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2) USDARIMENTOPAGRCULIRE ©

No. 203

Contribution from the Bureau of Plant Industry, Wm. A. Taylor, Chief.
April 30, 1915.

(PROFESSIONAL PAPER.)

FIELD STUDIES OF THE CROWN-GALL OF
SUGAR BEETS.

By C. O. TowNnsEND,
Pathologist in Charge of Sugar-Beet Investigations.

KINDS OF BEET GALLS.

There are at least two distinct but clearly related kinds of growths
occurring upon sugar beets which may be considered under the name
of galls. These have been designated as ‘‘tumors’’ and ‘‘tubercu-
losis’’ in Bulletin No. 213 of the Bureau of Plant Industry.t| While
these two kinds of outgrowths are similar in external appearance,
especially in their early stages of development, their internal ap-
pearance and their subsequent behavior serve to distinguish the
tumor from tuberculosis. Internally, the outgrowths known as
tuberculosis of the beet show small, brownish, water-soaked areas,
as mentioned on page 194 of the bulletin cited, while the tumor is
free from these discolored areas. Externally, both kinds of galls
are usually smooth at first, but the tuberculosis galls eventually
become decidedly rough, cracked, and very dark, and finally decay.
This decay of the galls often causes the beet itself to rot, thereby
entailing more or less loss on the grower, according to the prevalence
of the disease. On the other hand, the tumor remains comparatively
smooth, seldom cracks, does not usually decay, and frequently
retains its firmness until the beets are harvested. The quality of the
galls and their effect upon the beets from which they arise, as given
in this paper, relate for the most part to the tumor variety.

DISTRIBUTION OF BEET GALLS.

The abnormal outgrowths known in this country as crown galls
have been observed upon beet plants from time to time for more than
50 years. Indeed, as early as 1839 attention was called to these

1 Smith, Erwin F., Brown, Nellie A.,and Townsend, C.O. Crown-gall of plants: Its cause and remedy.
U.S. Dept. Agr., Bur. Plant Indus. Bul. 213, p. 105, 194. 1911.

82778°—Bull. 203—15

2 BULLETIN 203, U. S. DEPARTMENT OF AGRICULTURE.

peculiar growths upon beet roots, which were spoken of at that
time as warts. In 1859 some of these galls were described as larger
than the beet roots themselves and were looked upon as curiosities
and monstrosities. It appears from a study of the literature upon
the subject and from observations in the field from year to year
that this disease of the sugar beet has increased rapidly in recent
years and that it is still on the increase. Its presence has been re-
corded in many of the beet-growing countries of Europe, and in our
own country it has been found on the sugar beet from Virginia to
California. (Fig. 1.) In many localities where only a small number
of cases were observed a few years
ago, there are now hundreds and
sometimes thousands of galled beets
each year, especially if beets have
been followed by beets for several
years in the same field. On the
beet itself the galls may appear at
any point from the top of the crown
Fic. 1—Map of the United States, the shaded to the extreme up of the root. How-
portions showing the areas where sugar-beet e@Ve€TI, by far the largest number of
galls have been observed. galls are to be found at or near
the surface of the ground, and for this reason these growths have been
termed crown galls.

APPEARANCE OF BEET GALLS.

The outgrowths, or galls, on the beet do not usually appear until
the beets are from one-fourth to one-half grown; that is, until mid-
summer. From that time on, they may appear at any time until the
beets are harvested. Consequently, we may find at harvest time
galls in all stages of development, from tiny protuberances that have
just begun to grow to what might be called the full-grown gall,
several inches in diameter, as shown in Plate I, A to K. Frequently,
these galls push out from the surface of the beet without any distinct
line of demarcation between the gall and the beet.proper, as seen in
Plate I1, B. In other cases the outgrowths are attached to the beet
by very slender necks or threads, and between these extremes may be
seen the full range of variation in relative size of the connecting
tissue. Whether large or small, the connections are short, so that
the gall almost invariably lies close to the beet from which it springs.
Sometimes but a single gall is produced on one beet, as shown in ~
Plate I, / to K, while in other instances several or many galls may
develop on the same beet, as illustrated in Plate II, A, B, C, and F.
In the latter case the galls may be distinct and separate (Pl. II, A) or
they may occur in groups (Pl. II, B). It is not uncommon for the
entire crown of the beet to be covered with a mass of galls (Pl. I, £).

FIELD STUDIES OF THE CROWN-GALL OF SUGAR BEETS. 3

Usually the galls with slender attachments occur singly, although there
may be several on the same beet, while the galls occurring in groups
usually have broad bases without any distinct lne between the gall
and the beet. In the early stages of development—that is, when the
galls are young—their surfaces are bright, resembling the surface of
the beet proper and indicating active growth; but as the galls grow
older they become darker, especially if they are above the surface of
the ground. In this way their relative ages may be easily determined.
When galls have begun to form they usually increase in size most
rapidly on those beets that are making the most rapid growth.

CAUSE OF BEET GALLS.

The primary cause of the formation of crown galls on the sugar
beet and many other plants was for a long time in doubt. Few plant
diseases have given rise to more extended investigations than has
the so-called crown-gall. Different investigators have assigned the
origin of these abnormal growths to a great variety of causes, ranging
from slime molds to mechanical injuries. However, the investiga-
tions set forth in Bulletins Nos. 213 and 255 of the Bureau of Plant
Industry! prove conclusively that a bacterium or several closely
related bacteria are responsible for the origin and development of
these outgrowths belonging to the class of so-called crown galls.
The organism producing ‘‘tumors” is known as bactervum tume-
faciens (Smith and Townsend) and.the one producing ‘‘tuberculosis”’
is designated as B. beticolum (Smith)?

The most extensive work on mechanical injuries as the cause of
gall formations on sugar beets has been carried on by Spisar.* There
seems to be no proof, however, that the organism which is capable
of producing galls on sugar beets was not present in the fields in which
Spisar carried on his experiments. It is apparent that a mechanical
injury offers a favorable place for the organism to enter the plant,
yet the indications are that gall formations will not result from
mechanical injuries unless the gall-producing organism is present.
In the field studies on the crown-gall of beets carried on by the writer
for several years, it has been frequently noted that when galls begin
to appear on the beets in a given field they are at first few in number,
increasing from year to year if beets continue to be grown in that
field. It has also been noticed that if badly infested fields are
followed one or two years with a grain crop and then returned to

1 Smith, Erwin F., Brown, Nellie A., and Townsend, C. O., op. cit.

Smith, Erwin F., Brown, Nellie A., and McCulloch, Lucia. Thestructure and development of crown-
gall: A plant cancer. U.S. Dept. Agr., Bur. Plant Indus. Bul. 255, 60 p., 2 fig., 109 pl. 1912.

2 These organisms are described in Bureau of Plant Industry Bulletin 213, which may be obtained from
the Superintendent of Documents, Government Printing Office, for 40 cents, 10 cents additional being
required for postage to foreign countries.

3 Spisar, Karl. Uber die Bildung des Zuckerriiben-Kropfes. Jn Ztschr. Zuckerindus. Béhmen, Jabrg.
36, Heft 1, p. 1-17, fig. 1-6; Heft 2, p. 57-72, fig. 7-11. 1911.

4 BULLETIN 203, U. S. DEPARTMENT OF AGRICULTURE.

beets, the galls are greatly reduced in number after one year in grain
and practically eliminated after two years. It is true that when galls
are present in a field of beets they are frequently more numerous
near the ends of the rows, where the greatest amount of mechanical
injury is produced by the horses and cultivator in turning. (See
Pl. Il, HE and F.) However, adjacent fields and even parts of the
same field not previously in beets, but in which the beets at the time

of the observation were subjected to the same mechanical injuries, —
were free from galls (Pl. Il, D), regardless of the fact that the soil,
climatic conditions, and cultural methods were the same. While
certain plant galls and callus formations may be produced by other
agencies, all extensive laboratory, greenhouse, and field studies on the
crown-gall of sugar beets lead to the conclusion that in this country
the true crown-gall formations of this class are produced by bacteria.

QUALITY OF BEET GALLS.

In topping beets from which sugar is to be obtained, it is customary:
to cut off the crowns at the line of the lowest leaf scar. The reason
for rejecting the crowns, as generally known, depends on the fact
that as a rule they contain a high percentage of salts, which tend to
prevent the sugar from crystallizing in the mill. In the process of
topping the beets it frequently happens that a part or all of the galls
that occur on the beets are so located that they are left on the root
(Pl. I, F to K) and are, therefore, put through the mill. In order to
find out whether or not the galls might affect injuriously the juices
in the mill, a series of tests was made to determine the quality of the
galls as compared with the beet crowns and roots. The results of
these tests are given in Table I, the analytical work for this and the
succeeding table having been performed at the Garden City, Kans.,
aor con by Mr. C. A. Hauser.

Taste I.—Sugar tests of beets affected with crown-gall.

Average | solidsin | Sugar i
tae = : gar in | Coefficient | Sugar in
Part of beet tested. eee juice. juice. | of purity. | part tested.
Experiment 1: Ounces. Percent. | Percents>|\ Per cent. ”| Percent:
Peapienates fo Ne Ek tee I Ess | 3,2 15.32 9. 00 BRT 5.00
CID SAD Ss se Se Ae ee os oe Se De eae 13.6 14.10 10. 10 71. 63 8. 90
ERGs ch, As as eee Se Se 24.0 14.30 11.00 76. 92 10. 06
Experiment 2
CDI cee oe SE ee ie seo tee ets 4.0 15. 60 8.10 51.92 4.60
GRO WSte wen otis Se ea eee efone 2, 12.70 8. 20 64.56 7.50
LAGI 5 See ea ane ee eee a Sa ee 25.6 14.10 11.00 78. 01 10. 20
Experiment 3
CHIR Se SoC ae Dens Boe ease ne ee 352 16. 00 9. 20 57.50 7.10
CROWES SS peal cee ee ee Be eee 13.6 15. 40 10. 80 70. 20 10. 20
LE TORIES ee No Ss Rap ee Sree ar 29.8 15. 00 11. 40 76.00 10.70
Experiment 4
ACT Sipser: im reer NC PME de es 5.6 15.30 7.80 ' §0. 98 7.00
SPO WTS ee eee oe ee eer iors 18.4 13.10 9.00 68. 70 8. 40
POOLS se ee ec or fate ee nett so 32.8 13.10 10.10 77.09 10.00
Experiment 5
Galls= ye se sce eee tee eee tee 8.0 16. 60 9.10 54. 81 5.90
LOW TISS ee eras Ne Soo eee oe eee re 18.4 14. 60 9.90 67. 80 8.50
EU OO ES Been ae oes wishes bE Se 44.8 14.10 10.50 74. 46 9.00

Bul. 203, U. S. Dept. of Agriculture.

i. - Pe | a
, i. 3)
es "mg
Zink ee oe Ra aoe
Wey a i Eee: en Lhe gene, i

"4 Por) AE —_
4 + A,
Sigs ‘te Wye TF CAT Mai Die eet cit ed

GALLED SUGAR BEETS, SHOWING THE VARIOUS LOCATIONS OF THE GALLS.

Bul. 203, U.S. Dept. of Agriculture. —. (PEATE III:

SUGAR BEETS, SHOWING GALLS APPEARING SINGLY AND IN GROUPS (A, B AND C) AND
THE RELATION OF INJURY TO GALL FORMATION (D, E, AND F).

FIELD STUDIES OF THE CROWN-GALL OF SUGAR BEETS. 5

TaBLE I.—Sugar tests of beets affected with crown-gall—Continued.

Average | golids in Sugar in | Coefficient | Sugar i
Ds rei é l Sugar in
Part of beet tested. parttested juice. juice. of purity. | part tested,
Experiment 6: Ounces. Percent. | Percent. Per cent. | Percent.
CO ER Ae ee en oe 3.2 17. 46 11.10 63. 57 10. 40
Crowns. =f 22: I oS se re io, Sits 'o ass Min o/s 8.0 18. 40 14. 40 78. 26 12. 50
PIOUS ERAS ee Oe ee 30. 4 19.56 16. 80 85. 88 14.00
Experiment 7
ONES Oe a 2.4 17.20 9.10 52.90 8.30
OG RNEE Ri 5, 9 See thuy Salter nae eee 7.2 18.76 15.40 82. 08 13.10
EROS ERG Fore ee ee Sa are esate sseie cle ie 22. 4 17.76 14.80 83.33 13.50
Averages:
CRUG Se SO ee 4.23 16. 21 9. 05 55. 60 6. 90
RIDOWGISME DR Serre Se on occ civle Sees 12. 91 15. 29 11.11 72. 03 9. 87
SOD S onc ce Soe 29. 97 15. 41 12. 23 78. 81 11. 06

In preparing the material for the analyses which form the basis of
this table badly galled beets were taken in groups of five in order to
get a sufficient quantity of juice from the galls to make purity as
well as sugar determinations of the galls themselves. After removing
the leaves only, the beets were thoroughly washed and weighed. The
galls were then carefully removed and the beets again weighed. The
crowns were then removed and the roots weighed. The three lots of
material from each group of five beets were prepared and put through
the test for sugar and purity as quickly as possible after the beets
were taken from the ground. A study of Table I shows that the
galls are decidedly lower in both sugar and purity than even the
crowns. It is evident, therefore, that any considerable number of
galls on the beet roots would be decidedly injurious to the sugar
recovery in the mill, since the large amount of salts in the galls, as
indicated by the low purity coefficient, would keep approximately
one and a half times as much sugar from crystallizing. Hence, it
would be advisable to remove any galls that are attached to the
beets below the crowns at the time of topping the beets.

Tt might not be out of place in this connection to call attention to
the high quality of the crowns in some cases, as shown especially in
experiment 7. It is possible that the salts were taken up by the
galls to the improvement of the crowns to some extent. On the
other hand, it is possible that the quality of the crowns might be
greatly improved by proper selection, so that the matter of crown
tare would not be such an important factor in handling factory beets
as it is at present.

EFFECTS OF GALLS UPON QUALITY AND SIZE OF ROOTS.

In an effort to get some definite information regarding the effect of
the galls upon the quality of the roots to which they are attached, a
series of comparative tests was made between galled beets and beets
free from galls. In selecting the beets for these tests a badly galled

6 BULLETIN 203, U. S. DEPARTMENT OF AGRICULTURE.

beet was taken as soon as it was loosened by means of the plow at
harvest time, and for comparison another beet was chosen, free from
galls, but as nearly the same size and shape as could be found growing
in the same row, close to the galled beet. After selecting in the
manner described the 26 pairs of beets which form the basis of
Table II, each beet was topped, washed, and the galls carefully
removed from the galled beet of each pair. The individual roots
were then tested for sugar and purity, with the results shown in
Table II.

TaBLe II.—Comparison in sugar content and purity of galled beets with beets not so

affected.
Niels |. Solidsin | Sugarin | Coefficient | Sugar in
Condition of beets. | juice. juice. of purity. | the beet.

Test No. 1: Per cent. Per cent. Per cent. Per cent.

CO EPULNG YO LS RE Gye ee AE eas at 13. 70 10..00 72.99 8.30

Noro al Su teers Se ane Pe ie Mates 2 2) a ele ee 18.10 13. 80 76. 24 12. 20
Test No. 2 :

(OPI 1o 2k & ok 28h ee Re ee A i Soe TP Bi itearet 13.90 10. 50 (seh 9. 20

INO Sheers ae es ER ot oe eee cee 18.00 13. 80 76. 66 12. 80
Test No. 3

Caled eaten we see Sek een oe 2 eee ee ee 17. 60 14. 50 82.38 11.70

ANOS Stee preteen a Ue Ten 2 Se ee a eee 21.92 18. 60 84.85 16. 80
Test No. 4:

OPAC De 2 ar eg CR cn ete ae eee aN er ee 15577 13. 20 83. 70 11. 20

NOP ROSS eee oe ai ele re See Se eens 21. 50 18. 70 86.97 16. 80
Test No. 5:

Galle eee Sie ew a eee Se eee ty oe Aen ae 15. 70 12. 80 81. 52 10.90

INO} ool Soe oe Oe ee ee ee oe ae i wae 20. 20 L710 84. 65 16. 80
Test No. 6

CE CULES Uh ie ete eee el re ee i rete ry 16. 30 13.10 80. 36 11. 40

NOP ISSe: Kee eee ee Seine ree a eee 20. 20 Gali) 84. 65 16. 80
Test No. 7:

CETL D0 | ie ale, Ne Ie ci lpn Ree a ale 16. 47 13. 20 80.14 11. 20

INO eee cae par See ace ep eee see wooue 20.17 16. 40 81. 30 15.10
Test No. 8:

GEG ee SE i, Sie oe Sane ee See mene hn pene ort a 10. 85 9.30 85. 71 8. 50

IN Fo (eas US ee eS ee ee ere ae et BE eerie 18.90 15. 60 82. 53 14. 00
Test No. 9:

(CoE aC ee ee a ee FO eS eee eee ee Aa 15. 27 12. 40 81. 20 11. 70

ENVOYS tS eee ee one ein noc nse eee eee eee 16.77 13.90 82. 83 13. 40
Test No. 40:

Galled ee = ies ee oie se oie inet Sainte een eee 16. 40 13. 80 84.14 13.10

No galls. ...- elec am ee ae ree 15.77 12. 80 81.16 -12.00
Test No. 11

Gallego oe tee ee eee tee eee 16. 23 12.10 74.55 11. 20

INFO yg WS Sook Se ae ee Se See nee sooner eGo Se AOS 22. 56 19.10 84. 66 18. 50
Test No. 12:

Gallego eae ae oe eee 18.77 14.00 74, 58 11.50

IN|) ALIS 5 ee ee ee ae =e eo eeasenatao sce 22.99 18. 20 79.13 17. 30
Test No. 13:

(GEN Gol es ie shea ree TA eee Doo 18. 20 14. 80 81.31 14.10

ING) SSE eee 5 ooo SaaS" sce moo sooo e Gee Enoee= sce 18.77 15. 00 Wos0%: 13. 80
Test No. 14

(GEA =/o cote Rees a Siecle ee See al rt eR ees = 17. 60 15. 40 87. 50 14. 30

J Su ee NE I a pease SS 19. 40 16. 40 84. 53 16.00
Test No. 15: ;

(GMC h a 5 ee ear ae a eRe eae oon 18.07 14. 50 80. 24 13.10

INCOSE ee a eee 5 5 eee oo pope aOSeSosene a —— 19. 50 16. 10 82. 56 16. 00
Test No. 16:

NIG Lee ie ae, a ee Se cS tn a rt a ee 18. 27 15.10 82. 64 14.10

IS) IGE. at Bae ease BSoseosr na poser aeEpospac= 2005 16. 00 78.04 15. 70
Test No. 17: ;

Gallediay 3 re ao ae ema - toe eee 18. 60 @ 14.80 79. 56 13.10

BN Ogr alll sae ere eee on te oe ee en ene 22. 43 19. 00 84. 70 820 eke
Test No. 18:

allem asta 22 eh saci eh Solas oo aco bee 21.00 | 17.00 81. 42 15. 40

INFO GU. en Led eee Beas A Re eee Soren seee 21.30 17.90 84.03 16. 80
Test No. 19: ;

eae ee ae Ree eae Conse Saye 22. 40 18. 90 84.37 17. 20

INV AIRE Bete asa 5-SDaades sabe Segee Ea aace = 23.00 19. 20 83. 47 17. 20
Test No. 20: :

GENET ba Se ene oe ee ane eS 16. 92 14. 00 82. 74 14.00

INO lisesi ea. e Se ee Be Lee eee 14. 40 12.10 84,02 11.80

FIELD STUDIES OF THE CROWN-GALL OF SUGAR BEETS. (i

TaBLE I1.—Comparison in sugar content and purity of galled beets with beets not so
affected—Continued,

Gendition ot bects. Solids in Sugar in Coefficient | Sugar in
juice. juice. of purity. | the beet,
Test No. 21: Per cent. Per cent. Per cent. Per cent.
SENG! oo EMRE AS Ba ol 18. 80 16. 50 87. 76 16. 30
i oo eels 2d OSE QRS SORES SEC SESE Ss 18. 32 15. 90 86. 79 15. 20
es :
PEELE bane!) NB Si SAS 16, 42 13. 30 80. 99 12, 60
rs eS eas at cea ee EEE A ARE cae eee are 16, 62 13.90 82. 39 12. 40
est No. 2
Galled oie) icfatate: Sateen = Sage apni aiurs esate 16. 50 13. 40 81. 21 11. 70
a Re eats Re Pee ea rn ie wins Sains oe Cecile 2 WAP 14. 60 82. 39 13. 90
est No. :
UAE arg Let ae ie eet ices atts rete ea 18. 70 15. 30 81.81 14. 20
- roe galls [62 Qt aa SRS 5 San eres ae 22.12 18. 00 81. 37 16. 70
est No. 25
BUND LS | oo we Se het SS 2 a SEG ee ee ene 14. 87 13. 00 87. 42 12. 20
. ate aoe oo seine ob = Soe eR SED Ea een See one an 22.97 19. 30 84. 02 17. 60
es oO. B
“SUD SL 2 oe 2 i a ee rele OI a Ag 19.07 15. 20 79. 70 13. 90
Hs TG Gabi Ss is eS whey 2 a ei IS ae ee er 16.07 13. 60 84. 62 12. 50
verage:
SUNS). a. 2. 2a Be RR eg par re ee ee 17.01 13. 85 81.35 12. 54
WO) Sys cee os Pa ae i Se eae Sasa ae 19. 66 16. 21 82.15 15.18

A study of Table II indicates that gall formations on sugar beets
have a tendency to reduce both the sugar content and the purity
of the roots. The effect upon the sugar content seems to be more
marked than upon the purity. Everyone who has studied the indi-
viduality of the sugar beet knows that there is a difference in the
sugar content and purity of healthy beets growing side by side in
the same row. It is not surprising, therefore, that an occasional
pair shows qualities favorable to the galled beets, as in tests Nos,
10 and 21 of Table II. It is safe to say, however, that in the great
majority of cases the formation of galls upon the roots of sugar beets
has a decidedly injurious effect upon either the purity or the sugar
content or upon both these factors of quality in the beet root.

It seems to be practically impossible to obtain any accurate data
regarding the effect of galls upon the size of the roots affected. We
find the largest as well as the smallest beets more or less seriously
infested with galls, as shown in Plate I, A to #, and it is impossible
to know whether the galled beets would have been larger or smaller
if they had been free from galls. In some infested areas the larger
beets are more generally galled, while in other infested areas the
smaller beets are the ones most generally affected; and since the
individuality of the beet embraces the size and shape, as well as the
quality of the roots, a satisfactory comparison of the weights of the
galled and not galled beets has not been practicable in any of the
areas that have come under the observation of the writer. So far
as one can judge from general field observations, however, the galls
do not seem to have any marked effect upon the size of the beets.
Consequently the tonnage or yield of beets per acre does not seem
to be appreciably affected by the disease except in those cases in which
the galls cause the beet roots to decay.

8 BULLETIN 203, U. S. DEPARTMENT OF AGRICULTURE.

CONTROL OF BEET GALLS.

From our present knowledge of the cause of the crown-gall of
beets, combined with the field observations already made upon this
disease, its elimination or control becomes comparatively simple.
As already suggested, a beet field badly infested with the crown-gail
organism may be freed from the pest by growing some other crop
in that field for two or more years before returning to sugar beets.
It is necessary only that the rotation crops other than beets shall
be such as are not readily attacked by the crown-gall organism.
In Bulletin No. 213 of the Bureau of Plant Industry, already men-
tioned, it is pointed out that the crown-gall organism will attack
a large number of plants in a great variety of families, but there are
plants which are attacked with difficulty, if at all, by this organism.
In the test mentioned in this bulletin the crop grown was oats, but
it is safe to say that any of the small grains, corn, kafir, milo, or
sorghum would do well. If only those crops are grown upon which
the organism can not feed and thrive, it must eventually die out
and the field be left free from the pest. The elimination of crown-
gall is, therefore, a simple matter of wise crop rotation, which as a
matter of good farming should be practiced by every farmer regard-
less of the presence of crown-gall.

SUMMARY.

(1) There are at least two distinct types of sugar-beet galls.

(2) The crown-gall of sugar beets is caused by a bacterium or a
number of closely related bacteria.

(3) Sugar-beet galls appear to have an injurious effect upon the
quality of the roots.

(4) The galls themselves are low in purity and therefore detri-
mental in the milling processes.

(5) Sugar-beet galls sometimes cause the beet roots to decay,
but, so far as general field observations can determine, they do not
appear otherwise to affect the tonnage.

(6) This disease may be held in check by a proper system of crop
rotation with grain-producing plants.

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