History of Ecological Sciences, Part 44

Transcription

Click here for all previous articles in the History of the Ecological
Sciences series by F. N. Egerton
History of Ecological Sciences, Part 44: Phytopathology during the 1800s
Frank N. Egerton
Department of History, University of Wisconsin-Parkside, Kenosha Wisconsin 53141
E-mail: egerton @uwp.edu
Every ecologist knows the basic history of evolutionary theory—Lamarck’s inability to sway
many naturalists with his flawed arguments, and Darwin’s only gradual success with his masterful
presentation. Much less well known is the struggle to establish the germ theory of disease, with many
defenders and skeptics for well over a century before it gained acceptance. Although the germ theory
of disease is narrower in scope than evolutionary theory, its practical importance is at least as great, and
therefore its history merits greater familiarity. We saw in part 29 (Egerton 2008) that both mycology
and phytopathology made notable advances during the 1700s—with small steps toward a germ theory—
though these advances were not widely known or appreciated. Researchers during the 1800s built upon
the work of their predecessors. The science of phytopathology began with investigations of fungi, the
earliest known culprits, and fungi were already being accused during the 1700s, though there was a
hung jury rather than a clear verdict of guilt (Egerton 2008). If guilt was to be established, it was
important for botanists to have a standard system to classify what they knew and make clear what was
a new discovery when one occurred. Mycology and phytopathology had to advance together. Although
John Needham (1713–1781), whom we met in part 24 (Egerton 2007:147), discovered a “worm” (a
nematode now named Anguina tritici) in wheat galls (Needham 1743:640–641), D. J. Raski (1959:386)
dates the beginning of nematology to the Histoire naturelle des helminthes ou vers intestinaux (1845)
by Felix Dujardin (1801–1860). In 1854–1856, Dujardin’s fellow countryman, Casimir-Joseph Davaine
(1812–1882) studied the nematode that causes seed-gall disease in wheat, and in 1868 Davaine also
discovered a bacterium that caused a plant disease. Not until the late 1800s were there evidences of a
virus as a cause of plant disease. Nature is one, but scientists partition it into sciences they can master.
Although phytopathology and animal parasitology developed as separate sciences, their concerns
overlap in cases of fungal diseases of animals, including humans, and nematode diseases of plants;
and both phytopathology and parasitology overlap with bacteriology and virology concerning bacterial
and viral diseases. The history of nematode parasitism of plants was outside the scope of both Large’s
(1940) and Ainsworth’s (1981) histories of phytopathology, but nematodes are discussed in textbooks
on phytopathology (Heald 1926:831–851, Walker 1969:533–548, Agrios 2005:826–874). With many
simultaneous developments occurring, it seems best to discuss studies of fungi first, nematodes and
bacteria second and third, and viruses last, in Europe, then in North America.
Europe
The botanist who began to put the fungal house in order was Christiaan Hendrik Persoon (ca.1761–
1836), of Dutch heritage, from Cape Town, South Africa (Donk 1974, Ainsworth 1976:255–258, MagninGonze 2004:170–171). In 1775 he went to Germany, where he received his botanical education, and
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Fig. 1. (a) Christiaan Hendrik Persoon in 1796. (b) Elias Magnus Fries. Ainsworth 1976:256, 260.
in 1802 he moved to Paris, where he lived alone, in poverty. He corresponded with many botanists and
exchanged specimens with them, exemplified by his correspondence with James E. Smith (Ramsbottom
1934). In 1828 he gave his botanical collections to the Dutch government in exchange for a pension, and
when he died he left his remaining collections and his library to the Dutch government.
Persoon published his first article on fungi in 1793 (Schmid 1933), introduced his classification
system in 1794, and produced his Synopsis methodica fungorum in 1801, which became the foundation
of modern mycological systematics. Geoffrey Ainsworth (1976:258) found that “almost all the hundred
genera and subgenera he recognized are universally accepted genera of today.” Persoon brought together
the rusts and smuts in the Dermatocarpi, and he named Fontana’s small parasitic plants, brown and black
rust on wheat, Uredo linearis and Puccinia graminis (Fig. 2). However, Persoon accepted the prevailing
belief that these species could arise from abnormal sap or tissue in the host plant, and his description of
P. graminis was vague (Ainsworth 1981:34, 43).
John Ramsbottom (1913:81–84) evaluated Persoon’s classification of the Uredinales, which
classification is probably a fair sample of Persoon’s Synopsis. Persoon began a revision of his Synopsis
entitled Mycologia Europea (three volumes, 1822–1826), but left it incomplete, no doubt because a
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Bulletin of the Ecological Society of America
younger rival, Elias Magnus Fries (1794–1878)
had begun his authoritative Systema Mycologicum
(three volumes, 1821–1829 + 2 supplements,
1830–1832).
Fries quite literally followed in the footsteps
of Linnaeus: his father was a church vicar from
the southern Swedish province of Småland, he
studied at the Växjö secondary school, then at
the University of Lund, and eventually became
Professor of Botany at Uppsala University
(Eriksson 1952, 1972, Fries 1952, Ainsworth
1976:259–263). At Lund he studied fungi,
but before going there he had independently
learned between 300 and 400 species (compared
to Linnaeus’ listing of 92 species). He found
Persoon’s classification unsatisfactory, and Fries’s
Systema Mycologicum “did for mycology what
Linné did for phanerogamic taxonomy,” with
his descriptions being “models of accuracy and
conciseness” (Fries 1952:180). His Systema is the
starting point for fungal names, just as Linnaeus’s
names are for higher plants; however, Fries
worked without a microscope. He did not confine
his studies to fungi, but his other studies are
beyond the scope of this discussion. Elias Fries
was the first of four generations of Fries botanists.
We met Sir Joseph Banks (1743–1820) in
part 43 (Egerton 2012:200) as the botanist with
whom Thomas Andrew Knight (1759–1838)
corresponded, and who published Knight’s letters
in the Philosophical Transactions of the Royal
Society of London. Banks was president of the
Royal Society for 42 years, so he was a wellrespected member of the scientific community
(Foote 1970, Carter 1987, O’Brian 1997, Knight
2004). In 1805, in response to an extensive
outbreak of black rust (Puccinia graminis) in
1804 (Ordish 1976:115), Banks published A Short
Account of the Cause of the Disease in Corn,
called by Farmers the Blight, the Mildew, and
the Rust, in which he asked, “Is it not more than
possible that the parasitic fungus of the barberry
Fig. 2. Black rust Puccinia graminis, figures
57–59 (upper left corner). Drawn by John
Edward Sowerby (1825–1870). Cooke 1865:
Plate 4, facing 56.
and that of wheat are one and the same species,
and that the seed transferred from the barberry
to the corn [wheat], is one cause of the disease?”
(quoted from Ramsbottom 1913:85). He had read
Felice Fontana’s Observazioni sopora la ruggine
del grano (1767), which was little known in
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Fig. 3. (a). Joseph Banks. (b) Thomas Andrew Knight. Simmonds 1954:following 468.
Britain. One would think that Banks also would have read A Botanical Arrangement of All the Vegetables
Naturally Growing in Great Britain (edition 1, 1776) by William Withering (1841–1899), that claimed
the Berberis shrub “should never be permitted to grow in corn [grain] lands, for the ears of wheat that
grow near it never fill, and its influence in this respect has been known to extend as far as 300 or 400
yards across a field” (quoted from Ramsbottom 1913:81). However, Banks’ claim that “It has long
been admitted by farmers, though scarcely credited by botanists, that wheat in the neighbourhood of a
barberry bush seldom escapes the Blight” (quoted from Ramsbottom 1913:85) seems to indicate he had
not read Withering’s work. Banks’ pamphlet had two interesting enlarged color illustrations by Francis
Bauer (1758–1840) of the rust imbedded in the wheat tissue (one reproduced in Ordish 1976: plate 3;
see also Ainsworth 1969:15).
Knight had scooped Banks; in 1804 he had already sown wheat around a barberry bush and other
wheat from the same source a considerable distance away (Parris 1968:21–22). The wheat near the
barberry bush became diseased, as did the bush (quoted from Ramsbottom 1913:86).
Examining the barberry bush attentively, I found upon its fruit a species of fungus similar in
colour to that on the straws of the wheat, but its seed vessels were larger, and more spherical. I
was, however, much disposed to believe the parasitical plants of the same species, and that the
difference in the form and size of the seed vessels arose only from the difference of the nutriment
they derived from the wheat and from the acrid juice of the barberry.
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The distant (control) wheat was healthy until Knight took a diseased barberry branch to that wheat,
where he sprinkled the branch with water and brushed it over the healthy wheat, which 10 days later
became diseased. Simultaneously, he took some diseased wheat to healthy wheat, moistened the diseased
wheat and brushed it over the healthy wheat, but the healthy wheat remained healthy. Ramsbottom
(1913:86) suggested that Knight was “the first to try inoculation experiments on heteroecism.” Knight
did not write a letter to Banks about this experiment until after he had read Bank’s pamphlet, on 20
March 1806. In 1806, Banks published a second edition of his pamphlet and appended Knight’s letter
(Banks 1806:26–36, Ramsbottom 1913:85, Dawson 1958:501).
In Philadelphia, a retired physician and immigrant from Bath, England, Anthony Fothergill (1732?–
1813), read a paper on 11 November 1806 to the Philadelphia Society for Promoting Agriculture, arguing
that grain rust was caused by “clusters of a fungus or parasitical plant…[that] insinuate themselves
into the pores of the absorbent vessels of the stem, and deprive the grain of the sap destined for its
nourishment” (1808, quoted from Campbell et al.1999:33). In 1818, a Danish schoolteacher, Schoeler,
carried out the Knight experiment with the same results (Parris 1968:24). Knight recommended in 1818
the use of flowers of sulfur against pear scab (Lycoperdon cancellatum), and in 1834 he recommended the
sprinkling in early spring of peaches with sulfur and lime to control leaf curl. John Roberts recommended
in 1821 using soap to cause wetting of sulfur in water to control peach mildew, and in 1824 he reported
his successful experimental transmission of peach mildew (Knight 1818, 1842, Roberts 1824, Parris
1968:25–26, Ainsworth 1976:160, 1981:32–33, 110).
Phytopathologist George L. McNew divided the history of his profession into five periods, with
periods three and four occurring during the 1800s (McNew 1963:166–172). Period three he labeled
“The Predisposition Period,” during which the prevailing view was that fungi originated in diseased
plant tissue. Three authors published treatises on plant pathology and remedies in 1807. Freiherr von
Werneck and Filippo Ré represented the consensus in not accepting fungi as the cause of the diseases
(Whetzel 1918:32, Parris 1968:23, Walker 1969:20–22). Isaac-Bénédict Prévost (1755–1819) did think
fungi caused diseases (Keitt 1939a, Large 1940:76–79, Robinson 1975, Ainsworth 1981:30–32). He
was the son of a teacher-pastor in Geneva, but at age 22 he became a tutor in Montauban, France, where
he remained for the rest of his life. His interests included mathematics, physics, chemistry, biology,
and philosophy. In 1797 a member of the Society of Montauban read a memoir on the carie or charbon
(bunt or smut) disease of wheat, and the society asked other members to study the problem. Prévost
read previous studies on the problem, including those by Duhamel du Monceau (1728) and Mathieu du
Tillet (1755), discussed in part 29 (Egerton 2008), and especially Henri Alexandre Tessier’s Traité des
maladies des grains (Paris, 1783), which attributed rust to stoppage of transpiration, caused by mists.
Tessier repeated some of Tillet’s experiments and agreed that bunt was contagious (Parris 1968:19).
Tessier also stressed the importance of soaking wheat seeds in lime water before planting, which greatly
diminished the loss of wheat from bunt in France (Large 1940:76). The owner of a nearby estate made
available to Prévost land for large-scale experiments.
George Keitt (1939b) provided an excellent guide to and evaluation of Prévost’s achievement along
with his English translation of the Mémoire. Prévost could not discover everything he wished, but he
was remarkably thorough and clear in the presentation of his findings. Tillet had demonstrated in 1755
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Fig. 4. Germination and growth of bunt spores. Prévost 1807, 1939:
Plate 1, Figs. 1–32.
that bunt (carie, charbon) of wheat was spread by an infectious agent (Egerton 2008:237), but had not
demonstrated that the agent was a fungus. Prévost was able to induce the bunt dust to germinate in water
and illustrated its growth, thus demonstrating the dust was fungal spores. He illustrated its progressive
development on the first of his three plates.
To those believing a fungus arose within the wheat plant, he replied that he only observed branches
from the spore growing from the surface into the seedling (1939:34–35, paragraphs 44–46). Fruiting
bodies grew within ears of wheat and spores were disseminated by the wind. Prévost scattered spores
on soil where wheat seed was planted and concluded that it was only at germination, or soon after, that
infection occurred (Prévost 1939:38, paragraph 57). He discovered that copper dust suspended in water,
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or liquid copper compounds, inhibited spores from germinating (Prévost 1939:72–82, paragraphs 138–
177, Ainsworth 1976:160–161, 1981:124–126), and arsenic was reported in England to have the same
effect (Prévost 1939:83, paragraph 178). The Society of Montauban sent a published copy of Prévost’s
Mémoire to the Institute [de France?] in Paris for evaluation. Tessier reported for a review committee
favorably, but no actions were taken.
Mendel was by no means the only author of a clear publication that failed to gain the recognition
deserved upon publication. Lavoisier, Lyell, and Darwin achieved their scientific revolutions in part
because of prominence in their scientific communities. Prévost and Mendel were amateurs on the
periphery of their scientific communities, who were neglected, rather than being either challenged or
widely recognized. Nevertheless, Prévost gave the first experimental proof of the pathogenicity of a
microorganism, and he first described motile spores (zoospores) in fungi (Ainsworth 1976:62, 147).
In Italy, Agostino Maria Bassi (1773–1856), like Prévost, was on the periphery of science, but his
important parasitical work did attract the attention it deserved, because he promoted it (Belloni 1961:20–
26, Robinson 1970, Ainsworth 1976:163–168). He grew up near Pavia and attended its university. He
studied law there to please his parents, but his interests were in science, and he studied under Antonio
Scarpa, anatomist, Alessandro Volta, physicist, and Giovanni Rasori (1766–1836), pathologist. Rasori
(portrait in Belloni 1961:22) wrote a book defending the contagium vivum theory that appeared
posthumously in 1837. Both Rasori and Bassi were influenced by the writings of Lazzaro Spallanzani
(1729–1799), who discredited the theory of spontaneous generation and accepted the conclusions of
Fontana and Targioni-Tozzetti that small parasitic plants can cause diseases in vascular plants (Egerton
2008:235, 239).
The silk industry could flourish in northern Italy and in France, if it could eliminate an epidemic
disease, muscardine (calcinaccio). In 1807 Bassi decided to investigate; he grew silkworms, assuming
the disease (Bassi 1958:4)
…arose spontaneously in the silk worm and was due to some difference in the atmosphere, the
food, or the method of breeding, or rather to the various fumes emanating from the fermenting
litter…
He varied environmental conditions for different groups but failed to induce the disease in his
caterpillars. When he obtained diseased caterpillars, he found the disease spread at temperatures of 7°–
30°R, but not above 38°R (Bassi 1958:8). The disease could be transmitted to caterpillars of other species.
It was contagious, because it always began with one or a few caterpillars, whereas an environmental
or physiological disease could affect all individuals about the same time. Bodies of dead caterpillars
became covered with “a patina or efflorescence like flakes of pure snow” (Bassi 1958:7).
In 1835, Bassi announced that muscardine was caused by a fungus, and he added a footnote (Bassi
1958:10, note on 15)
The eminent and meritorious compilers of the celebrated Giornale Fisico-Chimico, Professors
Configliachi and Brugnatelli, were the first to put forward the hypothesis that the mark disease
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like smell that comes from worms which have died of that disease; and although, in that journal,
they appealed to breeders to undertake experiments to test this supposition, these last, perhaps
regarding it as the figment of a heated imagination, neglected it, whereas, had they taken the
trouble to examine it, they might easily have found, by experiment, that these able men were not
mistaken and had really hit on the truth.
Although Bassi blamed the silkworm growers for their indifference to the professors’ suggestion, one
may wonder how many growers ever read this physico-chemical journal or even knew how to undertake
an experiment. Neither Bassi nor his English editors gave a more precise reference to the professor’s
comments than is in this footnote. Nor does Bassi indicate whether his investigation was undertaken in
response to their comments, but if he had made his discovery without having read their suggestion, he
perhaps would have said so. In another footnote, he stated (Bassi 1958:17)
If this parasitic plant is observed with the great microscope of the illustrious De Amici, which
magnifies the object more than thirty million times (sic.), it will be possible to see all its minutest
ramifications in it, and perhaps its reproductory organs as well.
He explained that the “disease-bearing dust” was spread by air, water, and dogs, cats, rats, mice, flies,
and contaminated food for silkworms (Bassi 1958:23). The English translation of Bassi’s treatise is
limited to part 1, on theory (1835). In 1836 Bassi published part 2, on ways to combat the disease. These
included disinfecting or burning containers and materials used in raising them (Ainsworth and Yarrow
1958:x–xi). Bassi realized that his discoveries would be controversial, and he requested permission to
perform his experiments for faculty members at the University of Pavia. They agreed, and afterwards
provided a testimonial signed by nine professors, including Pietro Configliachi and Brugnatelli,
mentioned in the footnote quoted above. Bassi then published their testimonial in his preface to part
1 (Bassi 1958:1–2). Giuseppe Balsamo-Crivelli named the fungus Bassi discovered Botrytis Bassiana
(now Beauveria bassiana), and an abridged French translation of Bassi’s part 1 was published in Paris
in 1836. Both botanist Jean-Francois-Camille Montagne (1784–1866) and entomologist Jean-Victor
Audouin (1797–1841) investigated Bassi’s claims for the Académie des Sciences in Paris and published
confirmations, with a plate of illustrations of the fungus (Audouin 1836, Montagne 1836, Belloni
1961:25–28; Audouin’s illustration is reproduced by Ainsworth 1976:167). This was the first known
fungal parasite of an animal; Prévost had already demonstrated a fungal parasite of plants.
In the early 1840s, two Jewish physicians published evidence of fungal disease in humans. David
Gruby (1810–1898) was a Hungarian who obtained his doctorate in Vienna in 1840 and immigrated to
Paris, where he practiced and published (Kisch 1954:193–226, Ainsworth 1976:169–171). Although
the claim that he was one of the most brilliant biologists of the 1800s (Zakon and Benedek 1944:155)
seems an exaggeration, his discoveries were important for both medicine and phytopathology. His
six important scientific papers (1841–1844), all rather brief, are translated from French into English
(Zakon and Benedek 1944:157–168). In 1841 he published his microscopic observations showing that
favus was caused by a fungus; in 1842 he indicated that ringworm of the beard was caused by an
ectothrix trichophytosis. He named neither of these fungi, but in 1843 he identified the cause of human
microsporosis and named the fungus Microsporum audouinii to honor Victor Audouin. Even though his
discoveries were easily repeated by any competent microscopist, the Paris medical community remained
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Fig. 5. (a) Agostino Maria Bassi. Bassi 1958: facing title page. (b) David Gruby, age 75. Ainsworth
1976:169.
skeptical of his conclusions (Zakon and Benedek 1944:156). He also discovered several microscopic
invertebrate parasites during the early 1840s (Foster 1965:115–116), before devoting the rest of his life
to a very successful medical practice. Robert Remak (1815–1865) was Polish and immigrated to Berlin
at age 18. In 1837 he discovered the fungal nature of favic crusts but did not publish this until 1845. He
was then a physician in a clinic run by Professor J. L. Schönlein, and Remak named the fungus Achorion
schoenleinii, honoring Schönlein (Kisch 1954:227–296, Ainsworth 1976:168–169).
Following in Gilbert White’s footsteps as an English clergyman–naturalist, Miles Joseph Berkeley
(1803–1889) had a childhood interest in nature that led eventually to his becoming the foremost
British mycologist of the 1800s (Massee 1913, Whetzel 1918:55–57, Ramsbottom 1948, Ainsworth
1969:14, Taylor 1970, Stafleu and Cowan 1976–1988, I:192–195, Desmond 1977:60, Buczacki 1991,
Elliott 2004a). He attended Cambridge University, 1821–1825, but left two years before John Stevens
Henslow became its professor of botany. As an undergraduate, he collected algae, mosses, and mollusks.
Berkeley’s earliest published papers were on mollusks, and he only began to specialize on fungi in 1832,
when William Dalton Hooker invited him to write the fungal volume of The English Flora (Berkeley
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Fig. 6. (a) Miles Joseph Berkeley. Ainsworth 1976:155. (b) Anton de Bary. Jost
1930:facing 1.
1836). Berkeley’s introduction to plant pathology occurred in 1845, when the potato murrain, that had
been discovered in Liège, Belgium in 1842, and occurred in the eastern United States in 1843 (Stevens
1933, Campbell et al.1999:38–39) and in Western Europe in 1844, then spread to Britain and Ireland
(Berkeley 1948:14–17, Bourke 1964, 1969). When its seriousness in Ireland became evident, British
Prime Minister Sir Robert Peel formed a scientific commission to study the matter (Large 1940:26–27,
Woodham-Smith 1962:44–47), under London Botany Professor John Lindley (1799–1865; Stearn 1999)
and London Chemistry Professor Lyon Playfair (1818–1898), who had received his Ph.D. under Liebig
(Large 1940:26–27, Woodham-Smith 1962:44–47). The commission went to Ireland but did not find a
clear cause of the disease.
Montagne (mentioned above) found a fungus associated with blighted potatoes, which he named
Botrytis infestans (now Phytophthora infestans) on 30 August 1845 (portrait in Virville 1954:198;
Ainsworth 1976:154–155, 1981:54–55). He was uncertain about whether the fungus caused the murrain,
but sent his comments and drawings to his regular correspondent, Berkeley (Lamy 1989), who, at first
(1845) shared Montagne’s uncertainty (Walker 1969:23). However, Dr. C. F. A. Morren, head of a
school of agriculture in Liège, believed the fungus caused the murrain (1845), and he conducted crude
inoculation experiments to support the claim (Walker 1969:23). Berkeley agreed with Morren before
publishing his own study on the disease (1846).
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Berkeley also published Montagne’s drawings
(one of which is in Fig. 7). Berkeley presented
the case for the disease having an environmental
cause and cited the authorities who supported
this theory (1948:23–28), and then he argued the
case for “The decay is the consequence of the
presence of the mould, and not the mould of the
decay” (Berkeley 1948:28). He ended his article
(Berkeley 1948:35–37) with a Latin and English
description of Botrytis infestans Mont. and related
species (many of which, like infestans, are no
longer placed in that genus). Large (1940:32)
comments that Berkeley’s article swept away
many but not all objections. Berkeley did not
observe spore germination, nor did he document
the actual infection of the potato plants.
Berkeley became both mycologist and plant
pathologist, and he entered the fray in 1847 over
the cause of a new grapevine disease, discovered
by a gardener, Tucker, in 1845 near Margate,
England. Tucker reported that he controlled this
powdery mildew with a mixture of sulfur and lime
in cold water. Berkeley studied the mildew, named
it Oidium Tuckeri, and illustrated it with hyphae
growing within the leaves, as he had drawn B.
infestans. The French physician–mycologist
Joseph-Henri Lévillé (1796–1870, portrait in
Virille 1954:218) did not accept the claim that O.
Tuckeri caused the downy mildew, and he found
no hyphae within the leaf tissue, only on the leaf
surface (Large 1940:44–45). Berkeley translated
Lévillé’s discussion and added his own response
(Lévillé and Berkeley 1851). Lévillé was right
about O. Tuckeri not having any detectable hypha
within the leaf tissue, but wrong in thinking that
morbid tissue gave rise to the fungus. In 1851,
Dr. Zanardine, in Venice, argued that the surface
fungus, O. Tuckeri, was indeed parasitic, because
it had little suckers he called “fulcra” (now
“haustoria”) that obtained sustenance from leaves.
Skeptics could still argue that the fungus may not
have arisen if the leaf was not already moribund,
but the consensus was shifting toward Berkeley
Fig. 7. Fungus Botrytis infestans from potato
murrain. Fig. 5: horizontal section of diseased
cells, drawn by Jean Montagne; Fig. 9: part of
the underside of potato leaf showing fungal
hyphae with fertile shoots [sporangiophores with
sporangia]; Fig. 10: fungus growing out of leaf
stomata of potato plant. Figs. 9 and 10 apparently
by Berkeley. Berkeley 1846: part of plate 2, 1948.
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(Large 1940:44–49). Berkeley published 173 articles on “Vegetable Pathology” in the Gardner’s
Chronicle, 1854–1857, some of which are reprinted (Berkeley 1948:41–108). He also published a steady
stream of articles on fungi: 35 articles on 2050 British fungi, 1837–1885; 7 of these articles are only
his and 28 were coauthored by C. E. Broome; all are now reprinted (Berkeley and Broome 1967). In
addition, he was one of three European mycologists who described fungi brought back from elsewhere
in the world, the other two being Montagne in France and Elias Magnus Fries (1794–1878) at Uppsala
University, Sweden, whose Systema mycologicum (three volumes, 1821–1832) is recognized as one of
the two foundations of modern fungal nomenclature, along with Persoon’s Synopsis (Fries 1950:61–63
+ portrait on frontispiece, Eriksson 1972, Ainsworth 1976:273). Berkeley published his descriptions
of foreign fungi in batches of 10 descriptions per article (or at least 10 in the first article, and various
numbers after that, but all called “decades” nevertheless), for 27 articles, 1844–1856, three of which
were coauthored, and all of which are now reprinted (Berkeley 1969). Berkeley was a careful observer
but was not an experimentalist.
(Heinrich) Anton de Bary (1831–1888) was the leading physiological mycologist and a leading
phytopathologist of the 1800s and was an outstanding experimentalist (Whetzel 1918:45–47, Jost 1930,
Robinson 1971, Sparrow 1978). He was also one of the most respected and revered professors in that
century. He was the son of a physician in Frankfurt am Main who raised fruit trees and flowers, and as
a teenager Anton joined a local group of amateur naturalists on field trips into the nearby countryside. A
physician–botanist at the Senckenberg Medical Institute, Professor Georg Fresenius, introduced him to
the study of algae and fungi. After graduating from the Frankfurt Gymnasium (high school) in 1848, de
Bary studied medicine at Heidelberg, Marburg, and Berlin, receiving his M.D. degree in 1853. He had
published his first scientific paper in 1852, on the water mould Achlya prolifera, from a peat bog near
Berlin (illustration in Sparrow 1978:234). In 1853 he published his M.D. dissertation on the sexuality
of plants and a book on fungal rusts and smuts. Berkeley had published strong evidence that the potato
murrain was caused by a fungus, but without convincing all skeptics. De Bary’s Untersuchungen über die
Brandpilze und die durch sie verursachten Krankheiten der Pflanzen (1853) undermined the remaining
doubts that fungi do cause diseases. McNew’s fourth period of phytopathology is “The Etiological
Period,” which began with de Bary’s Untersuchungen (McNew 1963:167–168).
De Bary’s “phytopathological classic” is now translated into English (De Bary 1969), and one can
see why it ended doubts on disease causation. It is divided into three parts: I. Specific observations
concerning the form and development of the brand fungi; II. Systematic conclusion; III. Concerning the
relationship of the brand fungi to the brand and rust diseases of plants. By describing various life cycles
(I), he provided the basis for a new classification (II), which enabled botanists to discuss specific fungi
in relation to specific vascular plant hosts. Paradoxically, botanists who believed the fungi were products
of disease rather than the cause—he provided extensive citations in his notes (De Bary 1969:64–67)—
still thought of the fungi as distinct species. In contrast, he listed only 10 botanists who were convinced
that fungi were the cause of disease (De Bary 1969:68). De Bary pointed out that one can observe the
germination of spores and show their capacity to infect, the hereditary nature of the brand fungi, and
their relation to stomata of plants they infect. He also provided eight plates showing fungi in relation to
their hosts. Any remaining skeptics would have to show how the leading authority on these fungi was
wrong. None tried it.
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Fig. 8. Fertilization and overwintering stage of Cystopus candidus Lev.
fungus causing white rust of cabbage and mustard. De Bary 1863, from
Large 1940:102.
He returned to Frankfurt to practice medicine and realized he was interested in diagnosing illnesses, but
not in curing patients, and he lost interest in being a physician. In 1854 he discovered that a myxomycete
spore gave rise to a naked flagellated myxamoeba, not a hypha, and later he postulated that myxamoebas
fuse to form a plasmodium (Sparrow 1978:226).
De Bary attended a botany meeting at the University of Tübingen, where Professor Hugo von Mohl
helped him obtain a faculty position at the University of Freiburg im Breisgau, where he taught, 1855–
1866, and established the first teaching botanical laboratory. In 1866 he moved to the University of
Halle, where he expected to develop a botanical institute, which only materialized in his last year there,
1871–1872. After the German victory in the Franco-Prussian War, he moved in 1872 to the University of
Strasbourg and was elected the rector of the university. In October 1872, the first American who arrived
there to study with him found that university enrollment had shrunk from a prewar 1500 to 250, and
there were only three other botany students (Harris 1945:12–13). However, de Bary soon became head
of a fine botanical institute, and the building he had built for it was so spectacular that a later American
student wrote an article on it with two drawings and two floor plans and a drawing of its greenhouse
(Dudley 1888; photograph of institute in Sparrow 1978:248). De Bary excelled at leading field trips and
guiding laboratory research and the writing of dissertations (Ayres 2005:42–49). He advised 94 doctoral
students, with dissertations mainly in mycology, but some on other cryptogamic species (students listed
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Fig. 9. (a) Julius Kühn. Ainsworth 1981:36. (b) Louis-René Tulasne. Virville 1954:218.
in Sparrow 1978:245). He had his students study a species’ life cycle. He ended the debates about the
cause of late blight in potatoes (1861) and the heteroecious life cycle of wheat–barberry rust (1863).
He published the first textbook on the morphology and physiology of fungi, lichens, and myxomycetes
(1866, revised and enlarged edition, 1884). His achievements in mycology and phytopathology are too
numerous to list them all here, but Ayres devotes an excellent chapter to de Bary, which includes a succinct
outline of his achievements (Ayres 2005:44). De Bary also played an important role in lichenology
(Mitchell 2012), to be discussed in part 52 of this history, on symbiosis.
The other founder of modern plant pathology was Julius Kuehn (or Kühn, 1825–1910), who wrote
the first modern textbook on it (Whetzel 1918:47–53, Large 1941:93–94, Wilhelm and Tietz 1978,
Ainsworth 1981:36–38). He was a remarkable scientist because he was largely self-taught. He was from
Pulsnitz, Germany, the son of a landowner. He began his career, 1848–1855, as manager of a 750-ha
estate in Silesia. In 1850 he wrote to a prominent botanist, Matthias Jakob Schleiden (1804–1881),
who is remembered as a founder of the cell theory. Kühn was concerned about a beet disease, which
Schleiden thought was caused by unbalanced nutrition (Wilhelm and Tietz 1978:351). In 1851 he had
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Fig. 10. Stigmatea fragariae and Pleospora pellita. Tulasne and
Tulasne 1861–1865, II. From Virville 1954: Plate 11.
more luck when he wrote to G. Ludwig Rabenhorst (1806–1881), who later became the founding author
of Kryptogamen-Flora von Deultschland, Oesterreich und der Schweiz (11 volumes, 1884–1960). Kühn
sent him an alga that plugged his field drains, which turned out to be a new species Rabenhorst named
Leptothrix kuehneana (which did not, of course, unclog the drains).That began a valued correspondence
that lasted for many years. Kühn published three of his early scientific papers in Rabenhorst’s journal,
Hedwigia (1855–1858).
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In 1856 Kuehn published two articles on the life cycle of Claviceps purpurea, a parasite that causes
ergot in rye grains and causes ergotism in animals and humans that is often fatal. His biographers
(Wilhelm and Tietz 1978:353) think that the article in the Leipzig weekly, Agronomische Zeitung,
was a prototype for his textbook, Die Krankheiten der Kulturgewaechse (1858). The article refuted
Schleiden’s theory that cultivated plants have a disposition to disease, using Kuehn’s inoculation of
wild grasses with spores as evidence. In 1855–1856, he attended the Agricultural Academy at BonnPoppelsdorf, and in 1857 the University of Leipzig awarded him a Ph.D. on the basis of his published
papers. He then managed another Silesian estate for five years, and then his textbook enabled him to
obtain a new agricultural chair at the University of Halle in 1862, where he remained for the rest of his
career. He taught some 5000 students at his Agricultural Institute, though only one of them became a
phytopathologist, Reinhold Wolff, in 1873.
The Tulasne brothers, Louis-René (1815–1885) and Charles (1816–1884), collaborated on the study
of fungi, after an inheritance in 1839 freed the former from the study of law and the latter from the study
of medicine (Ainsworth 1976:26–28, Viennot-Bourgin 1976,). Louis attended botanical lectures in Paris
and then became an assistant naturalist at the Muséum d’Histoire Naturelle, and Charles became an
outstanding botanical illustrator. Berkeley had demonstrated that there are two fruiting forms in four
genera of Gasteromycetes, which Louis confirmed and expanded in a series of 10 memoirs; the last of
these memoirs, Fungi hypogaei (1851), “remains one of the foundations of the modern study of this
group” (Viennot-Bourgin 1976). In his second memoir on Uredinales (1854), Louis showed that Uredo
and Phragmidium represent different stages of the same species (Ramsbottom 1913:88–89). The brothers’
major contribution to mycology was “the establishment of pleomorphism among fungi by a wealth of
accurate and detailed observation, first in a series of papers dealing with pyrenomycetes, discomycetes,
and basidiomycetes published between 1851 and 1860” (Ainsworth 1976:28). The climax of their work
was a magnificent Selecta fungorum carpologia (three volumes, 1861–1865, English version 1931).
Another English amateur botanist who rose to professional prominence was Mordecai Cubitt Cooke
(1825–1914), but unlike Berkeley, he lacked a university education, though he did take botany courses at
the Science and Art Department at South Kensington (Ramsbottom 1915:172). He held a variety of jobs,
including teaching and museum work. In 1862 he helped found The Society of Amateur Botanists and
became its only president. In 1865 he became editor of the new Hardwicke’s Science Gossip, and a reader
suggested establishing an amateur microscopical group, since the Royal Microscopical Society did not
meet amateur needs. This led to the formation of the Quekett Microscopical Club, and Cooke was also
asked to become its president, but declined. He successfully urged the club to adopt the French metric
system of measurements. In 1861 he published an introductory botany textbook, the first of some 32
books he would publish, all on botanical topics, excepting one on reptiles and amphibians (Ramsbottom
1915:175–176, Freeman 1980:91–93). Cooke’s Rust, Smut, Mildew, and Mould: An Introduction to the
Study of Microscopic Fungi (1865) was possibly the first book on plant diseases written for a popular
audience, with 12 color and 4 black and white plates, containing 269 figures drawn by John Edward
Sowerby (Fig. 2). As mentioned above, Berkeley described 2050 species of British fungi; Cooke, in
his Handbook of British Fungi (1871), described 2810 species. He gave up editing Hardwicke’s Science
Gossip in 1872 to found Grevillea, devoted to cryptogamic botany, which he edited until 1892. In
1887 the Royal Botanic Garden, Kew bought his herbarium of 46,000 specimens and 22,000 drawings.
“Cooke had a greater influence on the study of fungi than any other Englishman with the possible
exception of Berkeley” (Ramsbottom 1915:184).
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Fig. 11. (a) Harry Marshall Ward. Ainsworth 1981:58. (b) Mikhail Stepanovich
Voronin. Voronin 1934: Frontispiece.
Harry Marshall Ward (1854–1906) was one of de Bray’s students. As a teenager, he joined the
Nottingham Naturalists’ Society. In 1872 he attended the Science School in London, founded in that year
and later renamed Royal College of Science (Thiselton-Dyer 1913, Desmond 1977:639, Junnker 2004,
Reisz 2004, Ayres 2005:27–29). He attended Cambridge University, 1876–1879, then went to Würzburg
to study plant physiology under Julius Sachs, where he met Francis Darwin, who went with him to visit
de Bary’s laboratory in Strasbourg (Ayres 2005:41). Rather brief training at Strasbourg qualified him
to be a British government cryptogamist, sent in early 1880 to Ceylon (Sri Lanka) to study coffee leaf
disease. The disease, which caused the loss of leaves, had been discovered on a single estate in 1869, but
had since spread rather widely. George Thwaites, Director of the Royal Botanic Garden, Peradeniya, had
sent infected leaves to Berkeley, who described from them a new genus and species, Hemileia vastatrix
(Ayres 2005:1–5). Ward used both de Bary’s methods and the technique of Dr. Charles H. Blackley
to study hay fever (1873), of hanging sticky microscope slides among coffee leaves to collect spores.
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Ward succeeded in two years in describing the life cycle of H. vastatrix, but he was unable to find an
effective means to end the epidemic, and coffee declined as a crop in Ceylon (McCook 2006:178–183,
2011:96–100). After leaving Ceylon, Ward returned to Strasbourg for two months of further study under
de Bray. After returning to England, he taught at Owen College, Manchester and became active in the
British Association for the Advancement of Science. In 1885 he became professor of botany at the Royal
Indian Engineering College.
Ward’s major contribution was establishing physiological plant pathology (Ayres 2005:39), though
he built upon de Bary’s work. “A Lily Disease” (Ward 1888), his most important paper, focused on
controlling disease by studying the interactions between plants, parasites, and environment. The fungus
secreted something from the tips of hyphae that broke down host cell walls. Kuehn had coined the word
“enzyme” in 1867, yet in 1888 there was no agreement on its meaning (Ayres 2005:80–81). Ward saw
two patterns of fungal attack: (1) biotrophic pathogens, such as rusts, that caused minimal disruption to
host, because enzymes released by hyphae enabled tips to grow into the cell wall, with host cells living
longer, and (2) necrotrophic pathogens, such as Botrytis, that released pectic enzymes disrupting cells,
allowing hyphae to grow rapidly, but killed host cells. Botrytis had been regarded as saprophytic, but
Ward showed it could also become parasitic. Ward spoke of ferments or enzymes (1888:365). Despite
the article’s importance, a modern scientific journal editor would likely ask the author of a 60-page
article of this sort for a much more concise presentation of evidence.
Francis Darwin had been lecturing on botany at Cambridge University since 1884, but when the chair
became vacant in 1895, he supported Ward for it. Ward received the position and remained active as
teacher, administrator, and researcher.
Casimir-Joseph Davaine (1812–1882), according to a Canadian phytopathologist (Estey 1975:549),
was “one of the most remarkable biologists of the nineteenth century,” with important contributions to
phytopathology, parasitology, and microbiology. He was the son of a distiller in St.-Amand-les-Eaux
and was educated in northwestern France before going to Paris to study medicine. He never held an
academic position, but did research while practicing medicine in Paris, beginning in 1838 (Théodoridès
1968, 1971). His research, 1854–1856, on the nematode Anguina tritici, which caused seed-gall disease
in wheat, was his “most comprehensive achievement in plant pathology” (Estey 1975:551) and the first
documented nematode disease in plants. In 1856, the Académie des Sciences awarded him 1500 francs
for his nematode studies. He was also first to document a plant disease caused by bacteria, which he
transmitted from one plant to another by inoculation (Davaine 1868a, translated in Estey 1975:549).
Among the succulent plants, or the vegetables with very tender and moist parenchyma, many
times I have seen a change that first appeared at the root and shortly invaded the rest of the plant
completely destroying it in a few days. This change, which reduced the tissues to a kind of rot, is
caused by the development of bacteria that differ from those of anthrax in as much as they have
movement. We can easily transmit this disease from one plant to another by inoculation: around
the inoculation point an oil-like spot appears that grows and takes over the whole plant if the
diseased part is not cut off.
Davaine knew that Pasteur heated wine to eliminate contaminating microorganisms, and therefore
he tried, successfully, to kill bacteria without killing the plant, at a temperature slightly above 52°C
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Fig. 12. Infected cabbage and turnips. Voronin 1934: Plate 30.
(Davaine 1868b). On another occasion, Pasteur acknowledged following Davaine’s lead.
Mikhail Stepanovich Voronin (Michael Stephanovitch Woronin, 1838–1903) was the founder of
Russian phytopathology (Chupp 1934, Senchenkova 1976). He was born into a wealthy family and
studied geology and botany at St. Petersburg University. He graduated in 1858 and went to Germany
for two years and studied under Holle in Heidelberg and de Bary in Freiburg, studying both algae and
fungi. Being independently wealthy, he returned to St. Petersburg and established his own laboratory
and only taught courses on mycology at the university in 1869–1870 and 1873–1875. He gave money
to the St. Petersburg University to build its Botanical Institute and subsidized other botanical projects.
Novorossysk University in Odessa awarded him an honorary doctorate in 1874, and in 1898 he was
elected a member of the Russian Academy of Sciences and headed its cryptogamous plants section.
Of Voronin’s 106 research publications, the most important for phytopathology were “Research on
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Fig. 13. (a). Adolf Mayer. (b). Tobacco leaves infected by mosaic disease.
Mayer 1886: Plate 3. From Johnson 1942: facing pages 9 and 12.
the Development of the Rust Fungus Puccinia helianthi, which Causes the Sunflower Disease” (1871),
“Organism Causing the Disease of Cabbage Known as Kila” (1877), and “On “Drunken Bread” in the
Southern Ussuri Region” (1890). A collected edition of his most important writings was published in
1961. In his study on ergotism from contaminated rye bread, he identified 15 fungi, 4 of which caused
the disease. He identified the cause of cabbage clubroot as a slime mold he named Plasmodiophora
brassicae, and his study on it (1877) is a classic, translated into English (1934).
Since fully-grown cabbage plants could get clubroot, all stages of growth were susceptible. The
growths were variable, as seen in two illustrations of whole roots (Fig. 12). He also provided four
microscopic drawings showing fungus and cabbage tissue.
The number of spores produced per fungus was unusually large, and their size unusually small. They
were released into the soil as the infected cabbage roots disintegrated. The spores burst and myxamoeba
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oozed out and with whip-like cilia moved
through soil water in quest of root hairs or root
epidermal cells. It had no cell wall, and Voronin
did not even find a cell membrane. Knowledge
of its life history enabled Voronin to suggest a
remedy: after cabbage was harvested, roots had
been left in the ground; they needed to be dug
up and burned. Cabbage seeds were grown in
beds and then seedlings transplanted into fields.
The seedlings should be carefully inspected to
eliminate any sickly ones being transplanted.
Finally, cabbage should be grown on a rotational
pattern—never the same field in consecutive
years (Voronin 1934:23–28).
Simultaneous with awareness of bacterial
plant disease was awareness of viral plant
disease (Johnson 1942, Waterson and Wilkinson
1978:23–31). By the mid-1800s, tobacco was an
important crop in The Netherlands, and in 1857
a college student working on a farm discovered
an unknown disease that eventually was made
known to the German director of the agricultural
experiment station at Wageningen, Adolf
Mayer (b. 1843; Fig. 14a). Mayer studied it for
several years, named it tobacco mosaic disease,
illustrated affected leaves in color, ruled out a
nutritional disease, but was unable to find either
fungi or nematodes in diseased plants. However,
he found that substance from diseased plants
ground up and mixed with water would infect
healthy plants, yet using Robert Koch’s methods,
he could not isolate bacteria. He filtered the
infective fluid through filter paper, and it was still
infective, but when filtered through double filter
paper it was not. Heating the infective fluid at
80°C for several hours killed its infectiveness. He
therefore concluded that it was likely a bacterial
disease (Mayer 1886, 1942).
Dmitri Iosifovich Ivanovsky (D. Ivanovski
1864–1921) (Ivanovski 1942) was an
undergraduate at St. Petersburg University in
1887, when he and another student were sent
Fig. 14. Martinus Willem Beijerinck.
Williams 1960: facing 84. Or Jost 1930:facing
1.
to Ukraine and Bessarabia to study wildfire
disease at tobacco plantations (Gutina 1973). He
concluded it was not contagious, and received
his degree in 1888. In 1890 another disease
appeared in tobacco plantations of the Crimea,
and the Department of Agriculture sent him to
study it. He thought that what Mayer interpreted
as two stages of tobacco mosaic disease were
actually two different diseases. Mayer seems
to have been correct on this point, but what
impressed other scientists was that Ivanovsky
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found that the sap of infected plants was still potent after it had been filtered through Chamberland filtercandles (Ivanovsky 1942). Like Mayer, he thought it likely was a bacterial disease.
Martinus Willem Beijerinck (1851–1931) was the son of a tobacco dealer who went bankrupt (Hughes
1978, Bos and Theunissen 1995). At the Delft Polytechnic School, his roommate was Jacobus van’t
Hoff, who won the first Nobel Prize for Chemistry in 1901. Beijerinck received a diploma in chemical
engineering in 1872, then switched in graduate school at the University of Leiden to botany and in 1877
received his Ph.D. with a dissertation on plant galls. He held several positions before returning to Delft
Polytechnic as a professor in 1895, where he became a popular teacher and active researcher (Williams
1960:73–87). His investigations in microbiology were quite diverse.
Beijerinck began teaching at the Agricultural School in Wageningen in 1876 and was a colleague
of Adolf Mayer, who showed him his experiments on tobacco mosaic disease in 1885. When Mayer
was unable to find a bacterium causing the disease, he went on to other things. Beijerinck (1898, 1899)
reasoned that the cause might be a liquid poison from a bacterium like tetanus, where the bacteria might
not be located at the site of the symptoms, and so not be discovered there (Beijerinck 1942:33–34).
However, diligent search did not discover them elsewhere in the plant, and he concluded that the disease
was not caused by microbes, but by a “contagium vivum fluidum.” However, that term was too long
and he soon called it “virus,” which was the first use of this term for this substance (Johnson 1942:6). It
remained infectious when filtered through porcelain that could remove bacteria. A very small amount of
the filtered virus could cause the infection of numerous tobacco leaves, which would not have occurred
with a small amount of toxin such as from tetanus bacteria. He was able to grow this virus on agar plates
and then infect plants with virus grown beyond the point of inoculation (Beijerinck 1942:35–36). His
curiosity exceeded Mayer’s, for he found that the virus can be dried without loss of strength and that it
could survive the winter in the soil. The virus was destroyed by boiling and even at 90°C (Beijerinck
1942:42–44). Since Erwin Smith (see below) had not found a bacterium that caused peach yellows in
1894, Beijerinck suspected that it also was caused by a virus, which Walker (1969:610) thought was
correct, but Campbell 1983:23) says it is caused by a mycoplasma.
In 1894, a privately funded Willie Commelin Scholten Phytopathology Laboratory was organized in
Amsterdam, and in 1895 Jan Ritzema Bos (b. 1850) became director for a decade (Faasse 2008). Bos had
been a professor at the Agricultural School in Wageningen, where he had developed a strong relationship
with farmers concerning plant diseases. Also in 1895 the Laboratory began publishing Tijdschrift over
Plantenzieken.
North America
Original contributions to phytopathology from North America were sparse and minor before the U.S.
Civil War (Stevenson 1959). In New York State, pomologist and agricultural editor John J. Thomas
published an article on “The Diseases and Insects Injurious to the Wheat Crop” (1844) in which he
drew upon the European literature to argue that fungi cause smuts and rusts, and he used a microscope
to see “a small plant of as regular and uniform a growth as the wheat itself” (quoted from Campbell et
al. 1999:34). America’s potato crop suffered from a blight in 1843, and in 1844 James Teschemacher
(1790–1853), an English immigrant to Boston, who published papers on several different sciences, wrote
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in a local agricultural journal that the blight was caused by a fungus, and he had used a microscope to see
its spores. He placed a diseased potato beside a healthy one under a bell jar and also buried a diseased
potato beside a healthy one and found the healthy ones still healthy after five days, but diseased after
two weeks (Peterson et al. 1992, Campbell et al. 1999:42–44). In 1847, Samuel Gookins reported to an
American horticultural and farming journal that Ruben Ragan, at his Indiana farm, had shown Gookins
where he had inoculated a healthy pear tree with sap from a pear tree that had fire blight, and some days
later the healthy tree showed signs of infection where it had been inoculated (Baker 1971:611–612,
Campbell et al.1999:56). Although Burrill would argue in 1879 that fire blight is caused by bacteria
(see below), before that time it was often assumed that all contagious plant diseases were caused by
fungi. By the 1850’s, America was gaining converts to the fungal theory of plant diseases (Campbell et
al.1999:57–59), and an influential Englishman (Berkley 1948:15) cited Teschemacher’s observations.
Although the Civil War in the United States was a setback to higher education, since young men went
off to war, the secession of the South enabled Congress to make significant progress, because sectional
strife within Congress ended. In 1862 Congress passed and President Lincoln signed bills creating both
the U.S. Department of Agriculture (USDA) and land grant colleges in every state, which would provide
agricultural education that included phytopathology (Elder 1962, Stefferud 1962a, b). Although a federal
bureaucracy was formed and a USDA building completed in 1868 (True 1937:41–48), there was no
progress in phytopathology during the 1860s either in Washington or in the states, but by the 1870s both
the USDA and state colleges became important for phytopathology in the United States and eventually in
the world (Baker et al. 1963, Campbell et al. 1999:62–65). In 1871, the USDA hired Thomas Taylor (b.
1820), who had designed and tested ordnance for the War Department during the Civil War, to man the
Department’s one microscope to study plant diseases. He quickly accepted the fungal theory of disease
for grape, pear, peach, and lilac diseases and published annual reports on these and other diseases,
1872–1877, but by the late 1870s, he was in competition with professionally trained mycologists and he
moved into the study of food adulteration (Stevenson 1959:19, Campbell et al. 1999:129–133, 407). In
1885, a botany committee from the American Association for the Advancement of Science petitioned
the new Commissioner of Agriculture, Norman J. Coleman, urging him to expand the research of USDA
to include the study of plant diseases. He supported the idea, and in turn urged academic botanists to
lobby Congress to appropriate funds for a Section of Mycology in USDA’s Division of Botany. Botanists
flooded Congress with letters, funds were appropriated, and Frank Lamson-Scribner became head of
mycology (Stevenson 1959:20, Campbell et al. 1999:136–139). Lamson-Scribner then wrote to Bessey
for guidance in establishing priorities for his research (Griffith et al.1994).
Yale graduate Samuel William Johnson (1830–1909) became Liebig’s outstanding American
student, 1854–1855; he returned to teach agricultural chemistry at Yale (Rossiter 1975:127–148).
Johnson embarked on a crusade to convince states to establish agricultural experiment stations (Rossiter
1975:149–171), and Connecticut did establish the first such station in 1876, and by 1880 California, New
Jersey, and North Carolina had followed suit. However, it was a struggle to keep them funded, and in
1887 Congress passed the Hatch Act to provide federal funds for state experiment stations (Byrd 1962,
Knoblauch and others 1962, Marcus 1985). In the 1890s, these stations began to make very substantial
research on plant diseases (True 1937, Campbell et al. 1999:181–203).
In 1872, William Gilson Farlow (1844–1919), a Harvard M.D. and Professor Asa Gray’s assistant,
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Fig. 15. (a) William Gilson Farlow. (b) Charles Edwin Bessey. Campbell et al.
1999:76, 78.
was the first American to go to Strasbourg to study mycology under de Bary (Setchell 1927, Harris
1945). Farlow’s father was a successful Boston businessman and state legislator, and he could afford
to send home a trunk full of books, including the Tulasnes’ Carpologia Selecta Fungorum, which he
commented to Gray was expensive (Harris 1945:18). When Farlow returned home in 1874, he became
assistant professor of botany at Harvard’s Bussey Institution, a school of agriculture and horticulture
founded in 1870 by the will of philanthropist Benjamin Bussey. While there, Farlow conducted research
on economically important plant diseases (Setchell 1927:5–6, Campbell et al. 1999:93–98 et passim,
Peterson 1999), though when he became a professor of botany at the Harvard campus in 1879, he
switched to mycological research. Nevertheless, he did train phytopathologists. His student and former
assistant, Roland Thaxter (1858–1932), served three years as a phytopathologist at the Connecticut
Agricultural Experiment Station, 1888–1891, before returning to Harvard to teach cryptogamic botany
(Weston 1933, Clinton 1936, Horsfall 1963, Lamb 1976).
Charles Edwin Bessey (1845–1915), son of an Ohio teacher, received his B.S. degree from Michigan
Agricultural College in 1869 and in 1870 became professor of natural history at Iowa State College of
Agriculture at Ames, to teach botany and horticulture (Pool 1915, Ewan 1970, Overfield 1975, 1992,
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Fig. 16. (a) Thomas Jonathan Burrill. Ainsworth 1981:65. (b) Erwin Frink Smith. Rodgers
1952:frontispiece.
Campbell et al. 1999:98–105, Shor 1999). He spent two winter vacations of three months each studying
botany under Asa Gray at Harvard and received an M.S. degree from his alma mater in 1872. Iowa
State College granted him a Ph.D. degree in 1879 based upon his botanical publications. In 1884 he
became professor of botany at the University of Nebraska in Lincoln, where he remained for the rest
of his career. Bessey developed strong ties with both the botanical profession and investigators of plant
diseases. He followed the teachings of de Bray and began publishing on parasitic fungi in 1877, and
continued doing so through 1903 (Pool 1915:513–516). However, he took all botany as his domain, and
he dominated botany in America by editing journals, publishing scientific papers and botany textbooks
for high schools and colleges, and by training other botanists. One of his three sons, Ernst Athearn
Bessey, became an internationally known mycologist (Bessey 1955).
Davaine’s reports of a bacterial plant disease were clear and succinct, but went unnoticed in
Germany and the United States. Thomas Jonathan Burrill (1839–1916), a farmer’s son, who was born
in Massachusetts and grew up in Michigan, graduated from the State University of Illinois in 1865,
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Contributions
age 26 (Smocovitis 1999). In 1867 he was botanist for John Wesley Powell’s first expedition to the
Colorado Rocky Mountains (Ewan and Ewan 1981:33), and in 1868 he began teaching at Illinois
Industrial University (in 1886 renamed University of Illinois) and soon became professor of botany and
horticulture. By 1874, he introduced plant pathology into his courses and his publications. Fire blight,
as noted above, was known to be contagious by the 1840s but assumed to be caused by a fungus. That
was still being assumed by Burrill in his first two papers on fire blight in 1877 and 1878, but in 1879 he
cautiously suggested that the bacteria he found in infected plant fluid caused the disease (Clark 1961:249–
250, Baker 1971:614–615, Ainsworth 1981:64–65, Campbell et al. 1999:107–116). In 1882, he named
the bacterium Micrococcus amylovorus (now Erwinia amylovora). Others confirmed his results (Matta
2007:198–199). For all this and much more, including 17 articles listed in their bibliography, Campbell
et al. (1999:109, 376–377) call Burrill “father of American plant pathology.”
Bacteriology was pursued for its relevance to both plant and animal diseases, and both pathologies
benefited from that. For example, Dr. George M. Sternberg (1838–1915), Surgeon-General of the U.S.
Army, published his own translation of a French textbook of bacteriology by Antoine Magnin (1880),
the first such textbook published in America (Clark 1961:51–52). He published an enlarged edition in
1884, and in 1892 published his own Manual of Bacteriology.
Burrill’s contributions to phytopathology were soon eclipsed by his younger contemporary, Erwin
Frink Smith (1854–1927). Smith grew up in Gilberts Mills, New York, near Syracuse, son of a tanner
and shoemaker. He had an early interest in fishing and in ants and was an avid reader (Rodgers 1952:1–5,
Clark 1961:228–232, Aycock 1975, Ainsworth 1981:66–70, Wolf 1999, Matta 2007:202–205). In 1870
his father bought a farm near Hubbardston, Michigan. In that town he met the druggist and postmaster
Charles F. Wheeler, who was interested in botany. Smith did not graduate from high school until 1881,
and in the same year he was junior coauthor with Wheeler of The Flora of Michigan, which described
1634 species (True 1927, Rodgers 1952:18, Campbell 1983:23). He graduated from the University of
Michigan in 1886, and in 1889 received his Ph.D., with a dissertation on the peach yellow disease. He
began work in the USDA Section of Mycology in 1886 and remained there for the rest of his career.
He became the world’s leading authority on bacterial plant diseases, though most of his renown came
during the 1900s, beyond the limits of this survey. In 1897–1901 he engaged in a well-known debate
with German botanist Alfred Fischer (1858–1913) about whether there were any bacterial diseases
in plants. Fischer had studied under Sachs and de Bary and was more physiologist than pathologist
(Matta 2007:207–212). That debate is now reprinted in English (Campbell 1981). Fischer might have
suspected he was skating on thin ice when he acknowledged in 1897 that nonpathological bacteria occur
in the roots of Leguminosae (Campbell 1981:1). Voronin had published his discovery of this in 1866
(Baker 1971:613, 633). Despite that, Fischer claimed that pathological bacteria could not penetrate plant
epidermal cell walls. Dismissively, he noted: “new descriptions of plant diseases caused by bacteria
keep springing up and, truly, what worthless descriptions and what non-critical trials.” Smith, in the
Centralblatt für Bakteriololgie, replied in kind: “It is seldom in a genuinely scientific book that one finds
so many unwarranted assumptions and serious misstatements in the space of a single page” (Campbell
1981:4) as in Fischer’s textbook on bacteria. Smith reported that eight plant diseases were attributed to
bacteria, and that six of them were established beyond a reasonable doubt; he furnished bibliographic
citations to the literature on those six (Campbell 1981:7–8). Fischer responded in the same journal to
Smith’s six-page account with a seven-page retort, in which he explained why he could not accept the
328
conclusions in the literature cited by Smith (Campbell 1981:9–16). Fischer no doubt hoped that he had
ended the debate, but Smith would have the last words, in a 34-page + illustrations response that the
Centralblatt für Bakteriologie published in 4 parts (1899–1901, cited from Campbell 1981:17–51 + 11
plates).
The Society of American Bacteriologists, covering the United States and Canada, barely got organized
during the 1800s—its organizational meeting was held on 27–29 December 1899 (McClung 1978).
Other countries
The study of phytopathology beyond Europe and North America was also important during the 1800s.
What I have found about it may not be exhaustive.
Australia was in the 1800s a British colony, and some of its immigrants were educated in Britain or
Europe. Wheat rust was a disease that struck wheat crops 10 times, 1799–1889 (Fish 1970:13). In 1864,
the Victoria Board of Agriculture appointed a committee, chaired by Government Botanist Ferdinand
von Mueller, to find the cause and prevention of rust disease. The committee recommended “early
sowing careful selection of wheat varieties as a means of control” (Fish 1970:13). In 1898, William J.
Farrer, a Cambridge University graduate who had immigrated to Australia in 1869, was botanist in the
New South Wales Department of Agriculture. He researched production of rust-resistant wheat varieties,
being advised by A. E. Blount, Colorado Agricultural Experiment Station. “The influence of Farrer’s
varieties was so great that the area sown to them rapidly increased as the overall production of wheat in
Australia expanded” (Fish 1970:14, quoting I. A. Watson, 1958). Scotsman Daniel McAlpine had studied
under Thomas H. Huxley and William T. Thistleton-Dyer at the Royal College of Mines, London. He
immigrated in 1884 and became lecturer in biology at the University of Melbourne (Fish 1970:14). In
1889 a rust epidemic led to McAlpine being appointed in 1890 as Consulting Vegetable Pathologist to the
Department of Agriculture, Victoria. He wrote a treatise on Australian fungi in 1895 and a monograph on
Australian rusts in 1906. He became known as “father of Plant Pathology in Australia” (Fish 1970:15).
An American, Nathan Augustus Cobb, had obtained a Ph.D. in Germany in helminthology and went
to Australia in 1889 and became Consulting Pathologist for New South Wales. While in Australia he
wrote about a dozen scientific papers on nematodes, and his 1895 demonstration that gumming disease
of sugarcane was caused by a bacterium impressed Erwin F. Smith, who had the USDA employ him in
1905. Cobb became “father of American plant nematology” (Buhrer 1969).
In Brazil, phytopathology began with the French naturalist–collector Augustin (Auguste de) SaintHilaire (1779–1853), who traveled there in 1816–1822 (Guerra 1975). He described wheat rust in several
of his writings, which Anna Jenkins quoted in French (1945). Jenkins, from the USDA, was a long-time
collaborator with A. A. Bitancourt at the Biological Institute of Saõ Paulo in phytopathological studies,
beginning in 1934 (Bitancourt 1978:12–13).
In Japan, descriptions of diseased plants can be traced back to 713 and later, but there was no
understanding of cause or remedy (Akai 1974:13–14). In 1649, the government prohibited foreigners
from entering Japan or Japanese from leaving, except that Dutch ships were admitted to one port once
a year. That situation ended in 1866, when a United States warship demanded its end. Afterwards, the
329
Contributions
government was anxious for Japanese scholars to be brought up to date in western science. Japanese
scholars studied in American and some European universities, and western professors taught for one
or more years apiece in Japanese universities. The earliest teacher on phytopathology was a German
professor from Brandenburg, Friedrich M. Hilgendorf, in 1873–1876 (Akai 1974:14). Shinnosuke
Matsubara, a professor at the Tokyo Imperial University, published the first textbook on botany based
upon western science in 1882, and it had a chapter on plant pathology. For the rest of the 1800s, Japanese
and foreign botanists described plant diseases in Japan and attempted to discover the causal fungi (Akai
1974:15).
Whatever is published about history of New Zealand phytopathology is unknown to me. E. H. C.
McKenzie was located at a government Plant Diseases Division when he wrote “Mycological History
and Exploration in New Zealand” (1983). Perhaps he also published on history of phytopathology.
Conclusions
During the 1800s, phytopathology made enormous progress. It benefited from progress made in plant
physiology—it helps in understanding diseased plants to first understand normal plant functioning.
Progress in mycology, bacteriology, virology, and nematology were also essential contributions from
related sciences. Many parasitic fungi turned out to be rather simple multicellular organisms that have
complex life histories. As knowledge of plant diseases increased, governments responded by supporting
agricultural research institutions and university instruction in plant diseases. In 1862, the United States
created both land grant colleges that taught agricultural sciences, and a Department of Agriculture.
Both creations grew to become centers for world leaders in phytopathology. Perhaps western European
countries still remained in the forefront of research by 1900, but if so, they would suffer a setback during
World War I that the United States did not experience, and by then the United States was leading the
world in phytopathology because of better research funding and more facilities.
In retrospect, one sees many steps toward a germ theory of disease in this story, which, as in the
1700s, were little appreciated at the time.
Literature guide
There are ample historical sources on the phytopathological literature of the 1800s. Excellent places
to begin are Geoffrey Ainsworth’s fine Introduction to the History of Mycology (1976) and Introduction
to the History of Plant Pathology (1981). However, earlier works provide additional details. Herbert
Whetzel’s Outlines of the History of Phytopathology (1918:32–106, 1977) has a good overview
and extensive references; Ernest Large’s classic Advance of the Fungi (1940, 1962) is well written,
illustrated, and documented. Narrower in scope and briefer is G. McNew’s “The Ever Expanding
Concepts behind 75 Years of Plant Pathology” (1963); Gert Orlob’s “Concepts of etiology in the history
of plant pathology” (1964:220–268); Garnet Carefoot and Edgar Sprott’s Famine on the Wind: Plant
Diseases and Human History (1967) is briefer than Large’s history and less well illustrated, but more
recent, with a good bibliography; George Ordish’s The Constant Pest: A Short History of Pests and
Their Control (1976) covers both fungi and insects, since the Neolithic era; and James Horsfall and Ellis
Cowling’s “The Sociology of Plant Pathology” (1977) is a history of phytopathologists in America,
with a three-page list of “Hall of Fame” pathologists’ achievements. Chronologies of phytopathology
330
by Karl Mayer (1959:18–21) and Keith Parris (1968:21–50) provide concise international panoramas
not found in the histories. The history of viral plant diseases is well covered in A. P. Waterson and
Lise Wilkinson’s Introduction to the History of Virology (1978). Histories of botany, such as Morton’s
(1981), place phytopathology within a broader context for the 1800s. There are also national histories of
phytopathology, including Anna Jenkins (1941, 1945) on Brazil, G. Viennot-Bourgin (1954) on France,
Ainsworth (1969) on Great Britain, S. Fish (1970) on Australia, Shigeyasu Akai (1974) on Japan, Don
Ellis (1976) on North Carolina, and Lee Campbell, Paul Peterson, and Clay Griffith (1999) on the
United States. John Schlebecker’s Bibliography of books and pamphlets on the history of agriculture
in the United States, 1607–1967 (1969) is also helpful. Histories of mycology are valuable resources,
including M. Chadefaud’s for France (1954), Ernst Bessey’s for the years 1853–1953 (1955), Giacomo
Lazzari’s for Italy (1973), McKenzie (1983) on New Zealand, and Donald Rogers’ for North America
(1981:7–27). The American Phytopathological Society during the 1900s reprinted works in English or
published English translations of foreign-language “Phytopathological Classics” from the 1800s, some
of which I collected and reprinted (Egerton 1977).
Phytopathology textbooks often have historical introductions (including Heald 1926:7–44, Walker
1969:14–46, Agrios 2005:8–28). Large (1940) and Ainsworth (1981) tell the interesting story of
phytopathology’s impact on agriculture. Raymond Doetsch’s Microbiology: historical contributions
from 1776 to 1908 (1960) is a sourcebook of fifteen authors.
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Acknowledgments
For their assistance I thank Professors Stuart McCook, Department of History, University of Guelph,
Guelph, Ontario and Steven Turner, Department of History, University of New Brunswick, Fredericton,
New Brunswick, Canada, and Dr. Jean-Marc Drouin, Muséum National d’Histoire Nationelle, Paris,
France.
339

History of Ecological Sciences, Part 44

Transcription

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