SPECIAL CIRCULAR 100 JUNE 1974

Survival of Plant
Pathogenic Bacteria

CURT LEBEN

OHIO AGRICULTURAL RESEARCH AND DEVELOPMENT CENTER

Wooster, Ohio
 CONTENTS
* * *
Introduction ________________________________________________ 3

Sh o rt -term Su rv iv a I ______ ____ _________ _____ _____ __ _______ - - __ 4

Inanimate Factors Affecting Survival _ _____ _ _______ _ ___ __ ____ 4

Animate Factors Affecting Survival _________________________ 6

Constructs _________________________________________________ 7

Hypobiosis ___________________________ _______________________ 7

Long-term Survival ___ _ ____ ______ _ ___ ___ _____________________ 8

Protected Positions _____ _ ___ _ ______ ___ __ ___ __________________ 9

Contributions of the Pathogen Life Cycle Phases to Survival ________ __ l l

Pathogenic Phase __ __ ________ ___________________ _____ ___ l l

Resident Phase _________________________________________ l l

Saprophytic Phase ___________ _ ______ - · - -· _______________ 13

Terminology _ __ ___ ____ _ ______ _____________ _ __ _____ __________ 14

Conclusions _____________________ .. __________________________ 14

Literature Cited ____ .. __ _____ ______ _____ _ _______ _ ____ _ __________ l 5

ACKNOWLEDGMENTS
Thanks are extended to G. C. Daft, H. A. J. Hoitink, P. 0 . Larsen, T. D.
Miller, J. P. Sleesman, and L. E. Williams for constructive criticism of the
manuscript.

AGDEX 630 6-74-2 M

 1
Survival of Plant Pathogenic Bacteria
CURT LEBEN 2

INTRODUCTION
Plant diseases incited by bacteria are not as numerous as those in-
cited by fungi and viruses, but many (probably most) plant species can
be attacked by bacteria at one or more stages in the plant life cycle. Some
bacterial diseases are of great economic importance; e.g., the vascular
wilts of musaceous, solanaceous, and other hosts (incited by Pseudomonas
solanacearum), fire blight of rosaceous plants ( Erwinia amylovora), and
the bacterial blights of cotton and rice ( X anthomonas malvacearum and
X. oryzae). Crops of lesser importance also may be limited because of
the depredations of bacterial pathogens.
It is useful to consider how plant pathogenic bacteria survive during
unfavorable periods, particularly season-to-season periods. This publi-
cation adds to the excellent efforts of the most recent reviewers of this
subject, Buddenhagen ( 7), Crosse ( 11), and Goto ( 25). Buddenhagen
and Crosse stressed survival in the soil and Goto emphasized survival in
relation to vegetation. In this publication, an attempt to link survival
at both sites is presented. An abstract of this paper has appeared ( 39).
When bacteria are considered as a whole, the subject of bacterial
survival is very large indeed, but when plant pathogenic bacteria are
considered, surprisingly little work has been done on this part of the life
cycle. Three constructs-frameworks of fact and speculation-con-
cerning survival are offered in this paper. These are based on work
with plant pathogenic and other bacteria. It is hoped they will offer
a degree of integration and will be useful in suggesting further research.
Bacterial pathogens must survive, of course, or the diseases they
incite would have vanished long ago. It is natural that much effort has
been devoted to the pathogen growth stage-the multiplication phase-
of the life cycle. After all, the products of bacterial growth are seen:
the wilts, the blights, the leaf spots, and all of the imperfections result-
ing in greater or lesser losses. More and more, plant pathologists are
trying to refine understanding of life cycles and modes of survival. This
knowledge may well lead to improved disease control.
There is no need to belabor a useful principle of plant and animal
pathology: if possible, one should attack the pathogen during diminished
1
Based on a lecture presented before the Second International Congress of Plant Pathol-
ogy, University of Minnesota, Minneapolis, Minn., U.S.A ., Sept. 6, 1973. Literature survey
was completed June 1, 1973 . Accepted for publication January 4, 1974.
2
Professor, Department of Plant Pathology, Ohio Agricultural Research and Development
Center, Wooster, and The Ohio State University.

3
survival times. With the plant pathogen, the low population point
usually is between seasons. In this paper, season-to-season survival is
referred to as "long-term". "Short-term" survival means survival for
minutes to days.
Refer·ences to spore-forming bacteria, whi ch do not appear to be
important plant pathogens, are excluded . Survival in arthropods is
also excluded. There are comparatively few instances of long-term sur-
vival of conventional bacteria in these animals. However, recent stud-
ies on "spiroplasmas" ( 16) .and "rickettsia-like" bacteria ( 24) suggest
that survival of unusual forms in arthropods is greater than previously
suspected. Then, too, "mycoplasma-like" organisms, which most rn-
semble bacteria, survive in insects.
At least two other important topics related to survival are men-
tioned only in passing. These are: methods used to detect survivors,
and the exchange of genetic entities between pathogenic and nonpatho-
genic bacteria associated with plants. Recent papers covering assay
methods have been written by Beech and Davenport ( 3), Dickinson
( 18), and Goto ( 25). Gibbins ( 23) has discussed genetic relatedness
with respect to the origin of plant pathogens. In this connection, Starr
and Chatterjee ( 73) have hypothesized that saprophytic Erwinia spp. or
Erwinias associated with plants may be the source of the Erwinias pro-
ducing diseased conditions in man and other animals.

SHORT-TERM SURVIVAL
Plant pathologists are mostly concerned with the practical effects
of long-term survival. However, it is appropriate that attention be giv-
en here to the survival of metabolically and physically active bacteria
for shorter periods-hours to days. Physical, chemical, and microbio-
logical factors affecting short-term survival also affect long-term survi-
val, but information about long-term survival is much more sketchy.
A brief discussion of inanimate and animate factors affecting short-
term survival of bacteria is appropriate.
Inanimate Factors Affecting Survival
Water governs the life of bacteria in vital ways and has a marked
effect on short-term survival. Free water probably is necessary for the
multiplication of bacteria in general, and it is of course required for
swimming. Metabolically active forms of most bacteria are killed by
drying, but there are differences in degree owing to species and condi-
tions of drying.
Active plant pathogenic bacteria existing independently probably
die quickly when they are dried, as would be expected of nonsporing
bacteria in general. There appears to have been no systematic study

4
of plant pathogenic bacteria in this respect, however. Some '70 years
ago, Jones ( 30) and Smith ( 71) placed active cells of pathogens in drops
on cover slips. Slips were dried and assayed later in vitro. Results in-
dicated that season-to-season survival was unlikely and that survival de-
pended on conditions of drying. Jones found that a potato soft rot bac-
terium he studied died within minutes when water suspensions were dried.
Work by Kauffman and Leben with Pseudomonas glycinea suspended in
buffer demonstrated that ca. l on cells per ml. of this pathogen died virtual-
ly as soon as drops of suspension dried on cover slips (unpublished). An-
other recent example of sensitivity to desiccation is that of Kikumoto and
Sakamoto ( 36), who found that Erwinia aroideae cells died rapidly when
soil particles colonized by this pathogen were dried.
The relative humidity during drying influences survival of many
kinds of bacteria. For example, Erwinia amylovora, in common with
the intensively studied Esch erichia coli, survived best at 40-90 % relative
humidity in air-borne particles ( 72) . A number of works indicated that
a low relative humidity limits multiplication of many kinds of macro- and
microflora on the plant surface, including plant pathogenic bacteria. On
the other hand, high humidity favors epiphytic growth in general ( 43).
Swimming may be far more important for survival of plant patho-
genic bacteria than present information suggests. l\1ovement toward or
away from a stimulus is an important fitness factor for many organisms
which swim. Many pathogenic bacteria are motile, and some experi-
ments have indicated they move toward seemingly favored positions ( 58,
62, 69, 80). It is not difficult to envision a bacterium and its progeny
swimming in a drop of water on a leaf toward a source of nutrients (at-
tractants) and that this location may provide a survival site, as suggested
below.
Repellants as well as attractants may help "guide" swimming bac-
teria ( 74, 76). The swimming of a virulent Pseudomonas solanacerarum
isolates appeared to be a fitness factor not for in vivo growth but for in
vitro growth, because Kelman and Hruschka ( 35) found that virulent
isolates were not motile. The subject of swimming of plant pathogens
needs much more investigation.
Numbers of plant pathogenic bacteria exposed on the above-ground
parts of plants probably are quickly reduced by the ultraviolet (UV)
rays in sunlight. Inactivation by UV is influenced by relative humidity
( 65) and other factors. Death from UV must be very large when patho-
gens washed from aerial lesions are spread widely during a wind and
rain storm. Many years ago, Jones ( 30) discovered the sensitivity of the
soft rot bacterium he studied to sunlight and recommended exposing po-
tato tubers to sunlight as well as to drying conditions for disease control.

5
There is much literature on the effects of UV and relative humidity on
the survival of bacteria. Little work has been done with plant pathogens,
however.
In addition to water and sunlight, two other inanimate factors, tem-
perature and the chemical environment at the survival site, undoubtedly
have a profound influence on pathogen survival. They also influence
other microorganisms, especially if water is present and the chemicals
serve as nutrients for these organisms.
The interactions among inanimate factors are varied, complex, and
little studied with respect to their influence on the survival of pathogens.
Animate Factors Affecting Survival
Prior to and during infection, and with the aging of a lesion, patho-
genic bacteria are subject to manifold pressures exerted by other micro-
organisms. Probably the only time when a pathogen is free, or nearly
free, from influences of these other organisms is during the early invasive
growth period within plant tissue. As a lesion ages, especially in a moist
environment and especially in contact with soil, omnipresent nonpatho-
genic microorganisms begin the decomposition of the lesion and the
masses of pathogen cells it contains. In addition to the microflora, many
kinds of fauna may be present, and some of these consume bacterial patho-
gens. It is probable that in regions where plant tissues decompose rapid-
ly, the survival time for a pathogen is shorter than in regions where de-
composition of organic materials takes place more slowly.
Evidence reviewed above suggests that the life expectancy of meta-
bolically active, independent cells of pathogenic bacteria likely is a short
one. Consider the soil environment. Assume that a bacterium began
its independent existence by being washed from a leaf lesion into the soil
or by swimming from a colony near a root. Gray and Williams ( 26)
reviewed work suggesting that microorganisms in the soil are largely in-
active, owing to the sparsity of energy sources and to other unfavorable
factors. More recently, Brown ( 6) presented additional evidence that
bacterial members of the soil flora are in a state of reduced metabolism
most of the time and refers to this condition as "bacteriostasis", a term
parallel to "fungistasis" ( 78).
An immigrating pathogen would encounter these adversities. If
an energy source should become available, very likely other organisms
would be better able to use it, because many plant pathogens are not nu-
tritionally versatile ( 5 7, 66). Other organisms would possess a greater
"relative competitive advantage" with respect to energy sources, to use
the term of Cook and Papendick ( 9). Thus, it seems likely that unless
a suitable niche could be found quickly, immigrating cells of bacterial
pathogens in the soil environment would expire within a short time.

6
Yet some pathogens are ''soil borne" and do persist m soil for years.
These are mentioned below.
Life expectancy of active, independent cells of a pathogen m an
aerial environment also would be expected to be short. In Ohio cli-
mate, bacteria washed from an active lesion on a leaf to another area of
the foliage probably would die soon, owing to desiccation or UV irradia-
tion. If the cell did not meet this end, it then would face the competi-
tion of other organisms already living on the plant. Crosse ( 12), Gib-
bins ( 23), Last and Warren ( 3 7), and Leben ( 43) have written about
the relationships among epiphytic organisms. A notable symposium cov-
ering many aspects of the microbial ecology of the leaf surface was held
in 1970 in Newcastle ( 63).

CONSTRUCTS
Three constructs are offered as a framework for thinking about the
survival of plant pathogenic bacteria.
1. Long-term survival of pathogens in nature takes place only in
association with living or dead plant tissues. This is a "critical trait"
( 1) which allows pathogens to survive in the face of recurrent or occa-
sional stresses.
2. Long-term survival is not likely to take place unless cells of the
pathogen are in aggregates or unless they are associated with living plant
tissues in "protected positions".
3. Pathogens in a state of reduced metabolism are more likely to
survive than are active cells.

HYPOBIOSIS
Bacterial cells in a state of reduced metabolism are designated hy-
pobiotic cells. The term is taken from the microbiological literature
( 52) and has had little use in plant pathology. Yet the concept bears
special significance. In the hypobiotic state, microorganisms may live
long periods without added nutrients and are more likely to survive the
physical and chemical stresses causing death than when metabolic ac-
tivity is high. Hypobiosis can be induced experimentally by low tem-
perature, loss of water, increasing salt concentration, and a variety of
other means.
It seems reasonable to conclude that pathogens surviving for long
periods are in a hypo biotic condition, having arrived at that state as a
result of the natural processes takjng place with the aging of diseased
tissues. Hypobiotic cells would be the survivors in dry leaf, stem, and
root lesions of annual plants: they would represent a small portion of
the masses of cells once alive within the lesion. Hypobiotic cells are, of
course, quite different from actively metabolizing ones, and they de-

7
serve much more study than has been given them. For example, cells
of Pseudomonas aeruginosa were more sensitive to desiccation in the ex·
ponental growth phase than when cells were 7 days old (70). Normand
et al. ( 60) found that the morphology of Pseudomonas phaseolicola cells
was different in the lesion center than at the lesion edges, where the bac-
teria were younger.

LONG-TERM SURVIVAL
Some bacterial pathogens can survive for many ·years if they are in
diseased plant tissues which are dry, as is well known. The surviving
cells in these tissues, in addition to being in a hypobiotic state, probably
are protected in varying degrees by the surrounding masses of dead bac-
terial and plant cells and by products of the pathogen-host interaction.

FIG. 1.-The filamentous form of exudate oozing from a cotyledon

lesion incited by Pseudomonas glycinea. Exudates consist of masses of
bacteria in a matrix.

8
On the other hand, some pathogens die when the lesion dries ( 31, 61).
If a pathogen survives in the dry lesion and the lesion is moistened and
exposed to microbial decomposition, as in soil, it is anticipated that sur-
viving cells surrounded by a quantity of diseased tissue would not be as
readily decomposed as when imbedded in smaller amounts. Further-
more, cells within tissues would be less subject to decomposition than
those in soft tissues.
Debris from diseased plants must always be considered a possible
source for seasonal carryover. However, two questions should be ask~d.
How likely is it that debris will decompose between seasons? If it is not
decomposed, what is the likelihood that surviving pathogen cells will come
in contact with susceptible plant tissue? In working with Pseudomonas
glycinea, for example, it was found that overwintering in quantities of di-
seased leaves was poor in soil, better on sod, and better yet if diseased
leaves were suspended in air ( 14). The differences are attributed to the
amounts of moisture and the relative numbers of decomposing micro-
organisms at the three sites. With ordinary crop cultivation methods, sur-
vivors in diseased leaves probably would occur near susceptible seedlings
rarely (if ever), particularly if crops were rotated.
On the other hand, diseased debris associated with seed likely would
he kept dry and thus not subject to deterioration. In this condition, cells
of a pathogen may well survive as long as the seed is viable. Baker ( 2)
has reviewed all aspects of seed pathology. This is an important subject,
because survival with seed is responsible for much damage produced by
bacterial pathogens of annual crops.
Lesion exudate is another type of debris resulting from disease.
Exudates consist of masses of pathogen cells in a matrix, which may hard-
en. Thus with Erwinia amylovora, exudate in the form of strands may
disseminate the pathogen via air currents within the orchard ( 33). Sur-
vival in dry strands for months has been reported.
Exudates are produced in lesions incited by many pathogens. For
example, when head-shaped exudates of Pseudomonas p,-Zycinea were
moistened, they were dispersed and formed flakes when dried ( 15) .
Flakes could be disseminated locally by the wind. It is suspected that
long-term survival of pathogens in exudates in nature is not common,
though, because most exudates would he .degraded by physical or chemi-
cal a-ction, or they would become moiste11ed and destroyed by m' :-' i
i croor-
-f •

garnsms.
PROTECTED POSITIONS
It is clear that many bacterial pathogens survive well ·under some
conditions if they are in aggregates. As indicated, they probably die
quickly as individuals in soil or on exposed sites above ground. How-

9
ever, recent evidence suggests that individuals or a few cells can survive
for varying periods of time in association with the healthy living plant
or plant part, at sites termed "protected". It is emphasized that plant
tissues are healthy as far as can be determined.
The protected position hypothesis first was deduced by experiments
in which water drops were placed on healthy cucumber leaves, and the
areas covered by these drops were examined later for bacteria ( 46, 4 7).
Sizeable populations of nonpathogenic bacteria developed in 48 hours in
many drop areas. In contrast, few bacteria were present in areas not
covered by drops. It was concluded that the progenitors of these bac-
teria were in some sort of sheltered locations on the leaf surface before
drops were placed over them. Probably they were hypobiotic. Blake-
man ( 4) and Blakeman and Fraser ( 5) also obtained a rapid buildup
of epiphytic bacteria in water drops on leaves of two plant species.
The bacteria just discussed were nonpathogenic. Recent work sug-
gests that pathogenic bacteria also survive in protected positions on
healthy leaves. Mew and Kennedy ( 55) found that Pseudomonas gly-
cinea multiplied and survived for at least 14 days on healthy soybean
leaves in the greenhouse. Scherff ( 67) and Kauffman and Leben ( un-
published) confirmed these results. In the latter tests, three levels of
relative humidity were used; multiplication and survival were observed
at each level: low, medium, and high. Multiplication or survival at the
low relative humidity was not expected because P. glycinea is sensitive to
drying and epiphytic growth of all types is reduced by a low relative hu-
midity ( 4·3). Consequently, it was concluded that the pathogen multi-
plied in a protected, moist site and survived at or near this location. Oth-
er workers have found pathogens surviving on healthy host or nonhost
leaves ( 10, 13, 22, 28, 32, 64, 77).
From these examples, one may extend the hypothesis of protected
positions to other locations associated with the healthy living plant or
plant part. For example, a few hypobiotic cells of a pathogen within
a seed would be in a well-protected and significant survival site. Buds
may be protected survival sites ( 41), and pathogens may survive within
fruit tree stems ( 8, 34). The short report of Meneley and Stanghellini
( 54) indicated that the interior parts of a number of fresh vegetable prod-
ucts served as protected survival sites for bacteria which caused tissue de-
composition when these products were warmed.
The rhizosphere also may serve as a protected position for the sur-
vival of some pathogenic bacteria. Pathogens have been found associ-
ated with apparently healthy roots of host and nonhost plants. Goto
( 25) has reviewed this literature, much of which is in Japanese. Possi-
bly these pathogens are living in the mucigel surrounding the root ( 27).

10
 It seems significant that most pathogens do not survive well in soil, and
the few that do persist, the soil-borne pathogens, possess wide host ranges.
Assume that a soil-borne pathogen is able to occupy a niche on the
rhizoplane of healthy host or nonhost plants. Pathogens at this location
then could exist in soil for as long as there were roots of suitable species
to provide sites for multiplication and survival. If suitable crop or weed
plants were not grown, the pathogen would disappear. Unless plants
became diseased, with the subsequent release of large amounts of inocu-
lum, populations probably would remain at relatively low levels. A
pathogen of this sort could be distributed in soil by contact between roots,
by moving water and agricultural implements, by swimming, or by nema-
todes ( 29) and other fauna.
A parallel with Rhizobia spp. is evident here. Rhizobia grow and
survive in association with nonhost as well as host roots, and it is suggested
that they survive in the soil as long as these roots are present ( 19). If
this suggestion is correct, an explanation is offered for the long-term di-
sease potential of certain pathogens in the soil in the face of what is de-
scribed as a hostile environment.
Protected positions probably are most significant for low numbers
of cells of either pathogenic or nonpathogenic bacteria. However, it is
possible that large numbers also could survive at these locations.

CONTRIBUTIONS OF THE PATHOGEN LIFE

CYCLE PHASES TO SURVIVAL
The life cycle of a plant pathogenic bacterium may be divided into
four phases or stages. These are the pathogenic, resident, saprophytic,
and survival phases. The first three-pathogenic, resident, and sapro·-
phytic-are growth phases. What is the contribution of each in produ-
cing surviving cells?
Pathogenic Phase
As far as known, the pathogenic phase, in which there is a large
increase in numbers of pathogen cells and the production of symptoms,
contributes most of the cells entering a seasonal survival period. The
larger the population of a pathogen entering the period, the greater the
chances for survival, other conditions being equal. Large populations
would be produced by compatible isolate-host combinations (those re-
sulting in typical disease responses) ; incompatible combinations ( resis-
tance responses), which produce lower populations, would be much less
likely to yield surviving cells.
Resident Phase
A decade ago it was suggested, on the basis of work with X antho-
monas vesicatoria, that some pathogenic bacteria possessed a "resident

11
phase" in their life cycle. This was defined as the capacity for multipli-:-
cation on the surface parts of the healthy shoot system ( 44) . A revised
definition is offered below.
It now appears as a result of recent studies that a number of bacterial
pathogens possess a resident phase, which may be associated with leaves,
buds, or flowers of host or nonhost plants ( 17, 20, 21, 22, 28, 38, 49, 50,
51, 53, 55, 56, 59, 67, 75). Earlier work is cited by Leben (43).
Pseudomonas syringae) which has a wide host range, is the subject of most
of these papers. Many leaf-spotting pathogens may be residents under

FIG. 2.-A scanning electron microscope view of the surface of a soy-

bean stipule, showing pillow-shaped epidermal cells, the base of a tri-
chome, and a stoma. Most of the small projections on the epidermal cells
are aggregates of bacteria. These organisms also are found in abun-
dance in depressions between epidermal cells, especially near the base of
trichomes. These locations are possible pro~ected survival positions.

12
some conditions. The increase of a pathogen in the absence of symptoms
may be of epidemiological importance by serving to build up an inoculum
as an immediate prelude to infection. That increase also could provide
pathogen cells which survived unfavorable times. Probably survival sites
would he at or near multiplication sites.
The location of these sites is speculative, hut there are several possi-
bilities. Leaves have been studied the most. In Ohio, relatively few
non pathogenic bacteria were detected on leaves of field plants ( 4 7). This
suggests that these organisms, and likely pathogenic bacteria as well, sur-
vive in protected positions. Probably there is a periodic (possibly diur-
nal) rise and fall in numbers of emigrants swimming or being washed
from these sites, with population levels being governed primarily by the
amount of water on the leaf, duration of periods of wetness, relative hu-
midity, and exposure to sunlight. The leaf surface, especially within
deep, moist depressions between epidermal cells, probably is a multiplica-
tion and survival site. The underside of the leaf would be expected to
have more protected positions than the exposed upper surface. The sub-
stomatal chamber and associated tissues also are suspect locations ( 79).
However, water does not readily enter stomata! pores ( 68). Protected
positions may be associated with trichomes ( 42) or with hydathodes or
other natural openings. These hypothetical locations could be occupied
by pathogenic and nonpathogenic bacteria. In addition, pathogens may
multiply and survive unnoticed as a result of an arrested infection of a
few epidermal cells. More work is needed to understand what is taking
place, particularly with pathogens.
Buds also are worthy of comment as sites of multiplication and sur-
vival of pathogenic bacteria ( 41 ) . In contrast to leaves in Ohio, the
bud habitat (the gemmisphere) of some plants may carry a high popula-
tion of a varied bacterial flora. The numbers and variety of nonpatho-
gens in soybean buds, flowers, and young fruits were remarkable ( 40).
Recently a Pseudomonas syrin!{ae type was found among the many bac-
teria in buds of healthy field soybean plants. Curiously, this pathogen
produced progressive lesions on the soybean hypocotyl and cotyledon, but
only incited a resistance response on leaves ( 49).
Saprophytic Phase
Do plant pathogenic bacteria have a true saprophytic stage in na-
ture? Are pathogens able to grow- multiply- on dead tissue derived
from either host or nonhost plants? Can pathogens multiply on other
material in natural habitats? Some pathogens in aggregations readily
survive in dead host tissue, as has been seen, but do they increase in these
tissues? No direct evidence is known for the proposition that they do
mcrease. On the other hand, there is much evidence that populations

13
decrease. With the soybean blight pathogen, for example, once leaf le-
sions became necrotic, the pathogen population decreased. This took
place if leaves were left on the plant or if they were detached and placed
in soil or other natural situations ( 14, 50) .
Probably saprophytic growth has been suspected to take place most
often in soil. If the explanation proposed above for long-term persistence
of soil-borne pathogens is accepted, saprophytism would not be essential
for survival. In general, soil seems to be particularly inhospitable for
bacterial pathogens, as has been mentioned. Consequently, it appears
that true saprophytic growth in this medium is limited. If so, it would
have little effect on survival. Obviously, more needs to be known about
this difficult subject.

TERMINOLOGY
In 1961 "resident" was defined as a member of the microflora multi-
plying on the surface of the aerial parts of the healthy plant proper ( 45).
Bacterial residents may be detected readily on the plant surface with suit-
able techniques. However, the possibility that some bacteria are within
the plant and reach the surface through openings is difficult to exclude.
In any event, workers have used "root resident" or "internal resident".
Consequently, it is now proposed that the term resident be expanded to
include all types of associations of microflora with healthy plants. This
includes the surface and interior parts, above and below ground. Per-
haps this alteration will avoid confusion. No strong brief for the original
term or this expansion is held; some may find it useful, however, to have
a short designation for an important ecologic grouping of the microflora.
With this terminology, a pathogenic bacterium could possess pathogenic,
resident, and saprophytic phases in its life cycle, as has been indicated, but
the resident phase would not be restricted to the shoot surface.

CONCLUSIONS
Plant pathogenic bacteria are poorly adapted for survival in nature
away from plant tissues. They are likely to survive seasonal periods in
aggregates in association with dry diseased plant tissues, or as individuals
or a few cells in protected positions associated with the healthy living plant
or plant part. Surviving cells likely are in a hypobiotic state rather than
in the active, vigorous conditions most often studied in the laboratory.
In nature, the pathogenic phase of the life cycle probably contribute-,
most of the cells carrying the pathogen through adverse periods. The
contribution of the resident phase may well be important, but the extent
of its natural occurrence is unknown. On the other hand ) the saprophy-
tic phase seems to contribute few cells which survive.

14
 If these ideas have validity, it is clear that further studies are needed,
especially of hypobiosis, and the nature of protected survival sites and how
pathogens get into them. With more understanding, plant pathologists
may learn how to alter hypobiosis and to expose protected positions to
advantage.

LITERATURE CITED
1. Alexander, M. 1971. Biochemical ecology of microorganisms.
Ann. Rev. Microbiol., 25 :361-392.
2. Baker, K. F. 1972. Seed pathology, pp. 317-416. In Seed biol-
ogy. T. Z. Kozlowski [ed.]. Academic Press, New York and
London.
3. Beech, F. W. and R.R. Davenport. 1971. A survey of methods
for the quantitative examination of the yeast flora of apple and
grape leaves, pp. 139-157. In Ecology of leaf surface micro-or-
ganisms. T. F. Preece and C. H. Dickinson [ed.]. Academic
Press, New York and London.
4·. Blakeman, J. P. 1972. Effect of plant age on inhibition of Botrytis
cinerea spores by bacteria on beetroot leaves. Physiol. Plant Path-
01., 2:143-152.
5. Blakeman, J. P. and A. K. Fraser. 1971. Inhibition of Botrytis;
cinerea spores by bacteria on the surface of chrysanthemum leaves.
Physiol. Plant Pathol., 1 :45-54.
6. Brown, Margaret E. 1973. Soil bacteriostasis limitation in growth
of soil and rhizosphere bacteria. Can. J. Microbiol., 19: 165-199.
7. Buddenhagen, I. W. 1965. The relation of pl ant-pathogenic bac-
teria to the soil, pp. 269-283. In Ecology of soil-borne plant patho-
gens. K. F. Baker and W. C. Snyder [ed.]. Univ.of Calif. Press,
Berkeley.
8. Cameron, H. R. 1970. Pseudomonas content of cherry trees.
Phytopathol., 60:1343-1346.
9. Cook, R. J. and R. I. Papendick. 1972. Influence of water poten-
tial of soils and plants on root disease. Ann. Rev. Phytopathol.,
10:349-374.
10. Coyne, D. P., M. L. Schuster, and K. Hill. 1973. Genetic control
of reaction to common blight bacterium in bean ( P haseolus vul-
garis) as influenced by plant age and bacterial multiplication. J.
Am. Hort. Sci., 98 :94-99.
11. Crosse, J. E. 1968. Plant pathogenic bacteria in soil, pp. 552-572.
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This page intentionally blank.
BETTER LIVING IS THE PRODUCT
of research at the Ohio Agricultural Researc:h and Development Center.
All Ohioans benefit from this product.
Ohio's 110,000 farm families benefit from the results of agricultural
research translated into increased earnings and improved living condi-
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of Ohio consumers. They enjoy the end products of agricultural science
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changing the name to Ohio Agricultural Research and Development Cen-
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Center's research program today .
Research at OARDC deals with the improvement of all agricultural
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The State Is the Campus for
Agricultural Research and Development

GREEN SPRINGS ~
CROPS RESEARCH UNIT NORTH CENTRAL
e BRANCH
NORTHWESTERN • MAHONING CO.
BRANCH MUCK CROPS e FARMe
BRANCH
WOOSTER

•
CENTER
HEADQUARTERS

e POMERENE FOREST
LABORATORY

COLUMBUS

•
THE OHIO STATE EASTERN OHIO RESOURCE
WESTERN• UNIVERSITY DEVELOPMENT CENTER
BRANCH , •

SOUTHEASTERN
BRANCH

JACKSON•
•
BRANCH

Ohio's major soil types and climatic Jackson Branch, Jackson, Jackson
conditions are represented at the Re- County: 344 acres
search Center's 13 locations. Thus, Mahoning County Farm, Canfield:
Center scientists can make field tests 275 acres
under conditions similar to those en- Muck Crops Branch, Willard, Huron
countered by Ohio farmers. County: 15 acres
Research is conducted by 15 depart- North Central Branch, Vickery, Erie
ments on more than 6500 acres at Cen- County: 335 acres
ter headquarters in Wooster, nine
branches, Green Springs Crops Re- Northwestern Branch, Hoytville,
search Unit, Pomerene Forest Labora- Wood County: 247 acres
tory, and The Ohio State University. Pomerene Forest Laboratory, Keene
Center Headquarters, W o o s t e r, Township, Coshocton County: 227
Wayne County: 1953 acres acres
Eastern Ohio Resource Development Southeastern Branch, Carpenter,
Center, Caldwell, Noble County: Meigs County: 330 acres
2053 acres Southern Branch, Ripley, Brown
Green Springs Crops Research Unit, County: 275 acres
Green Springs, Sandusky County: Western Branch, South Charle~ton,
26 acres Clark County: 428 acres