common to
find resting-spores in the hard new tubers even when
discoloured by disease ; still it is quite possible to find
them even in new potatoes. Eipe resting-spores of the
potato fungus may be found with great ease in the spring
and early summer, in the fragments of diseased and
decayed potatoes picked up in the fields or about manure
and refuse heaps by hedge sides.
A germinating resting-spore may be compared with a
germinating seed of dodder. The dodder has enough
nourishing material stored up within its outer integument
to support an infant dodder plant for a short time. If
no suitable host plant is near, the young dodder perishes.
The first fruiting branch from a germinating resting-
spore of the potato fungus is in an exactly similar con-
dition, for, unless the spores or conidia are aided by the
wind to reach a potato or some other suitable plant, the
first-produced conidia perish at once. The resting-spores
of the potato fungus germinate in and upon the ground
at the precise time of the year when the potato plant is
in the best condition for infection. Habits of this nature
are extremely common and well known amongst parasitic
fungi.
We have as far as possible in this work avoided con-
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 311
troversial matter, only referring to disputed opinions
and deductions by giving without bias the views held on
both sides. Nothing is more damaging to the position of
science than disrespectful and hasty criticism and anim-
adversion. It is, however, necessary to inform our readers
that our views, as here advanced, in reference to the
nature of the oospores of the potato fungus and of
Artotrogus, have been criticised by Professor A. de Bary
of Strasbourg, in the Journal of the Royal Agricultural
Society of England, second series, vol. xii. p. 239, 1876.
Professor de Bary's objections to our views were the
following : —
1. He disapproved of our comparison of the potato
fungus oospores with the oospores of Protomyces ;
see our notes under Protomyces macrosporus, Ung.,
in this work.
2. He could not accept our drawing as illustrating the
potato fungus at all, as it presented an important
difference, he said, from the real Peronospora in-
festans, Mont.
3. Our assumed " oogonia," " antheridia," and " oospores,"
he said, were " bladders," and did not belong to
the potato fungus.
4. The mycelium, he said, was wrong, as the threads
bearing "oogonia" and "antheridia" were only
shown in local and not in anatomical relation with
each other.
5. He objected to the septa shown by us in the
mycelium.
6. He objected to the habitat we gave for the oospores,
i.e., in decayed potato material.
7. He stated in reference to Artotrogus that there was
no evidence of its nature as an oospore.
8. That there was no reason for considering it as be-
longing to the potato fungus.
9. That the smooth form of Artotrogus was a different
fungus from the echinulate one found with it.
312 DISEASES OF FIELD & GARDEN CROPS. [cri>
10. That the mycelium of Artotrogus was not septate, as
illustrated by us, but that it was like the my-
celium of Pythium vexans, D.By., i.e., in one con-
tinuous piece without septa.
11. That our "bladders" were more like the oospores
of the Peronospora of the vine or some Pythium.
Many other minor objections were advanced. Keplies
will be found in Nature for 27th April and 25th May 1876,
and in various numbers of the Gardeners' Chronicle for
1876. A report from the pen of the Rev. M. J.
Berkeley will be found in the Gardeners' Chronicle for
1st April 1876, p. 436. Our original illustrated papers
were published in the Gardeners' Chronicle for 17th and 24th
July 1875. Micro -photographs from nature, of potato
oogonia, and antheridia were published by us in the
Quarterly Journal of Microscopical Science for October
1875. The first sketches made by us are at Kew,
and the more carefully finished drawings are in the
British Museum. A moderate, independent, and just
resume of the whole subject was published by "W. Peard,
M.D., LL.B., in the Journal of the Bath and West of
England Society and Southern Counties Association, vol.
viii., third series, 1877.
For the discovery of the reproductive organs and
oospores of Peronospora infestans, Mont., the Royal Horti-
cultural Society of England awarded us their Knightian
medal in gold.
Some time prior to 1881 Professor de Bary changed
his views in reference to the nature of Artotrogus, as in
the latter year he agreed with us in illustrating and
describing the organism as a true oospore, as supported
on septate mycelium, and with an antheridium in "local"
and not "anatomical" relation to the thread of the
oogonium, precisely as we originally illustrated it.
The accompanying illustration, Fig. 137, enlarged 600
diameters, is copied from the Beitrage zur Morphologic und
Physiologic der Pilze, 1881, and it represents, from pi. 1,
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 313
Artotrogus from Professor de Bary's own hand ; it shows
the septate mycelium at AAA, the supporting thread of
the oogonium at B, and the distinct supporting thread
(not anatomically connected) of the antheridium at C.
At D the antheridium has projected a fecundating tube
through the outer wall of the oogonium to the oosphere
within. Professor de Bary found these bodies in
Lepidium, so they cannot be the true Artotrogus hydno-
sporus, Mont, which is borne only on the mycelium of
Peronospora infestans, Mont. They are the second form
of Artotrogus, peculiar to cruciferous plants, as first
B
•X-400
Fio. 137.
Artotrogus, as illustrated by Professor De Bary in 1881.
Enlarged 400 diameters.
detected by Mr. C. Edmund Broome, M.A., F.L.S., of
Batheaston, Bath, in. 1849, and referred to Artotrogus by
Dr. Montagne. They grow upon the mycelium of
Peronospora parasitica, Pers., and are the oospores or
resting-spores of the putrefactive fungus of the cabbage
tribe, as pointed out by the Kev. M. J. Berkeley in the
Gardeners' Chronicle for 1854, p. 724. They are illus-
trated, from nature, in this work in Figs. 31, 37, and 38.
No agricultural subject is more difficult to approach
than the possible curative or preventive treatment of the
potato disease. Cure, we may say, is utterly impossible,
314 DISEASES OF FIELD & GARDEN CROPS. [CH.
but prevention or palliation may be considered fairly
within reach, and this prevention can only be attained
by skilful culture and perfect winter storage.
Sometimes growers keep their potatoes in enormous
underground heaps called "pies ;" in these positions the
tubers frequently heat and rot ; in other instances diseased
potatoes are interbedded in dunghills, or dug into the
ground ; in all such cases the best means have been taken
for successfully propagating the disease. From all such
positions many millions of conidia of the potato fungus are
dispersed each June, whose special mission is to devastate
potato crops. The warmth and moisture of " pies " and
manure-heaps are the exact conditions required by resting-
spores for their maturation.
To prevent the annual recurrence of the potato mur-
rain it is in the highest degree necessary to destroy the
material which is undoubtedly swarming with myriads of
disease germs. This destruction should be effected by
burning, or, where burning is not practicable, deep burial
might be resorted to. No more fatal mistake can be made
by potato growers than leaving dead stems, leaves, and
tubers about in their fields, especially after a potato crop
has suffered from disease.
When cut sets are used at planting, the cut surface
should perhaps be allowed to heal or dry before planting,
or, if this is not convenient, the cut surfaces might be
quickly passed over a hot iron. It frequently happens
when diseased sets are used that the produce grows in a
healthy manner, with no trace of the murrain. There can
be little doubt of the existence of perennial spawn in
some of these examples, if not of resting-spores ; but in
some instances it would appear that neither spawn or
spores work much mischief direct from the seed tuber
when buried. In some instances both mycelium and
oospores must be dead.
Any cure of the murrain in invaded potato plants is
quite hopeless, for in this disease the substance of the
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 315
potato tuber is decomposed, and it is impossible to replace
rotten tissues with sound. With the object of prevention
in view, hardy varieties which have not exhibited disease
should be selected and reselected. They should be grown
where possible in well- drained dry soil, and mineral
manure should be used. As darkness, heat, and humidity
are highly favourable to the growth of the Peronospora,
all potatoes should be stored in perfectly dry, airy places,
in positions where light is not entirely excluded. Potatoes
should never on any account be stored in heaps or in the
damp holes in the ground termed " pies."
During the last year or two great attention has been
directed to the potato plant and its treatment under
disease, chiefly by Mr. J. L. Jensen of Copenhagen, and
Mr. C. B. Plowright, M.K.C.S., in the pages of the
Gardeners' Chronicle, and more lately by Mr. J. G. Baker,
F.R.S., in the Journal of the Linnean Society, and to these
communications it is necessary for us to advert.
The paper by Mr. J. G. Baker, F.K.S., F.L.S., written
at the request of Earl Cathcart, in the Journal of the
Linnean Society (" Botany," vol. xx. p. 489), is called " A
Review of the Tuber-bearing Species of Solanum" Mr.
Baker reviews the species and varieties of tuber-bearing
Solanum geographically, beginning under Chili with — 1.
Solanum tuberosum, L. ; 2. S. etuberosum, Lind. ; 3. S.
Fernandezianwn, Phill. ; 4. S. Maglia, Sch. ; and 5. S.
Collinum, Dun. Brazil — 1. S. Commersoni, Dun., and its
var. S. Ohrondiiy Carr. Peru, Bolivia, Ecuador, and
Colombia — 1. S. tuber osum, L. ; 2. S. immite, Dun. ; 3.
S. Colombianum, Dun. ; 4. S. Valenzuelce, Pal. (S. Maglia
Sch.) Mexico — 1. S. verrucosum, Sch. ; 2. S. suaveolens,
K. and B. ; 3. S. stoloniferum, Sch. ; 4. S. demissum, Lind. ;
5. S. utile, Klot. ; 6. S. squamulosum, M. and G. ; 7. S.
cardiophyllum, Lind. ; 8. S. oxycarpum, Sch. South- West-
ern United States — 1. S. Fendleri, A. Gray ; 2. S. Jamesii,
Torr. Of these plants Mr. Baker considers there are only
six genuine species in a broad sense, viz. — 1. Solanum
316 DISEASES OF FIELD & GARDEN CROPS. [en.
tuberosum, L. ; 2. S. Maglia, Sch. ; 3. S. Commersoni,
Dun. ; 4. S. cardiophyllum, Lind. ; 5. S. Jamesii, Torr. ;
and 6. S. oxycarpum, Sch. Mr. Baker states that of all
known species of Solanum, only six produce potatoes,
and the remainder propagate themselves by their flowers,
fruits, and seeds. In reference to the Mexican S. demissum,
Lind., Dr. Lindley, in his notes on the wild potato, pub-
lished in the Journal of the 'Royal Horticultural Society,
vol. iii., 1848, says that when grown in England it was
attacked by the disease in July, and exhibited the charac-
teristic black blotches in a worse degree than any other
in the garden. The runners were also affected. The
Mexican S. cardiophyllum, Lind., on the other hand, was
not attacked by the disease. Dr. Lindley concluded that
neither renewal of seed, introduction from foreign countries,
or treatment in the earth afford any guarantee against the
attacks of the disease. Sir Joseph Hooker has stated that
S. Maglia, Sch., when grown at Kew, did not yield tubers
for the first two years. It has now been grown there for
twenty years side by side with the common potato, S.
tuberosum, L., and maintains its individuality. It does
not, however, produce berries. Both plants are natives
of Chili ; but Mr. Baker points out the very important
fact that whilst S. tuberosum, L., is a plant of the hills of
the interior, S. Maglia, Sch., grows in the near neighbour-
hood of the coast. This is the potato found by Mr.
Charles Darwin in the Chonos Archipelago in south lati-
tude 44°, 45°; and Mr. Baker throws out the happy sug-
gestion, and one which we hope will be generally adopted,
that S. Maglia, Sch., should become popularly known as
Darwin's potato, a plant which, as far as climate is con-
cerned, Mr. Baker thinks without doubt is better fitted to
succeed in England and Ireland than S. tuberosum, L., a
plant belonging to a comparatively dry climate. Both &
Maglia and S. Commersoni, Dun., yield an abundant
supply of edible potatoes. Mr. Baker suggests that these
two species should be brought into commerce and thor-
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 317
oughly tested as regards their economic value, both as
distinct types and when hybridised with the innumerable
forms of S. tuberosum, L.
Mr. Charles Darwin described Solanum Maglia, Sch.,
in the 1835 octavo edition of the Voyage of the Beagle, p.
288. He there writes: — "Chonos Archipelago. — The
wild potato grows on the islands in great abundance on
the sandy, shelly soil near the sea-beach. The tallest
plant was 4 feet in height. The tubers were generally
small, but I found one of an oval shape 2 inches in dia-
meter. They resembled in every respect and had the
same smell as English potatoes ; but when boiled they
shrunk much and were watery and insipid, without any
bitter taste. They are undoubtedly here indigenous.
They grow as far south, according to Mr. Low, as latitude
50°, and are called Aquinas by the wild Indians of that
part." We give an illustration, natural size, of the
flowers, foliage, and tubers of Darwin's potato, Solanum
Maglia, Sch., in Fig. 138 (frontispiece). Sir Joseph
Hooker, in writing of this species in the Botanical Maga-
zine for May 1884, says the tubers were first sent by Mr.
Alexander Caldcleugh from Chili to the Royal Horticul-
tural Society in 1822. Mr. Caldcleugh's tubers were
cultivated in manured soil at the Royal Horticultural
Gardens, where two plants yielded about 600 tubers of
about the size of a pigeon's egg and under, which had,
when boiled, the flavour of a common potato. Tubers of
the same species were given to Kew in 1862 by Dr.
Sclater, F.R.S., — these were grown in the sandy soil of
the pleasure-grounds without manure. Experiments are
now being carried out under the auspices of the Royal
Agricultural Society to improve the qualities of the
potato, especially in its power of resisting attacks of the
potato disease, by crossing S. tuberosum, L., with its allies,
and amongst them with S. Maglia, Sch. Sir Joseph
Hooker says that 8. Maglia, Sch., flowers freely every
autumn at Kew, and yields watery, scarcely edible potatoes,
318 DISEASES OF FIELD & GARDEN CROPS. [on.
Mr. Baker lias not seen S. Commersoni, Dun., in a
living state, but he says that 8. Ohrondii, Carr., recently
described and illustrated in the Revue Horticole, 1883, pp.
496-500, Figs. 99, 100, and afterwards adverted to at
some length by the horticultural press of this country, is
the same plant. Sir Joseph Hooker, in the Flora Antarc-
tica, reduces it to a mere form of the common edible
potato, 8. tuberosum, L. Tubers of this plant were lately
brought by M. Ohrond, a French naval surgeon, from the
island of Goritti, at the mouth of the Kio de la Plata,
and grown at Brest by M. Blanchard, gardener-in-chief of
the Marine Hospital, who writes as follows : — " From the
time of its importation I have cultivated the plant, or
rather left it to itself to grow, for it is almost impossible
to destroy it when once it has become established in a
piece of ground. Each year, at the end of June or the
beginning of July, I have collected the tubers ; but the
rootstock creeps so widely that always plenty have re-
mained in the ground to furnish stock for another year.
It is my belief that it would be easy to improve the
tubers by simply cultivating them. Already the culti-
vated tubers are much better than those which I received
from M. Ohrond. The wild tubers were scarcely bigger
than small walnuts, but some of those of the cultivated
plants have attained the size of small hen's eggs. I may
add that the tubers are quite palatable, with a taste of
chestnuts, but leaving in the mouth a slight flavour of
acidity, like that of a potato that has sprouted. My
workmen and I have tried them both boiled and baked
in the oven ; the latter are preferable. As to the hardi-
ness of the plant it is complete — at least here at Brest.
During the winter of 1881, when the thermometer fell
two degrees centigrade below freezing point, the tubers
took no harm, and up to the present time the plant has not
been found to suffer in the least from disease."
Mr. Baker thinks that our present method of potato
culture unfits the plant to resist disease by exciting the
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 319
plant grown in crowded positions, to a large production of
tubers. He thinks, in agreement with suggestions often
made by writers on horticultural subjects, that the absence
of flowers and berries on cultivated potatoes is a proof
that the plant is in an unnatural and disorganised con-
dition. Mr. T. A. Knight has shown, in the Philosophical
Transactions for 1806, p. 297, that the varieties of pota-
toes which uniformly produce neither flowers or berries
may be caused to produce them by preventing the growth
of tubers and runners amongst the fibrous roots.
The writings of Mr. J. L. Jensen of Copenhagen, as
laid before the horticultural world by Mr. C. B. Plow-
right, M.R.C.S., have chiefly had reference to what Mr.
Jensen has termed " Protective Moulding."
It is a common practice amongst potato growers to
earth-up potatoes, usually by driving a plough between
the rows. This earthing-up not only helps to support
the potato haulms in an upright position in the rows, but
it keeps the potato tubers from the light and consequent
greening ; it obviously keeps the potatoes free from the
numerous injuries they sometimes sustain from the attacks
of wire-worms, slugs, snails, rabbits, rats, moles, and other
animals, and from cracking after exposure to sun, hail, rain,
and wind. A potato when scratched, bitten, or bruised,
and with its inner substance exposed, is much more liable
to the attacks of fungus parasites than examples with the
natural armour of a perfectly whole skin. We have
shown that the mycelium of the fungus enters the potato
plant by the organs of transpiration, and sometimes even
pierces the epidermis or bark in its effort to reach the
interior. It follows, then, that any injury to the leaves,
stem, or tuber, even if the injuries are of the most micro-
scopic proportions, must aid the parasite in its efforts to
gain access to the inner tissues of the host. The fact has
long been accepted by a large section of potato growers
that earthing-up has also a marked tendency to keep
potato tubers free from the murrain, even when the
320 DISEASES OF FIELD & GARDEN CROPS. [OH.
haulms have been completely destroyed by the parasite.
A prize essay on this subject by Dr. Jeffrey Lang will be
found in the Journal of the Royal Agricultural Society for
1858. Dr. Lang experimented to the full depth of a
spade or fork, and double ploughed, the potatoes being
early earthed-up, with the result that few or none of the
potatoes so grown were diseased. Dr. Lang mentions the
case of " a man at Whilborough " who on dry days, in
order to save his potatoes, instead of digging them tip,
earthed the stalks up very high, and so effectually saved
his crops. "It was observed," writes Dr. Lang, "that
no potato covered with more than 3 inches of soil was
ever diseased," and " I have seen scores of potatoes dug,
but I have never seen or heard of one diseased potato
being found 4 inches under the surface of the ground.
It will be at once seen — and too much stress cannot be
laid on the fact — that the disease is in an exact ratio to
the proximity of the tubers to the surface." Dr. Lang
also experimented with tubers in the following manner : —
Three series were planted three deep, and covered with
two-and-a-half inches of soil. On two series diseased potato
leaves were placed, and then supplied with water through a
fine rose ; the third set were covered with a slate. In all
three series the under layers of potatoes were found un-
diseased ; all were undiseased under the slate ; whilst the
upper layers covered with diseased potato leaves were
found to be much affected or quite rotten. He ends the
essay by saying, " Earthing-up repeatedly with fine earth
is the only effectual preventive to the ravages of the
disease."
During the last quarter of a century the subject has
been frequently adverted to in the horticultural and agri-
cultural papers. In some instances a good result has been
recorded, in others a negative one. As commonly prac-
tised, the harvest of large tubers is said to be lessened,
and in some quarters the extra expense, care, and labour
has been greatly objected to. Professor W. G. Farlow, in
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 321
the Bulletin of the Bussy Institution, part iv., refers to deep
planting, which is much the same with earthing-up, and
writes, p. 336 : — " Theoretically, it would appear to be
an advantage to plant deep, that the tubers may have less
chance for being infected from spores which have fallen
from the surface. Practically this does not work well,
but potatoes planted near the surface do best. However,
the plan tried by some cultivators in England, with
apparently good result, of hoeing the earth up over a
good part of the tops as soon as the rot appears, is worthy
a trial."
During the last two years Mr. J. L. Jensen, a gentle-
man of Copenhagen, has, chiefly through the mediumship
of Mr. C. B. Plowright, of King's Lynn, again placed the
subject of earthing-up prominently before the agricultural
and horticultural public of this country.
Some of Mr. Jensen's views have been opposed by Mr.
William Carruthers, F.K.S., the Keeper of the Depart-
ment of Botany at the British Museum ; by Mr. George
Murray, also of the British Museum ; and by ourselves, as
contrary to fact, and contrary not only to the experience
of botanists, but, what is of more importance, to the ex-
perience of practical potato-growers and dealers. Person-
ally, we advocate, and have always advocated, careful
earthing-up ; it agrees with the practice approved by
many potato growers. That earthing-up is, however, not
esteemed by all seems shown by the fact that neither Dr.
Lang's suggestions here, or Professor Farlow's in America,
have been generally adopted.
Mr. Jensen says a high and sharp ridge of earth
should be thrown up round the potato plants a little
before the disease has appeared in the foliage, or at least
at the very first appearance of it. " The usual moulding
practised in all countries," writes Mr. Jensen, " is a flat
moulding, by which the uppermost tubers are only
covered by one or two inches of earth ;" but the Jensen-
ian system requires, after a preceding flat moulding, a
y
322 DISEASES OF FIELD & GARDEN CROPS. [CH.
high and sharp moulding, by which the upper surface of
the uppermost tubers is covered with about 5 inches of
earth. To effect this, it is necessary that the ridge be so
high that the top of it is 10 inches or 12 inches above
the surface of the adjoining furrow, whilst the ridge must
be very broad at the bottom : this system also requires
that the tops of the potatoes shall be moderately bent to
one side, with a view to prevent the rain-water from run-
ning down the stems and thus carrying the spores outside,
as Mr. Jensen thinks, to the tubers. By this contrivance
more spores will fall between than upon the ridges. Mr.
Jensen advises that the potatoes be not lifted before the
diseased foliage has quite withered, because the tubers
will become sprinkled with the fungus spores from the
leaves and stems. " For six days," Mr. Jensen writes,
" the harvested tubers will appear to be sound, but on
the seventh or eighth day, according to temperature, they
will suddenly show marks of the disease. It is not even
sufficient that the leaves are withered before the lifting ;
they must have been so for three or four weeks, otherwise
many spores will be found capable of germinating, and
thus be dangerous to the tubers when the latter are taken
out of the ground." Mr. Jensen sums up his views with
the following general rules : —
1. The ground must be thoroughly worked, so that the
potatoes may be planted in friable earth, which
affords a better means of protection than a lumpy
soil.
2. The potatoes should be planted (pretty early) at a
distance between the rows of at least 28 in. or 30
in. A greater distance is not required by the
system, but if closer it would impede the protective
moulding.
3. The first moulding must be flat, so that the formed
ridge be broad on the top and only about 4 in.
high. This moulding may be repeated if it is
thought advisable.
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 323
4. The protective moulding must be applied as soon as
the disease-blotches make their appearance on the
leaves of the haulm. If this has not occurred
before wheat-harvest-time, the moulding ought to
be executed then, without waiting for the appear-
ance of the disease-blotches.
5. The protective moulding is performed by throwing
up from one -side of the row of plants a high ridge
with a broad base, and running to as sharp a point
at the top as possible. The covering of earth
thereby produced over the upper surface of the
uppermost tubers must be about 5 in. to begin
with ; later, by the settling of the earth, and by
sliding down, it will, as a rule, preserve a thick-
ness of about 4 in. Simultaneously with this
moulding, the potato-tops are gently bent over
towards the opposite side of the row, so as to give
the top at least a half-erect position.
6. The flat and the protective moulding, where potatoes
are only grown on a small scale, may be done
with a hand-hoe ; on a larger scale these opera-
tions ought to be performed with a moulding-
plough, the " Protector," which is constructed to
meet the necessities of the described system.
7. In order to prevent after-sickness, which may often
be exceedingly great, the potatoes must not be
lifted before about three weeks after the last
leaves in the potato-field are withered.
8. If the potato-tops are cut off and carried away, which,
for the sake of the quantity and quality of the
crop, ought not to be done before the leaves, in
the main, are withered, the lifting may, as it
seems, without danger of after-sickness, take place
about six days after such removal.
Where this method of culture can be conveniently
practised, we think the result can be no other than bene-
ficial. We look upon the difficult process of bending the
324 DISEASES OF FIELD & GARDEN CROPS. [CH.
haulm to one side as not dissimilar in its aim and results
from the common practice of removing it altogether. In
both instances it reduces the yield of large tubers, and at
the same time, as we think, has a tendency to prevent
the mycelium of the potato fungus reaching the tubers
by the inside of the haulm.
Every one who has experimented with potatoes knows
that it is possible to infect tubers with the disease from
the spores produced on the leaves. This infection is more
readily produced in the eyes where the skin or bark of the
tuber is thin and delicate. Infection, however, from the
outside of the tuber inwards, is the exception and not
the rule. From our own experience, we believe the
disease generally reaches the tubers by travelling down
the interior of the stem, and that in the majority of in-
stances the interior of the tuber is the first part affected,
and the disease then works from the inside outwards.
It commonly happens that potatoes are harvested in
an apparently sound condition, but during the winter or
early spring the stored tubers are destroyed by the fungus
of the murrain bursting through the skin or bark from
the inside to the outside. It is also a fact of common obser-
vation that when a large number of apparently sound
potatoes are cut for seed, disease patches, either large or
small, may be seen in the central parts of the tuber, with
no apparent connection with the sound parts outside. At
the beginning of January 1884 we received a letter from
one of the largest potato dealers in this country, com-
plaining of a large crop of unsalable potatoes ; the tubers
were apparently perfectly sound outside, but full of disease
within. A selection of the tubers was also sent on for
our inspection, and the sharpest searching failed to detect
any disease patches outside ; the interiors of the tubers,
on the contrary, were full of dark-brown corroded mur-
rain patches. In this bad case, — and we know of very
many similar ones, — it seems impossible that the disease
could have been derived through the bark of the tuber,
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 325
which, in every instance, was apparently perfectly intact.
Another instructive case is given in the Gardeners'
Chronicle for 1st March 1884, p. 283.
Mr. Jensen states that it may be said with full certainty
that the disease either never reaches the tuber by growing
through the stems, or, if it does so happen in a few single
plants, which he says, to his knowledge, has never been
proved, it is of so rare an occurrence as to be of no prac-
tical consequence.
A series of test experiments has been instituted this
year at the garden of the Royal Horticultural Society at
Chiswick, where Mr. Jensen's instructions will be rigidly
adhered to. At the time of planting, when some of the
tubers were cut for sets, it was observed that the disease
was apparent in the interior, with no trace whatever of
disease on the skin or bark. The bending over of the
brittle haulm we consider a delicate operation, which is
hardly suited for unskilled rustics ; and if the easily-
broken stems become severed instead of bent, the process
becomes identical with the old plan of removing the haulm
altogether. A broken haulm is synonymous with the
destruction of leaves, and if the leaves are lost no starch
can be formed for subsequent storage in the tubers. The
mere gathering of the stems together for bending over
must be injurious to the potato plant, as it prevents the
leaves from receiving their requisite amount of light.
We consider the immunity from disease of earthed-up
potatoes, with bent haulms, is less owing to the power
possessed by the earth of filtering the fungus spores, and
so preventing them from reaching the tuber, than to the
effect of the earth in keeping the tubers whole and sound.
Mr. Jensen has also devised what he terms a disinfecting
apparatus. This has been described and illustrated by
Mr. C. B. Plo wright in the Gardeners' Chronicle for 5th
April 1884. According to Mr. Jensen, a temperature of
77° F. kills both the mycelium and spores of the Perono-
spora, provided the heat be continued for a sufficient
326 DISEASES OF FIELD & GARDEN CROPS. [on.
time, and that a considerably higher temperature does
not impair the germinating power of the potato. Mr.
Jensen states that the exposure of tubers to a dry heat of
100° or 105° F. for four or five hours is the best mode of
disinfecting them. This disinfection, it appears, should
be carried out in an oven, which has to be carefully
watched the whole time, and it is necessary that the heat-
ing process be continued for some hours to ensure the
deeper parts of the tubers being raised to the requisite
temperature. Mr. Jensen recommends for small experi-
ments the construction of a double box capable of hold-
ing water in its interior ; the space between the inner
and the outer box being filled with some non-conductor
of heat, as chaff. The inner box is filled with water at
a temperature of 100°- 120° F. Into this are placed tin
or zinc cylinders, about five inches in diameter, containing
the tubers to be disinfected, and in these cylinders the
tubers are allowed to remain. By the employment of
narrow cylinders like the above, the potatoes are more
readily heated than is the case if wider ones are used. A
thermometer in the water is essential to ascertain its
temperature. It may require an addition of warm water
once or twice, according to the heat of the surrounding
atmosphere, and the efficacy of the non-conducting medium
employed. In a later communication Mr. Plowright
states that the tubers must, for not less than four hours,
have a temperature not below 104° F., and if this rises^
to 115° F., no harm will be done to them, and that it is
not safe to go beyond 130° F.
The apparatus has, of course, been founded on the idea
that many potatoes apparently sound at the time of plant-
ing, yet contain living hibernating fungus mycelium, and
that this mycelium grows with the young potato plant,
and such diseased plants act as centres from which the
potato fungus spreads. Our observation and experience
of growing potatoes does not lead us to look upon this as
always the case ; and we must leave it for practical men
xxxvi.] POTATO DISEASE, II.— PASSIVE STATE. 327
to decide whether repeated earthing -up, skilful haulm-
bending, and disinfecting with a hot-water apparatus and
assistants watching a thermometer, can be made a com-
mercial success.
Disinfection, even if effectual, merely secures a possible
healthy start of the young potato plant, — it by no means
secures the potato from the attacks of spores in June or
July ; these spores may come from neighbours' fields,
where the potatoes have not been disinfected, or from
tomatoes or other plants, and so all the labour of disin-
fection may be lost. It is obvious that, unless all the
seed-potatoes in Britain are disinfected, little or no good
can accrue from the use of a few sets of apparatus. Mr.
Jensen thinks this part of the subject " cannot be con-
sidered an unworthy object for legislation" (Gardeners'
Chronicle, p. 616, 10th May 1884), which we suppose
means that farmers should be compelled by law to use a
hot- water disinfecting apparatus before planting potatoes.
Mr. Jensen appears to believe, judging from Mr Plow-
right's communications, that the planting of diseased sets
causes an early appearance of the disease, and that disin-
fection has a tendency to make the disease late. He
seems to conclude from this that constant disinfection
would at last make the fungus so late in its appearance
that the potatoes would be mature before the fungus could
grow. We are inclined to think, however, that the fungus
would change its nature so as to agree with the new habit
of the potato. Parasites always modify their habits to
suit any change of nature in their host.
We have shown that Mr. Jensen's views were more or
less anticipated by Dr. Jeffrey Lang more than a quarter
of a century ago, and the earthing-up system has been
advocated both here and in America. We are inclined
to think that practical agriculturists would never have
dropped this treatment if it had contained the elements
of commercial success.
Mr. Jensen's views will be found reported at length,
328 DISEASES OF FIELD & GARDEN CROPS. [CH.
with various comments from correspondents, in the
volumes of the Gardener^ Chronicle for 1883 and 1884.
Personally, we are sorry to say that we esteem many of the
statements put forward by Mr. Jensen as contrary to fact.
"We say this without disrespect to Messrs. Jensen and
Plowright, who have certainly made a vigorous attempt
to ward off the attacks of the potato disease ; and for this
they deserve the hearty thanks of all practical men.
Their communications are, however, too voluminous and
involved for any complete reply here. Practical potato
growers, if so inclined, must sift, weigh, and compare the
numerous statements brought forward for themselves.
We highly esteem Mr. Baker's suggestion regarding the
potato termed "Darwin's Potato," Solanum Maglia, Sch.,
and 8. Commersoni, Dun. The evidence brought forward
by Mr. Baker seems to indicate that S. Maglia, Sch.,
would well suit our humid climate ; and £ Commersoni,
Dun., appears to naturally resist the Peronospora.
In 1874 and 1875 a report was widely spread in this
country through one of the scientific societies that Pro-
fessor De Bary of Strasbourg had discovered an alternation
of generations in the life cycle of the Peronospora of the
potato murrain, — that he had found that the fungus
passed one part of its existence on clover, just like the
rusts and mildews of corn are assumed by some to live on
barberry bushes and borage. The scare had no effect on
the men of science in this country ; the statement was
received in silence, like the statements regarding the Colo-
rado beetle. In America, however, the case was different,
for when the report of the assumed discovery reached
that country it was believed, and notices appeared in the
agricultural reports published at Washington, and in
several agricultural journals in different parts of the
country, warning farmers — according to Professor W. G.
Farlow in the Bulletin of the Bussy Institution, part iv. —
that in consequence of Professor De Bary's discoveries no
potatoes should be planted after clover and other fodder
xxxvi.j POTATO DISEASE, II.— PASSIVE STATE. 329
crops. Professor Farlow has stated, and no doubt cor-
rectly, that Professor De Bary never made the statements
attributed to him by a few of his friends in this country.
It ought not to be difficult on well-conducted farms
to keep the fields clear from rotting potato refuse. The
working men and boys should be taught, as a rule of
the first importance, that all potato refuse should be
scrupulously gathered together and either burnt or
deeply buried. Stones are always gathered together on
farms by boys and girls. When vegetable refuse is in-
corporated with dung, this material by its warmth and
moisture keeps the germs of nearly every known plant
disease alive and in good condition through the winter
for a renewed burst of vitality in the spring or early
summer. It has been said that when potatoes are grown
near chemical works they are frequently free from disease,
as sulphurous acid gas or some other gaseous impurity
proves fatal to the fungus of the murrain.
CHAPTER XXXVII.
PARASITIC FUNGI AS FOUND IN A FOSSIL STATE.
IT is a curious fact that representatives of some of our
common parasitic fungi are found in a silicified state in
fossil plant stems and roots of great antiquity. Some of
these parasites were in existence in company with the
higher cryptogams in Palaeozoic times. Indeed it is prob-
able that some fungi, not dissimilar in structure from
fungi which are now parasitic, led in remote geological
times a non- parasitic life upon the ground. We still
have a Botrytis named B. terrestris, Pers., which is fre-
quent on the naked ground. The species belonging to
Botrytis are very similar with the species described under
Peronospora, and they were till quite recently all grouped
together.
The late Mr. Charles Darwin informed us that more
than forty years ago, Mr. Robert Brown, then Keeper of
the Department of Botany at the British Museum, showed
him silicified fungus mycelium in slices of fossil wood.
Mr. William Carruthers, F.R.S., the present Keeper of
Botany at the British Museum, South Kensington, has
described silicified fungus mycelium resembling that of
a Peronospora found in the tissues of a fossil fern named
Osmundites Dowkeri, Carr., from the lower Eocene strata
of Herne Bay. The same gentleman has also detected a
fungus in a fossil Lepidodendron from the coal measures ;
and Mr. Butterworth, of Oldham, has also met with a
fungus in the vascular axis of Lepidodendron. A portion
of the latter example was drawn by us, and the drawings
are now in the Museum of Practical Geology in Jermyn
OH. xxxvii.] PARASITIC FUNGI. 331
Street London. Engravings from the same transparent slice
were published by us, with a description, in the Gardeners'
Chronicle for 20th October 1877. Mr. Carruthers also
published an extremely small engraving and a brief de-
scription of the fungus in his printed address, read before
the Geologists' Association in 1876. We have named
this parasite Peronosporites antiquarius, W.Sm. To us
the mycelium appears to be distinctly septate, and the
large globular oogonia or zoosporangia appear to show clear
traces of zoospores within their walls.
Mr. J. T. Young, F.G.S., the owner of the transparent
slice of fossil Lepidodendron, has recently replaced the
example in our hands for a new illustration ; and our
engraving at Fig. 139, enlarged 400 diameters, has been
made direct from the microscope.
Notwithstanding criticisms to a contrary effect, we have
no hesitation in repeating, after a renewed and prolonged
examination of the preparation, that traces of zoospores are
distinctly visible in many of the oogonia ; there is no reason
why they should not exist, but good reason why they
should ; the mycelium is septate ; and the oogonia, as in
all Permosporew and Saprolegniece, are cut off from the sup-
porting threads by distinct septa. The slice of Lepidoden-
dron from which our illustration is taken has a large
number of free oogonia in different parts of the silicified
tissue ; such free oogonia or zoosporangia are very com-
monly seen in Peronospora, as in P. ganglioniformis, B.
The zoospores in some of these free isolated examples are
much more distinct than in the characteristic group, en-
graved to show the oogonia only, in Fig. 139. The
traces of zoospores, seen in Peronosporites, exactly agree in
size with the zoospores of Peronospora infestans, Mont.,
Figs. 128 and 136. The genus Peronospora is in close
and obvious relationship to the Saprolegniece, one member
of which, Saprolegnia ferax, Kutz., is the cause of the
salmon disease. Professor de Bary has even said that
facts do not exclude the possibility of the fungus of the
332 DISEASES OF FIELD & GARDEN CROPS. [CH.
FIG. 139.— FOSSIL PARASITIC FUNGUS.
Peronosporites antiquarius, W.Sm., in a slice of fossil Lepidodendron from
the Coal Measures. Enlarged 400 diameters.
potato disease being one of the Saprolegniece (Journal of
the Royal Agricultural Society, p. 249, 1876). It is some-
XXXVII.]
PARASITIC FUNGI.
333
times (as when the plants are not fully developed) im-
possible to distinguish between one genus and the other.
Peronosporites, therefore, has without doubt relations with
the Saprolegniece, as correctly pointed out by Professor W.
•X-400-
FIG. 140.— SPORANGIUM OP A FOSSIL FUNGUS.
Protomycites protogenes, W.Sm., in a slice from a rootlet of a fossil
Lepidodendron. Enlarged 400 diameters.
C. Williamson, F.K.S., in the Philosophical Tratisactions of
the Royal Society, 1881.
We have a second representative of fungi of enormous
antiquity in a transparent silicified slice of a rootlet of
Lepidodendron from the coal measures, now in the British
334 DISEASES OF FIELD & GARDEN CROPS. [en.
Museum at South Kensington. This slice exhibits
numerous unusually large sporangia of a fungus not to
be distinguished from Protomyces. Very little mycelium
can be detected ; and many of the sporangia of the fungus
are situated in positions where the tissues of the host
plant have apparently, but perhaps not really, decayed.
We have illustrated one sporangium of this fungus, which
may be named Protomycites protogenes, W.Sm., at Fig.
140, enlarged 400 diameters (protogenes, first produced or
primaeval). In most of the silicified examples an outer
or exospore, and inner or endospore are distinctly visible.
This fungus presents some analogy with the alga
named Chlorochytrium Lemnce, Cohn., which grows within
the fronds of duckweed, the spores from the zoosporangium
FIG. 141.
Diagram showing development of simple fungi by cell-division.
of which conjugate or fuse in the style of the zoospores
(or zygozoospores) of Protomyces, and so produce zygospores.
The observer should, however, be ready to distinguish,
between mere fusing, which is very common in fungi,
and true conjugation, which is by no means common.
Although these two fungi have been detected in
Palseozoic rocks, it must not be concluded that they
are the simplest known forms of primal fungi. In
Peronospora and Protomyces alike, sexual organs occur;
and the fact of a separation of sexes shows a great ad-
vance upon a primordial form. Besides, the members
belonging to the two genera are parasites, and doubtlessly
lived in Palaeozoic times, as their representatives do now,
upon the living tissues of more highly-organised plants.
xxxvn.] PARASITIC FUNGI. 335
But there must have been a long antecedent time, when
the lowest fungi were non-parasitic, and grew upon the
moist warm ground as Botrytis terrestris, Pers., sometimes
does now. In those far-off times the primordial plant
was probably a mere microscopic cell or thin sac resting
on the moist surface of the earth, as illustrated at A,
Fig. 141. It probably increased by division, as at B,
and redivision, as at C ; each of the four parts soon be-
coming distinct, as at D, and each segment speedily reach-
ing the original size and form, as at E. Or it might have
increased by budding, like yeast. From this simple
beginning many observers believe it probable that all
plants have been developed. The primal cell might
have been a fungus, an alga, or a form occupying an inter-
mediate position between fungi and algae, as both fungi
and algae may have originated from a primal and at pre-
sent unknown stock
CHAPTER XXXVIII,
CONCLUSION.
WE have now rapidly passed in review some of the most
familiar forms of disease as seen in our field and garden
crops, and not a few of our readers may possibly think
the details as described both complicated and difficult.
Yet on careful study it will be found that the courses of
all diseases more or less follow one or two simple general
plans. The details may vary and the colours may be
changed, but the chief outlines are not essentially different
from each other.
It is only in the knowledge obtained after completely
mastering the life history of each disease that any pre-
ventive remedy against disease can be hoped for. With
a full knowledge of the character and habit of an enemy,
it can be fought under favourable circumstances, as in a
bright light. Without the proper knowledge it is like
fighting against a powerful, unknown, and merciless foe
in the dark.
One point that must impress every reader is the
extreme, almost inconceivable, smallness and attenuation
of the parts of some of the most destructive of our field
and garden fungi. To give an idea of this smallness, we
have in Fig. 142 engraved the foot of a common house-
fly, with its hairs and claws, enlarged 100 diameters. At
AA are seen six of the spores or conidia of the potato
fungus, Peronospora infestans, Mont. Each of these spores
contains within itself, on an average, eight other little
spores or zoospores, illustrated as free from the investing
spore or conidium at B, and each of these smaller spores
CH. XXXVIII. ]
CONCLUSION.
337
has two inconceivably fine cilia or vibrating hairs, by
which it can propel and guide itself over any moist
surface. The large spores of the putrefactive fungus of
lettuces, Peronospora Schleideniana, Ung., are shown at C ;
others of the putrefactive fungus of clover P. exigua,
X-IOO
FIG. 142.
Foot of house-fly, with the spores of various parasitic fungi.
Enlarged 100 diameters.
W.Sm., at D ; and the spores of the smut fungus of
grain, Ustilago carbo, Tul., at E.
Every one who has walked amongst potato plants must
have noticed the small green fly or plant-louse, Rhopalosi-
phum dianthi, Schrank, shown, the natural size, at B, Fig.
143. Some of the female lice possess wings, as shown,
Z
338
DISEASES OF FIELD & GARDEN" CROPS. [CH.
but so small and gauzy that they may be readily over-
looked. One of these small and inconceivably thin
wings is shown enlarged to 20 diameters at Fig. 143,
•20
FIG. 143.
Green fly, from potato plant, natural size, and wing with spores or conidia
of the potato fungus, Peronospora infestans, Mont., at A.
Enlarged 20 diameters.
and on the wing at A, engraved to the same scale, are a
number of conidia or spores of the potato fungus. Small
as these germs are as seen on the greatly enlarged wing,
xxxvin.] CONCLUSION.
yet each atom under favourable circumstances encloses no
less than eight other atoms, each furnished with two
vibrating hairs, and endowed with the power of sailing
rapidly about in any non-corrosive film of moisture.
It need hardly be said that various insects and flies,
both large and small, commonly eat or imbibe fungus
spores. The spores are not only to be seen dusted over
the wings or sticking amongst the hairs of the legs, but
they are quite as commonly seen inside the insects as
out ; this is especially well seen in such small transparent
insects as plant-lice or aphides. The spores are carried
about with the juices inside the bodies of the insects, and
may not only be found in the body, but inside the limbs,
and even within the almost invisible antennae or horns.
The spores of various fungi not only stick to the bodies
of insects, but they germinate upon them and produce
mycelium outside, and sometimes, inside their bodies.
Mr. G. B. Buckton, F.R.S., in commenting upon our
observations on this subject in his Monograph of British
Aphides, says the facts need cause no surprise.
Small as some of these organisms or parts of organisms
may be, it must not be assumed that we are acquainted
with the smallest objects of nature. On the contrary,
every new and true observation about minute things indi-
cates that what we already know regarding small things
is as nothing when compared with what is not known,
and what at present we seem to have but little prospect
of knowing. The most perfect and powerful telescopes
cannot resolve the more distant nebulae into stars,
neither can the most perfect microscopes display to our
sight numerous atoms which are believed to exist, but
which cannot be seen. Persons possessed of strong
vision can often see, both with the telescope and micro-
scope, objects that are invisible to persons of ordinary
sight It would be very rash, therefore, for any observer
to say that certain objects or characters do not exist
simply because that observer cannot see them. Neither
340 DISEASES OF FIELD & GARDEN CROPS. [CH.
is it always wise to say too positively that doubtful
characteristics are certainly present ; it is safer under
critical circumstances to say the characters appear to be
present or non-present.
The cilia or vibrating hairs of the zoospores of the
potato fungus are so excessively attenuated that, when the
highest magnifying powers are used, and with the cilia close
to the object-glass, it commonly happens that both hairs
cannot be seen at the same time : an alteration of the focus,
small beyond description, is necessary to see first one, then
the other. When a zoospore becomes quiescent and ger-
minates, the cilia vanish. They either dissolve or break
up into the finest dust — dust so small that no figures can
express the minuteness of the particles. Now it is well
known that all parts of the potato fungus are so potent
with life that every visible atom will grow and reproduce
the fungus. It is quite possible, then, that, just as every
atom of a mycelial thread of this fungus will continue its
growth to the perfect form, so every atom of a broken up
flagellum — perfectly invisible to the eyes even when the
highest powers of the microscope are used — may be
capable of carrying the poison and at length reproducing
the perfect form of the fungus in the potato plant.
We think it would be well if all agriculturists would
set apart a small portion of each farm or garden for ex-
perimental purposes, each farmer taking a personal,
practical, and scientific interest in his own special crops.
Seeds of all sorts should be selected from the healthiest
parents. Indiscriminate seed planting should never be
practised. By constantly selecting seed from plants free
from disease, hereditary disease might at length exhaust
itself and be extinguished. We think it impossible to
over-estimate the importance of the fact of the hereditary
nature of disease in plants and animals. Mr. Charles
Darwin, writing in his Animals and Plants under Do-
mestication, vol. ii. p. 7, says : " Unfortunately it matters
XXXVIIL] CONCLUSION. 341
not, as far as inheritance is concerned, how injurious a
quality or structure may be if compatible with. life. No
one can read the many treatises on hereditary disease
and doubt this. ... A long catalogue could be given of
all sorts of inherited malformations, and of predisposition
to various diseases." Under "Cataract of the Eye" Mr.
Darwin writes at p. 9 : " When cataract affects several
members of a "family in the same generation, it is often
seen to commence at about the same age in each — e.g., in
one family several infants or young persons may suffer
from it, in another several persons of middle age." This
latter observation has a direct bearing on the hereditary
diseases of plants which notoriously appear at certain
stages of the plant's growth. Under " The Horse " Mr.
Darwin quotes Youatt on p. 9, who writes : " There is
scarcely a malady to which the horse is subject which is
not hereditary." At p. 11 : "Andrew Knight, from his
own experience, asserts that disease is hereditary in
plants, and this assertion is confirmed by Lindley." And
again : " Seeing how hereditary evil qualities are, it is
fortunate that good health, vigour, and longevity are
equally inherited. ... As to the inheritance of vigour
and endurance the English racehorse offers an excellent
instance. Eclipse begot 334 and King Herod 497
winners."
Facts like the above should serve as key-notes, and
strongly impress all practical agriculturists. Every
diseased plant or seed should be mercilessly struck out
and destroyed. No owner of herds and flocks would
allow badly diseased animals to breed, and, in the same
way, no agriculturist should take his seed from plants
notoriously infected. By a constant selection of seeds
from plants — the freest from disease — at length races
might be obtained almost entirely free from disease. If
it should some day be proved that disease does not exist
in seeds, it will not be denied that certain plants inherit
a strong tendency to become diseased.
342 DISEASES OF FIELD & GARDEN CROPS. [CH.
Sometimes, after several healthy generations of plants,
the progeny may revert back to disease in an analogous
way with the many similar instances so well known
amongst men and other animals. The disease in these
cases is sometimes derived from some remote and perhaps
forgotten ancestor.
A rotation of crops is of high importance as regards
disease, for a fungus that destroys turnips or cabbages
will probably not injure corn, and neither will injure
clover. As a rule any given destructive fungus keeps to
one Natural Order of plants, often to one genus, some-
times to one species. Any fungus capable of invading
plants belonging to several Natural Orders is an exception
to the rule.
In writing these concluding words We are strongly
impressed by the fact of how little has really been described.
The mere margin of each subject has been barely ap-
proached, and the anatomy and physiology belonging to
each disease only glanced at. The subjects discussed have
only been presented in bare outline, and many of great im-
portance have not been mentioned at all — as the diseases of
beans, beet, cucumbers, mint, hops, etc. A large subject
awaits description and illustration in the nature of canker
and the diseases of our fruit crops, our grapes, peaches,
nectarines, apples, pears, gooseberries, strawberries, figs,
melons, etc. Another of equal importance presents itself
in the diseases of our timber trees, our firs, pines, larches,
oaks, elms, and latterly, yews.
In closing, we advise all students of nature to think as
well as observe, for a man may be a good observer and
not a good thinker, and a good thinker may be but a poor
observer. Some observers, by always applying themselves
to the elucidation of minute things, have apparently made
themselves mentally incapable of broad generalisation and
the understanding of great ones.
The greatest possible caution is necessary in making
deductions from observed facts. So long as an observer
xxxviii.] CONCLUSION. 343
keeps to a record of facts, lie is doing useful work, and
work that cannot be questioned ; but as soon as theories,
hypotheses, and deductions are introduced, an element of
uncertainty creeps in. The mere superficial appearances
presented by Nature are seldom to be depended upon.
They are often deceptive, and exhibit a tendency to lead
an observer in the wrong direction. We may refer as
examples to the structure of the blue mould fungi of our
provisions, named Eurotium repens and E. (Aspergillus)
glaucus, Lk. In these fungi sexual organs have been
described and illustrated by Professor De Bary, but
rejected as such by M. Ph. Van Tieghem (Bulletin de la
Socie'ttf Botanique de France, p. 96, 1877) ; to the supposed
sexual condition described and illustrated by Tulasne and
De Bary under Erysiphe and Peziza; both the latter
observations greatly need confirmation ; and to the cases
of erroneous interpretation mentioned elsewhere in this
volume. With a full knowledge of the deceptive super-
ficial appearances frequently presented by natural objects,
Mr. Charles Darwin once said that Nature "will tell
you a direct lie if she can " (Trans. Essex Field Club, vol.
iii. p. 67).
It is not wise to become an unqualified believer or dis-
believer in any hypothetical views. Our knowledge of
Nature is at the best extremely imperfect, and the very
little we know of her and her ways is as nothing when
compared with what we do not know.
INDEX.
AcMllea Ptarmica, L., 183
Achlya racemosa, Hild., 296
Acosta, M. Joachim, 277
jEcidiospores, 186
JEcidium Allii, Grev., 44
asperifolii, Pers., 141, 143, 144,
145, 181
bellidis, B.C., 179
Berberidis, Pers., 159, 160,
161, 163, 165, 166, 167, 176
E, development of, 162
nice, Lenz., 206
uperans, Vize., 197
Euphorbice - sylvaticce, B.C..
190
magelhanicum, B., 197
meaning, 143
tragopogonis, Pers., 175, 207
Tussilaginis, P., 179
violce, Schum., 197
JEgilops ovata, L., 158
JEgopodium Podagraria, L. , 239
Agaricus nauseosus, Fr. , 66
ostreatus, Jacq., 66
tuberosus, Bull., 21
Agrostis, 113, 215, 253
alba, L., 158
pulchella, 73
vulgaris, With., 158
Aira ccespitosa, L., 158, 214, 262
flexuosa, L., 214
Alliaria officinalis, B.C., 85
Allium acutangulum, Schrad., 43
Cepa, L., 43, 49
fistulosum, L., 49
flavum, L., 43
oleraceum, L., 195, 196
ophioscorodon, Bon. , 43
palustre, Pourr., 43
Porrum, L., 43
rotundum, L., 43
sativum, L., 43
schcenoprasum, L., 38, 43
Allium stellatum, Gawl., 43
ursinum, L., 44, 195, 196
Alopecurus fulvus, Sm., 158
pratensis, L., 158, 214
Amelanchier, 204
American onion smut, 54
Amoeba, meaning, 8
Anchusa arvensis, Bieb., 146
officinalis, L., 141, 146
Anda, Herr, 22
Andropogon hirtus, L., 262
Anemone nemorosa, L., 195
Angelica sylvestris, L., 239
Aniseed, 244
Antheridium, 297
meaning, 91
Anthina flammea, Fr., 57
Anthocercis viscosa, R.Br., 276
Anthoxanthum odoratum, L., 214
Anthriscus cerefolium, HoflFm., 244
Aphides, 96
Aphis, 339
Apple, 204
Aquinas, 317
Arabis perfoliata, L., 85
Arrhenatherum avenaceum, Beauv.,
214, 262
elatior, L., 142
Artotrogus hydnosporus, Mont., 84,
100
meaning, 295
Artotrogus in cruciferous plants,
313
Ascobolus, 28
trifolii, Riv., 7
Ascus, meaning, 27
Aspergillus glaucus, Lk., 343
Australia, 180, 181, 182, 184, 189
Avena elatior, L., 158
fatua, L., 158
flavescens, L., 142, 158, 262
pratensis, L., 158
pubescens, L., 262
sativa, L., 158, 262
Awn, meaning, 107
346
INDEX.
Bacteria, 225
Baker, J. G., F.E.S., 281, 315
Baldmoney, 244
Balfe, Ambrose, 15
Banks, Sir Joseph, 172, 173
Barberry blight, 159
its possible connection with
corn mildew, 169
Barley, 215, 253, 262
grass, 262
Barren brome grass, 136
Bary, Professor de, 23, 84, 95, 175,
176, 186, 296
Barya aurantiaca, P., 237
Bastian, H. C., 109
Beagle, voyage of, 317
Berberidacece, 159
Berberis glauca, D.C., 196
Uicifolia, Forst., 197
Berkeley, Kev. M. J., 24, 31, 35, 55,
87, 175
Bishopweed, 239
Bitter-sweet, 275
Blanchard, M., 318
Blight of barberries, 159
of borage, 143
of straw, 69, 71
Blytt, Professor, 23
Bonninghausen, 174, 178, 185
Borage, blight of, 143
Boraginacece, 159
Borrago officinalis, Tour., 146
Botrytis, meaning, 82
crustosa, Fr., 241
terrestris, Pers., 330, 335
Boussingault, M., 277
Brachypodium ciliatum, P.B., 262
Brassica campestris, L., 85
oleracea, L, 85
Brefeld, Dr. Oscar, 250
British Museum (Nat. Hist.), 307
Bromus asper, L. , 142
mollis, L., 142, 158
sterilis, L., 136
tectorum, L., 158
Broome, C. Edmund, M.A., 84, 296,
300
Brown, Robert, 330
Bryum Jiornum, Sw., 63, 64
Buckman, Professor James, 76, 125
Buckton, G. B., F.R.S., 96, 193, 339
Bunt, 170, 171, 184, 245
meaning, 245
remedies for, 252
Burnet saxifrage, 244
Bussey Institution, bulletin of, 321,
328
Butterworth, Mr., 330
72
C
CABBAGE, 85
Cabbages, club -root in, 94, 97, 98,
100, 101
putrefactive mildew of, 80, 81,
83, 84
white rust of, 86, 88, 91
i Calamagrostis Epigejos,~Roih., 142,158
I Caldcleugh, Alexander, 317
Camelina sativa, Cranz., 85
Cancer, 198
Candytuft, 236
Capnodium, 57
Footii, B. and D., 195
Capsella Bursa-pastoris, D.C., 85, 86
impatiens, L., 85
Cardamine Mrsuia, L. , 85
Carduus arvensis, Curt., 273
Carroll, Thomas, 23
Carrathers, Wm., F.R.S., 110, 172,
180, 321, 330
Caspary, Dr., 296
Cathcart, Earl, 315
Catstail-grass, 214
Cedar apples of United States, 203
Cerinthe alpina, Kit., 146
minor, L., 146
Chcerophyllum sylvestre, L. , 244
Cheiranthus Cheiri, L., 85, 236
Chiswick, garden at, 294
Chives, disease of, 38
Chlorochytrium Lemnce, Colin., 334
Cholera, 252
Chrysomyxa Ledi, 193
Cichorium Endivia, L., 273
Cilia, 340
meaning, 95
Cladosporium, 6
Claviceps, 60
microcepTiala, Tul., 238
purpurea, Tul., 214, 219
purpurea, Tul., var. Wilsoni.
W.Sm., 233
meaning, 219
structure, 220, 221
Clog wheat, 229
Clover dodder, 115
mildew, 6
seeds, 117
seeds germinating, 118, 118
sickness, 6
Club-root, 94, 97, 98, 100, 101
Cobbolcl, Dr. T. S., 189
Cocksfoot grass, 214
Ccema Rubigo, L., 138
Colorado beetle, 328
INDEX.
347
Comfrey, tuberous, 144
Conidiophore, meaning, 9
Conidium, meaning, 8
Conium maculatum, L., 244
Consortism, 241
Cooke, Dr. M. C., M.A., 176, 181,
183, 207
Coral root, 85
Cordiceps, 65
meaning, 60
Corn mildew, 135, 147
its possible connection -with
barberry blight, 169
spring rust and mildew of, 135
Cornu, Dr. Max, 302
Cow-parsnip, 239, 244
Cracking of potatoes, 35
tomentosa, L., 204
Creeping soft grass, 136, 214
Crimson clover, 7
Cupressus, 205
Curl disease of potatoes, 277
Curtis, J., 97
Cuscuta, meaning, 115
Trifolii, Bab.,* 115, 116, 117,
118, 119, 120, 122, 124
Cynodon Dactylon, L., 262
Cynoglossum officinale, L., 146
Cystopus candidus, Lev., 85, 86, 88,
91, "
Dactylis glomerata, L., 158, 214
Dactylium tenuissimum, B. , 31
Darnel grass, 215, 262
Darwin, Charles, 317, 330, 340
Decaisne, 251
Dentaria bulUfera, L., 85
heptaphyllos, Clus., 85
Devaine, 110
Dianthus, 182, 185
Diarrhcea, 198
Diatrype, 213
Digraphis arundinacea, Trin., 44
Disease, hereditary, 341
mode of prevention, 341
new, of grass, 55
Disinfecting potatoes, 326
Dodder, anatomical connection with
stem of clover, 122, 124
host, plants of, 117
meaning, 115
of clover, 115
seed of, 117
seeds germinating, 118, 119
DotMdeatrifolii,Fr,, 7
Downy oat-grass, 262
Drdba verna, L. , 85
Dry rot of potatoes, 277
Duchamel, M., 227
Duckweed, 334
Du Port, Rev. Canon, 231
EAR- COCKLE in wheat, oats, and
rye, 105, 106, 108, 109, 110, 112
Earthing-up potatoes, 321
Echium vulgare, L., 146
Eczema, 198
Eels of stale paste, 109
Elaphomyces muricatus, Vitt., 60, 61
meaning, 64
Elymus arenarius, L., 158, 215
giganteus, Vahl., 215
glaucifolius, L., 158
Empusa muscce, Cohn., 66, 194
Endophyllum, 188
Entozoa, 187, 188, 189
Equisetum, 185
arvense, L., 301
Ergot, 214
effects of, 227
meaning, 219
of rye, 216
remedy against, 231
structure of, 217
Ergotcetia dbortifadens, Qk., 225
Erysiphe, 79, 343
communis, Schl., 7
graminis, D.C., 7, 126, 127,
129, 130, 131, 132, 133, 266
Martii, Link., 7, 263
meaning, 126
Pisi, Grev., 266
Essex Field Club, Transactions of,
343
Eurotium repens, 343
FAIRY RINGS, 292
Farlow, Prof. W. G., 203, 280
Fescue-grass, 262
Festuca elatior, L., 113, 142, 215
gigantea, Vill., 158
ovlna, L., 55, 56, 57
pratensis, Huds., 251, 262
spectabilis, Jan., 158
tenella, Willd., 158
Finger-and-toe, 94
Floating sweet-grass, 215
Fossil fungi, 330
Foxtail grass, 214
348
INDEX.
Fries, Elias, 178
Fumagine, 195
Fumago, 195
Fusarium graminearum, Sch., 209
Fitsicladium, 183
Fusisporium, meaning, 30
atro-virens, B., 49
culmorum, W.Sm., 208, 210
nordei, W.Sra., 208, 210
inosculans, B., 251
insidiosum, B., 73
Lolii, W.Sin., 208, 212
mucophytum, W.Sm., 213
oUusum, Ck., 213
roseolum, Steph., 31
Solani, Mart., 30, 32, 33, 208,
277
G
GALLS, 109
Gardeners' Chronicle, 175, 176, 183
Garlic, 43
mustard, 85
Gasteromycetes, 295
Geologists' Association, 331
Glume, meaning, 105
Glyceria fluitans, R.Br., 215, 233
Goritti, 318
Gout, 198
Goutweed, 239
Graminece, 159
Grass mildew, 126
new disease of, 55
Green fly, 338
Grevillea, 238
Groundsel, 272, 273
Grove, W. B., M.A., 197
Gymnosporangia of United States,
203
Gymnosporangium Mseptum. Ellis,
204
Juniperi, Lie., 206
macropus, Lk., 204
HAIR-GRASS, 262
Hairy bittercress, 85
Hallier, Professor, 252
Hansen, Dr. Chr., 209
Hardingham, 170
Harkness, Dr. H. W., 134
Haustoria, 267
meaning, 80
Hawthorn, 204
Hemlock, 244
Henbane, 275
Hen slow, Prof. J. S., 171
Heradeum Sppndylium, L., 239
Hereditary diseases, 341
Hernia, 94
Hetercecia, 189
Hetercecism, 205
in Peronospora, 328
in Puccinia, 141, 168, 169
in Tilletia, 250
in Ustilago, 250
Hogweed, 239, 244
JIolcus lanatus, L., 136, 142, 158, 214
mollis, L., 136, 142, 214
Honey Guide, Great, 65
Hopkins, Matthew, 174
I Hordeum distichum, L., 142, 158,
215, 253, 262
murinum, L.. 136, 142. 158,
262
secalinum, Trin., 142
sylvaticum, Huds., 158 *
vulgare, L., 142, 158, 262
Horticultural Society, Journal of
Royal, 182
Hydnum, 295
Hyoscyamus niger, L., 275, 276
Hypericum, 268
Ileris, 236
Indicator major, Steph. , 65
Inflammation, 198
Isaria, meaning, 55
Juciformis, B., 55
JAMIESON, Prof., 103
Japan, 182
Jensen, J. L., 174, 279, 319
Juniperus communis, L., 207
Jussieu, 171
KEW, 312, 317
Knight, T. A., 171, 281, 319
Krunitz's Encyclopedia, 172
Lfibrella Ptarmica, D., 183
iMctarius controversus, P., 178
Lactuca altissima, M.B., 273
sativa, L., 273
Lang, Dr. Jeffrey, 320, 327
Lapsana communis, L., 273
Lathyms macrorrhizus, Wimm., 265
Leek, 43, 49
Le Maout, 251
INDEX.
349
Lepidodendron, 330, 333
Lettuce mildew, 269
Leveille, 134
Lindley, Prof. John, 125
Unnean Society, Journal of, 179
Lithospermum arvense, L., 146
Lodicules, meaning, 107
Lolium, 253
perenne, L., 158, 212, 215, 262
temulentum, L., 142, 215, 262
IjOtus corniculatus, L., 13
Lucern, 7
Lychnis, 260
Lycopersicum esculentum, Mill., 275
Lycopsis arvensis, L., 141
Lyme-grass, 215
M
Mdhonia, 159
Aquifolium, Lind., 166, 167,
184, 185
M'Cullough, Dr., 178
Mangels, club-root in, 94, 97, 98,
100, 101
Manures, mineral, 198
Martius, 35
Massachusetts, Province Law, 174
Masters, Dr. M. T., F.R.S., 262
Mat-grass, 214
Matthews, Chas. Geo., 209
Meadow fescue-grass, 215
soft-grass, 214
Medicago sativa, L., 7, 9
Medlar, 204
Melica, 262
Metcecia, 189
Meum Athamanticum, Jacq., 244
Microcera coccophUu, Desm., 66
Mildew, and spring rust of corn,
135
corn, 147
of grass, 126
putrefactive, of cabbages, 80,
81, 83, 84
putrefactive, of turnips, 80,
81, 83, 84
surface of turnips, 75, 77, 78
Millet, 262
Mnium hornwm, Heclw., 63, 64
Molinia ccerulea, Moench., 158
Molluscs, 193
Montagne, Dr., 84, 294, 295
Morchella Smithiana, Ck., 178
Morren, Dr., 277
Mouillefert, M., 13
Moulding, protective, 319
up potatoes, 321
Mould of onions, 51
Mucor, meaning, 51
mucedo, L., 51
subtilissimus, B., 21, 51
Mullein, 75
Murray, George, F.L.S., 321
Mushroom spawn, 291
Mycelium, perennial, 290
Mycetozoa, meaning, 95
Mycologia Scotica, 202, 207
Mylitta, 19
Myxomycetes, meaning, 94
Myzocytium, Sch., 302
N
Nardus stricta, L., 214
Narrow-leaved bittercress, 85
Nematode, meaning, 109
Neslia paniculata, Des., 85
Neuralgia, 198
New Zealand, 180
Nipplewort, 273
Noel, M., 229
Nonneapulla, D.C., 146
OAT-GRASS, 214
Oats, 262
Oersted, Prof. A. S., 203, 204
Ohrond, M., 318
Oidium dbortifatiens, B. and Br., 225
Balsamii, Mont., 75, 77, 78, 80
meaning, 75
monilioides, Lk., 127
porriginis, Mont., 66
Tuckeri, B., 76
Onion, 43, 45, 49
fly, 50
mould, 51
smut, 54
Onygena apus, B. and Br., 66
equina, Pers., 66
Oogonium, meaning, 47
Oosphere, 90
meaning, 297
Oospores, 83, 91
formation of, 298
of Peronospora inf-
Mont., germination, 308
Orolus, 265
Oryza sativa, L., 215, 262
Osmundites Dowkeri, Carr., 330
Ovularia, 286
splicer o idea, 14
Ozonium, 72
350
INDEX.
Pachymacocos, Fr., 19
Pale, meaning, 106
Paraphyses, meaning, 28
ofPuccinia, 139
Parsnips, mildew of, 239
Pea mildew, 266
remedy against, 268
Pea mould, 263
remedy against, 264
Peard, Dr. W., LL.B., 281, 312
Penny cress, 85
Peppercorn in wheat, oats, and rye,
105, 106, 108, 109, 110, 112
Peridium, meaning, 162
Periola tomentosa, Fr. , 31
Perithecium, meaning, 61, 130
Peronospora exigua, W.Sm., 12, 337
ganglioniformis, B., 269
grisea, Ung., 12
Hyoscyami, P., 276
infestans, Mont., 15, 23, 275, 336
meaning, 7
nivea, Ung., 239
parasitica, Pers., 75, 80, 81,
83, 84, 100, 101
Schleideniana, Ung., 45, 46, 48,
337
trifoliorum, D.By., 6, 10, 13
umbelliferarum, Gasp., 239
vicice, Berk., 263
Peronosporites antiquarius, W.Sm.,
331
Petroselinum sativum, Hoff., 244
Petunia, 276
Peziza, 343
ciborioides, Fr., 6
Fuckeliana, D.By., 20
meaning, 24
postuma, B. and W., 15, 25
sclerotiorum, Lib., 23
tuberosa, Bull., 23, 195
Phacidium trifolii, Bond., 7
medicaginis, Desm., 6
Plialaris arundinacea, 44, 158, 214
Phillipar, 170, 172, 250
Phillips, Wm., F.L.S., 38
Phleum pratense, L., 158, 214
Phthisis, 198
Phytopathology, 252
Phytophthora, 286
Pies, of potato growers, 314
Pirn, Greenwood, M.A., 67
Pimpinella Anisum, 244
Saxifraga, L., 244
Pisum, 265
Plasmodiophora, meaning, 94
Brassicce, 94
Plasmodium, meaning, 94
Pleospom, 6
Plowright, C.B., M.R.C.S., 167, 171,
173, 174, 319
Poa annua, L., 158, 179, 184
aquatica, L., 260
fluitans, Scop., 260
nemoralis, L., 158
pratensis, L., 158
trivialis, L., 237
Podisoma Juniperi-Sabince, Fr., 202
Polyactis dnerea, B., 20
Polycystis, 35
Polygonum, 253, 262
Polyporus, 21
Polythrincium trifolii, Kze., 7
Potato disease, 275
prevention of, 314
scab, 35
Potatoes, " Champions," 15
cracking, 35
culture of, 15-16
new disease, 15-17
" Protestants," 15
Prevost, 288
Protective moulding, 319
Protomyces, meaning, 241
macrosporus, Ung., 241, 242
Protomy cites protogenes, W.Sm., 333,
334
Protoplasm, meaning, 90
Protozoa, 94
Pucdnia, meaning, 38
cegra, Grove, 197
allii, Rud., 43, 195
apii, Corda, 183, 244
graminis, Pers., 147, 153, 154,
155, 156
heraclei, Grev., 244
lychnidearum, Lk., 182
malvacearum, Mont., 38, 51
mixta, PI., 38, 40, 41, 42, 51
obscura, Sch., 193
Poarum, K, 179
Rubigo-vera, D.C., 135, 139,
140, 141
sparsa, Ck., 207
sessilis, Schum., 44
straminis, PL, 142
striceformis, West., 142
tragopogi (Pers.), PL, 207
tragopogonis, Corda, 175, 207
violai, D.C., 197
Schum., 197
violarum, Lk., 197
Pulmonaria officinalis, L., 146
tuberosa, Schrk., 146
Pupce, 59
Purple clover, 7
INDEX.
351
Purples in wheat, oats, and rye, 105,
106, 108, 109, 110, 112
Pycnidiiim, meaning, 134
Pyracantha, 183
Pyrus, 204
Pythium equiseti, Sdbk., 185, 301
incertum, Ry., 302
megalacanthum, D.By., 303
micracanthum, D.By., 303
proliferum, D.By., 303
vexans, D.By., 303
QUINCE, 206
B
Ramularia, 286
sphceroidea, 14
Rayer, Dr., 294
Reed canary-grass, 214
Resting-spores, 83
Revets, 229
Rheumatic fever, 198
Rhizomorpha, 72
Rhizosporium solani, Rab., 35
Rhopalosiphwndianthi, Sch., 195, 337
Rice, 215, 262
Robin, Charles, 60
Rocambole, 43
Rcestelia, 188
aurantiaca, Pk., 206
cancellata., Reb., 202, 203
cornuta, Tul., 206
lacerata, Tul., 206
Jiosacece, 76
Rot of potatoes, 277, 280
Royal Agricultural Society, Journal,
172, 175
Royal Horticultural Society, Journal,
175
Royal Society, Proceedings, 167, 185
Rum, Island of, 278
Rumex Acetosa, L., 262
oUusifolius, L., 253
Rust, spring of corn, 135
summer, 147
white of cabbages, 86, 88, 91
Rye, 214, 262
grass, 215, 262
S
SADEBECK, Dr., 301
Saprolegnia, 237
asterophora, D.By., 296
fercas, Kutz., 66, 194, 331
meaning, 67
philomukes, W.Sm., 67, 68
Savin, 202
Saxifrage, burnet, 244
Scab, of potatoes, 35
Scallions, 40
ScMzanthus Grahami, Gill, 276
Schceler, 174
Sclater, Dr., F.R.S., 317
Sclerotium, 20, 21, 51, 217
Cepce, 51
cepcevorum, B., 21, 51
clavus, D.C., 218
complanatum, Tode, 20
durum, P., 20
echinatum, 20
fungorum, P., 21
meaning, 18
scutellatum, A. and 8., 20
stipitalum, Fr., 21
varium, P., 24
Scrofula, 198
Scrophulariacece, 76, 276
Secale cereale, Walld., 142, 158, 214,
262
Senecio vulgaris, L., 272, 273
Septum, meaning, 13
Serrafalcus secalinus, B., 142
Setaria italica, Beauv., 262
Sharpe and Co., 117
Shepherd's-purse, 85, 86
Shingles, 198
Silene, 260
Smee, Alfred, 206
Smooth meadow-grass, 214
Smut of corn, 254
remedies against, 261
of onions, 54
Solanacece, 276
Solanum cardiopliyllum, Lind., 275,
315
Collinum, Dun., 315
Colombianum, Dun., 315
Commersoni, Dun., 315
demissum, L., 275, 315
Dulcamara, L., 275
etuberosum, Lind., 315
Fendleri, A. Gray, 315
Fernandezianum, Phill., 315
immite, Dun., 315
Jamesii, Ton-., 315
Maglia, Sch., 315
Ohrondii, Carr., 315
oxycarjmm, Sch. , 315
squamulosum, M. and G., 315
stoloniferum, Sch., 315
suaveolens, K. and B., 315
tuberosum, L., 281,315
utile, Klot, 315
Valenzuelce, Pal., 315
verrucosum, Sch., 315
Sorghum vulgare, P., 262
352
INDEX.
Sonchus arvensis, L., 273
Sorus, meaning, 40
Spermatia, 188
meaning, 164
Spermozdia davus, Fr., 218
Kpermogonia, 160, 164, 165
development of, 162, 164
Spermogonium, meaning, 164, 186,
188, 194, 205
Sphacelia, 238
meaning, 224
structure, 225
Sphceria, 65
herbarum, Pers., 6
Spircea Ulmaria, L., 268
Spore, meaning, 8
Sporangium, meaning, 51
Sporidium, meaning, 27
Spring rust and mildew, 135
Stilbum, 60
St. John's wort, 268
Strawberries, 75
Straw blight, 69
Strigula Babingtonii, B., 195
Stroma, meaning, 61
Stuart, Duncan, 278
Stylospore, meaning, 134
Stylospores, 226
Summer rust, 147
Button and Sons, 117
Symphytum officinale, L., 146
tuberosum, 141, 144, 146
TALL FESCUE-GRASS, 215
Tanner, Prof. Henry, 231
Tare or vetch mould, 263
Teleutospore, meaning, 41
Teleutos pores, development of, 192
Thecce, meaning, 61
Thecaphora, 184
Thlapsi arvense, L. , 85
Thuya, 57
Tieghem, Ph. Van., 341
Tilletia bullata, Fl., 253
caries, Tul., 35, 54, 184, 245,
248
Icevis, Kuehn., 253
Lolii, Awd., 253
Tillet, Matthieu, 245
Tomato, 275
Torrubia, meaning, 60
militaris, Tul., 65
ophioglossoides, Tul., 60,'. 61, 65
sphecocephala, K.I., 60
Tower mustard, 85
Tariuon, garden at, 171
Trichobasis glumarum, Lev., 138
Rubigo-vera, Lev., 138
Trifolium alpestre, L., 7
incarnatum, L., 7
medium, L., 7
Triticiim canimim, Huds., 158
repens, Vill., 158, 215
sativum, L., 215, 253
turgidum, L., 262
vulgar*, Vill., 142, 158, 253 262
Truffle, 60
Tubercinia, meaning, 35
scabies, B., 35, 54
Tuberous comfrey, 144
Tuckahoo, 19
Tulasne, 296
Turfy hair-grass, 214
Turnip, 85
Turnips, club-root in, 94, 97, 98, 100,
101
putrefactive mildew of, 80, 81,
83, 84
surface mildew of, 75, 77, 78
Tylenchus, meaning, 109
devastatrlx, 6
Havensteinii, 6
tritici, Bast., 105, 110, 112
vital tenacity in, 112
Typhula, 20
Umbelliferce, 268
Uredo, meaning, 54
alliorum, D.C., 43
a Wall., 244
'.a, Baeur, 247
ris, Pers., 147, 148, 149,
150, 151
porri, Sow., 43
RuUgo, B.C., 138
Rubigo-vera, B.C., 135, 136,
137, 138
Urocystis, 35
cepulcB, Tar., 54
colchici, Tul., 54
meaning, 54
occulta, Pre., 252
violce, B. and Br., 54
Uromyces alliorum, D.C., 43
appendiculatus, Lev., 7, 190
Ustilaginei, 250
Ustilago antherarum, Fr., 260
Candollei, Tul., 262
carbo, Tul., 35, 54, 254, 259,
337
grandis, Tul., 252
Kuhniana, Wolff, 262
INDEX.
353
Ustilago longissima, Tul., 260
meaning, 254
VACCINATION, 198
Valvatidce, 193
Verbascum montanum, Schrad., 75
nigrum, L., 75
Vernal grass, 214
Vetch, bush, 265
common, 265
slender, 265
Viciasativa, L., 265
sepium, L., 265
tetrasperma, Mosnch., 265
Vinegar eels, 109
Vize, Bev. J. E., 47, 281, 299, 307
W
WALDHEIM, A. F. DE, 250
Wall-barley, 136
Wallflower, 236
Wasp, with fungus growths, 59
Waved hair-grass, 214
Webb, Edward, and Sons, 117
Wheat, 215, 253, 262
grass, 215
Whilborough, man at, 320
White-rust fungus, 85
White-rust of cabbages, 86, 88, 91
Whitlow grass, 85
Williams, B. 8., 184
Williamson, Prof. W. C., F.R.S., 333
Wilson, A. Stephen, 22, 300
Wilson's Claviceps on ergot, 233
Witchcraft, 170
Witchfinder, 174
Woronin, M., 94
Wright, Dr. E. P., A.M., F.L.S., 67
YEAST, 335
Yellow oat-grass, 262
Young, J. T., F.G.S., 331
ZEALAND, NEW, 180
Zoosporangia, meaning, 67
Zoospore, meaning, 88
Zygospores, 243
Zygozoospores, 334
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