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| UNITED STATEB “~~
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DEPARTMENT or AGRICULTUR

DEPARTMENT CIRCULAR J. Bepeparement

— 44+—
——= od

| Washington, D. C. Revised November, 1928

_

A METHOD FOR THE CONTROL OF CROWN:
GALL IN THE APPLE NURSERY

M. B. WAITE

Principal Pathologist in Charge

and

E. A. SIEGLER

Associate Pathologist, Office of Fruit Diseases, Bureau of Plant Industry

CONTENTS
Page Page
Perch = 2 ee = i>) Discussion) == ee es A Ee ee 5
Experiments and results________--_-_~ 2 | Recommendations for control]_________ 6
INTRODUCTION

The problem of crown gall in the root-grafted apple nursery is a
very serious one. The nurseryman who has to discard only 5 to 10
per cent of his trees on careful inspection at digging time is usually
considered fortunate. Losses of 25 to 50 per cent are not uncommon,
and certain lots of trees in extreme cases have even shown a loss
of 95 per cent. ‘These serious losses to nurserymen and also the
general interest of orchardists in obtaining disease-free stock for
planting have made the writers feel the importance of this phase
of the crown-gall problem. Present-day nursery stock, especially
when grown by the root-grafting method, appears to contain a
higher percentage of infection than formerly, and there also appears
to be more trouble in recently planted apple orchards than in those
planted a generation ago. Whether this supposition is true or not
may be open to question. It may be that nursery and orchard trees
are examined more critically as increased attention is given to
orchard diseases.

Before the bacterial cause of this disease was known George G.
Hedgcock, in his extensive studies of crown gall. tried many experi-
ments in the propagation of root-grafted apple trees and made many
observations which resulted in progress toward the control of this
disease. The discovery of the bacterial nature of this previously
obscure disease by Erwin F. Smith and his associates in 1907 marked

15586°—28

RECEIVED |

of Agridult
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2 CIRCULAR 376, U. S. DEPARTMENT OF AGRICULTURE

a real epoch in the history of crown gall and appeared to open possi-
bilities of finding satisfactory and practical remedies for the trouble.
Nevertheless, in so far as the control of crown gall in the apple
nursery is concerned, no very extensive practical applications of
these important discoveries have as yet materialized.

This circular is published for the purpose of making immediately
available to nurserymen and others interested in the propagation
and growing of root-grafted apple trees in nurseries a new method
of greatly reducing the attacks of crown gall. This preliminary
report, to be followed later by a technical paper containing full
details, also discusses briefly certain problems concerning the diag-
nosis of crown gall. It is hoped that the results of these experiments
will tend to answer some
of the questions concern-
ing the identification of
this disease.

EXPERIMENTS AND
RESULTS

As early as 1909 the
senior writer used a for-
maldehyde solution as a
dip for apple stocks and
scions before grafting, as
a means of controlling
crown gall. Observations
always indicated the prac-
ticability of this method,
and nurserymen accord-
ingly have been advised in
correspondence as to this
treatment for at least a
decade. The need of
more exact knowledge

Fic. 1.—Typical trees classified as clean was so apparent that in
1921 experiments were

initiated in an attempt to obtain more definite information on this
phase as well as on related phases of the subject. This investigation,
covering a period of five years, appears to confirm conclusively the
observations of the writers extending over a much longer period. In
these experiments this formaldehyde-solution treatment with modifi-
cations has consistently proved of value in controlling the crown-gall
disease on apple grafts. For the past three years one of the organic-
mercury compounds has also been tested as a dip and has proved dis-
tinctly more efficacious than the formaldehyde-solution treatment.
Mercury compounds are among the most powerful of germicides, but
the inorganic compounds, such as corrosive sublimate, are often very
injurious to plant life, even in extremely dilute solutions. A number
of new organic-mercury compounds have recently been put on the
market and have proved of value in the control of other plant diseases.
This particular one was tried in the hope that it would be effective
in controlling crown gall and would be noninjurious to the apple

°

METHOD FOR THE CONTROL OF CROWN GALL 3

grafts. This proved to be the case. In a number of experiments
performed in 1925 complete confirmation of results of experiments
initiated in 1921 with the formaldehyde solution and in 1923 with the
organic-mercury preparation has been obtained. Im practically all
of these experiments the grafts were wrapped either with raffia or
with a cheap grade of muslin in such a manner as to cover completely
the graft unions. The trees were dug as 1-year grafts. Neither the
stand nor the amount of growth was reduced by the use of the or-
ganic-mercury compound as compared with untreated grafts grown
as checks. The trees were classified into three grades: (1) Galled
trees. with further subdivision into those with large and those with
small galls; (2) doubtful trees, that is, trees exhibiting symptoms so
slight as to be open
to question or differ-
ence of opinion; and
(3) clean trees, that
is, those trees which
showed no symptoms
of gall or hairy-root.
It should be stated
that practically all of
the so-called doubtful
trees would have been
classified as clean
trees in an ordinary
inspection. The ac-
companying illustra-
tions were made from
photographs of
trees grown in the
experiments of 1925.
Figure 1 illustrates
examples of clean
trees. Figure 2 illus-
trates the type of
tree which has been
classified as possess-
ing a small gall, Fic. 2.—Type of tree classified as possessing a small gall
whereas Figure 3

illustrates two types of trees classified as having large galls, which
have invariably encircled more than 40 per cent of the circumference
of the tree. Figure 4 represents one of the types of trees classified
as doubtful.

TABLE 1.—Comparison of results obtained with all grafts treated with the
organic-mercury compound and untreated grafts grown as checks

Treated grafts | Untreated grafts
| Percentage of galls Percentage of galls
Number = = eae Number |
Large | Total | Large Total
| eases( |
Ei) be Sal een rite Re ae ie 8 2.0 | OG ce 0a ae a ee 28. 7 32. 6

A CIRCULAR 376, U. S. DEPARTMENT OF AGRICULTURE

Fic. 3.—Typical trees classified as possessing large galls, one (left) showing the
woolly-knot type and the other a simple form of crown gall

In the experiments of
1925 the organic-mercury
treatment was given to 2,619
grafts, representing a num-
ber of different varieties,
with the result that the
total number of trees ob-
tained at digging time with
both large and small galls
amounted to 6.1 per cent,
as compared with 32.6 per
cent on 2,619 untreated
grafts of the same varieties
which were used as checks.
The proportion of the total
number of large galls in the
treated grafts of all varie-
ties was 2 per cent and in
the untreated or check
grafts 28.7 per cent, as
shown in Table 1.

Figure 5 represents a typ1-
cal lot of trees resulting
from the use of the organic-
mercury compound. ‘This
lot was selected because it
illustrates about the average
Fig. 4.—One of the types of trees classified as control obta ined in the ex-

of doubetal periments in 1925, whereas

ce

METHOD FOR THE CONTROL OF CROWN GALL 5

Figure 6 shows the corresponding lot of trees resulting from un-
treated grafts which were used as checks. The variety used in each
case was Summer Rambo. In both illustrations the galled trees are
on the left, the doubtful trees in the middle, and the clean trees on
the right. Figure 7 shows a lot of Ben Davis trees selected as a
sample of some of the best results obtained from grafts treated with
the mercury compound, and Figure 8 shows the corresponding
lot of Ben Davis trees resulting from untreated grafts used as checks.
Reading from left to right, in both illustrations, it will be seen
that there were no galled trees grown from the treated grafts, as
compared with 48 ‘oalled
trees in the check lot.
This, of course, represents
an example of absolute con-
trol that has not been met
with very often in the ex-
periments.

DISCUSSION

Recently Riker and
Keitt* have advanced the
working hypothesis that
“the malformations dealt
with in the rejected nur-
sery trees were not induced
by the crown-gall organ-
ism.” This hypothesis * was
formulated because of their
negative results in attempts
to isolate the organism
from 175 trees which had
been rejected on account
of malformations suppos-
edly due to crown gall.
Since apparently similar
malformations were not fic. 5—One of the lots of trees which after

onlv controlled in size but classifying, represented approximately the aver-
“ age of results obtained with the organic mercury-
also were largely prevented compound treatment. Variety, Summer Rambo.

— - - Reading from left to right: Galled t
from forming by the action, ee. ee ti ed trees (5)

of germicides in the writ-

ers’ experiments, their results are not in accord with the hypothesis
of Riker and Keitt. It is difficult to assume that these malformations
are due to any agency other than a pathogenic organism, since they
are so definitely prevented by germicides.

Referring to Riker and Keitt’s statement that malformations occur
on graits treated as aseptically as possible and “ that gall-like forma-
tions may develop without the intervention of Bacterium tumefaciens,”
attention is called to the fact that Hedgcock ? performed almost iden-
tical experiments more than 15 years ago and arrived at the following

1 Riker, A. J., and Keitt, G. W. Crowngall in relation to nursery stock. Jn Science,
vol. 62, p. 184-185. 1925.

2 Hedgcock, G. G. Field studies of the crown- gall and hairy-root of the apple tree.
U. S. Dept. Agr., Bur. Plant Indus. Bul. 186, 108 pp., illus. 1910.

6 CIRCULAR 376, U. S. DEPARTMENT OF AGRICULTURE

significant conclusions: “ The two regions where the formation of
callus in root grafts was most abundant * * * coincide exactly
with the position of more than 90 per cent of the galls, * *
indicating a close relation between the formation of callus and the
development of the disease.” All of the writers’ observations and
experiments furnish additional evidence to support these conclusions
of Hedgcock.

It should also be noted that the failure to find a parasitic organism
or agency in diseased tissue at any given time does not necessarily
justify the conclusion that the organism or agency is not present or
that it has not been present at some previous time. Until conclusive

evidence is brought forth proving that malformations “ supposedly
crown gall” are not due
to the strongly pathogenic
organism Bacterium tume-
taciens Sm. and Tn., the
present rigid inspection and
rejection of crown-gall trees
should be maintained. It is
considered that this partic-
ular question is of vital im-
portance to the orchardist.

RECOMMENDATIONS FOR
CONTROL

Seedling apple trees of
l-year’s growth ordinarily
show a certain amount of
hairy root and crown gall.
These seedling trees with
either type of the disease
when lined out for budding
or when used for grafting
continue to carry the dis-
ease, in the great majority
of cases at least. The expe-
Fig. 6.—Lot of untreated Summer Rambo trees rience of the writers is that

gown ne, checks, #05 thoes aie agers 3 ete oD a: eee a

doubt: (ClAyicleaeet aa * inspect the seedling stocks

rather critically. Only an
occasional gall and a few hairy-root trees may escape their scrutiny.
However, when these diseased stocks are planted out or cut up into
pieces for piece-root grafting, the nurseryman is simply propagating
the disease. The best practice, therefore, is to inspect critically the
seedling stocks before using them for orafting and to burn all the
obviously infected ones. It has been generally accepted by both
pathologists and nurserymen, at least since the work of Hedgcock,
that the selection of roots free from crown- -gall and hairy-root ‘infec-
tion is an important factor in the control of this disease.

Since this disease is caused by a germ easily killed by germicides
when it can be reached and since this germ may be present over the
surface of the roots, the problem of killing the organisms present
on the surface and of disinfecting the material resolves itself into

~ tO ETS e a ae e r

METHOD FOR THE CONTROL OF CROWN GALL Z

the question of finding a suitable germicide or fungicide and a
suitable dosage which will kill the bacteria and not materially injure
the roots of the trees. In this respect the organic-mercury compound,
used as a dip in these experiments, represents a distinct improvement
over the formaldehyde solution.

Well-made grafts properly fitted, especially those which have
no overhang at the lower end, were shown by Hedgecock to be less
susceptible to crown gall. They have, in fact, less exposed callus.
The wrapping, of course, should be thoroughly done, whatever
method is selected. Since the material has to be handled so much
and since the freshly made grafts will stand this disinfectant, part
of the writers’ experiments also included dipping® the grafts the
same day they were made,
in order to disinfect and
saturate the wrappers with
the germicide and to give
a second coating to the
grafts themselves.

Since there is still a
possibility, after the grafts
have been stored and the
young tender callus devel-
oped, that this may carry
incipient infections, and as
it is desirable to give an-
other dose of the germi-
cide just before planting.
a third dipping * was tried
and found to be noninju-
rious to the graft and help-
ful in the control. Melhus
and Maney* also report
the use of Bordeaux mix-

ture as a dip for apple
grafts. Fic. 7.—Lot of Ben Davis trees selected as showing
A Aaa some of the best results obtained in controlling
There 1S; 5 possibility crown gall by the use of the organic-mercury com-
that the grafting knives or 3002, ,ReMding from left to right Galled, trees
other tools and the benches
and tables may become infected. It is not probable that all obscure
cases will be eliminated in the inspection, and dormant infection
would, of course, escape notice. Furthermore, the knives may cut
through bacterial infection beneath the thin outer bark which was
not reached by the disinfectant. Although there is no definite proof
that stock can be contaminated with crown gall by means of grafting
knives, there is proof that the pear-blight organism can be carried
on grafting and pruning knives. In view of the fact that it is rather
easy occasionally to disinfect the tools and grafting benches, it is
thought good practice to utilize this method of control, even though

its value is not fully proved.

2Later experiments in a repetition of these treatments have tended to minimize the
importance of the second and third dipping and to increase the importance of the first
dipping.

t Melbus, I. E., and Maney, T. J. A study of the control of crown-gall on apple grafts
in the nursery. Iowa Agr, Exp. Sta. Research Bul. 69, pp. 159-172. 1921.

8 CIRCULAR 376, U. S. DEPARTMENT OF AGRICULTURE |

The following treatment of apple grafts for the control of crown
gall is recommended as a result of a large number of experiments @
over a number of years:

(1) Select seedlings free from hairy-root and crown-gall infection.

(2) After washing, dip for
10 minutes uncut seedlings
and scions in a solution con- a
sisting of 1 part of hydroxy- |
mercurichlorophenol * to 400
parts of water (approxi-
mately at the rate of 1 ounce
to 3 gallons). In this as
well as in subsequent dips
do not rinse with water, and
keep the solution in either a
wooden or a nonmetallic
container. The uncut seed-
lings and scions should be
allowed to become surface
dried before cutting for the
grafting operation.

(3) Dip the grafts, which
should be well fitted and
carefully wrapped, in this
same freshly made solution

x 6
Fic. 8.—Trees from untreated Ben Davis grafts for about 2 seconds.

grown ag coe for en ot wus brated in ae (4) Store erafts under
ure 7. eading from le o right: Galle rees “78 :
(48), doubtful (4), clean (61) cool conditions and dip

them for about 5 seconds
in a freshly made solution of the mercury compound immediately
before planting.°®
(5) Keep the bench on which the grafting is done, as well as all
grafting tools, disinfected by frequent washings with a germicide.

5 Obtainable on the market in powdered form under the trade name “ Semesan.”

6 Later experiments have shown excellent control with the first dipping alone and very
little additional benefit from the second and third dippings. Since there has been found to
be some danger of injury, under unfavorable conditions of storing and growing the grafts,
by the second and third dippings here suggested, it is possible to eliminate these addi-
tional dippings without appreciable loss of control.

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OF THIS PUBLICATION MAY BE PROCURED FROM
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AT
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