BACTERIA INFECTIONS OF PLANTS
INTRODUCTION
Anyone who has ever planted a garden knows not only the rewards of
beautiful flowers, fruit, and/ or vegetables, but also the disappointment when plants
become diseased or damaged. Many factors cause plants to exhibit poor vigor,
changes in appearance, or even death. Both abiotic (non-living) and biotic (living)
factors can negatively impact plant heath. Disorders that result from non-living
factors (such as nutrient deficiencies, over/under watering, temperature stress, and
chemical damage) are discussed in subsequent chapters. This chapter focuses on
those living organisms that cause disease: fungi, water molds, bacteria, viruses,
nematodes, phytoplasmas, and parasitic plants.
History & Significance
Plant loss to homeowners may result in frustration and minor monetary cost.
However, on a global scale, plant diseases cause an estimated $38 billion1 in annual
losses. History also provides some perspective on the impacts of plant disease.
One of the most notable historical impacts of plant disease was caused by late
blight of potato. This disease was a major contributing factor in the Irish potato
famine of 1845. During this time, approximately one million people perished from
starvation; a million and a half more are believed to have left Ireland and immigrated
to the United States. The late blight pathogen is still present in production systems
today, but it is managed by resistant varieties, sanitation, and fungicides.
Near complete loss of the American chestnut was caused by a fungal disease
that nearly wiped out forests in the eastern United States. The trees once grew to
majestic heights within their native forest habitat, as well as in urban plantings. They
provided high quality hardwood for building construction and nuts as a food source
for people and wildlife. However, in the late 1800s, chestnut blight was accidentally
introduced to the United States through imported Chinese chestnut trees. While
Chinese chestnuts are tolerant to the blight, American chestnut trees are not, and in
less than 40 years, approximately 30 million acres of chestnut trees died. Chestnut
blight remains a problem, and researchers are still seeking options for management.
Disease management changed drastically when the French wine industry gave
way to the first fungicide, Bordeaux mixture. France’s grape production had long
been devastated by powdery mildew and downy mildew. Then in 1882, the discovery
of a copper sulfate and lime mixture helped manage these vineyard diseases.
Bordeaux mixture is still used in a modified form to manage powdery mildew,
downy mildew, and other fungal diseases on numerous types of plants. These
examples represent situations in which plant diseases have reached historical
proportions. However, the amount of damage that plant diseases cause varies
depending upon factors such as environment, host health and susceptibility, and
pathogen biology. There are many options for managing disease development and
spread. The effectiveness of management techniques begins with proper
identification of the disease and/or causal organism.
Pathogens
A plant disease is any physiological or structural abnormality that is caused by
a living organism. Organisms that cause disease are referred to as ‘pathogens,’ and
affected plants are referred to as ‘hosts.’ Many organisms rely on other species for
sources of nutrients or as a means of survival, but are not always harmful to the host.
For example, saprophytic organisms obtain nutrients from dead organic material and
are a vital part of many ecosystems. Plant pathogens, on the other hand, utilize hosts
for nutrients and/or reproduction at the hosts’ expense. Disease causing organisms
include fungi, oomycetes (fungus-like organisms called water molds), bacteria,
viruses, nematodes, phytoplasmas, and parasitic seed plants.
Once a pathogen infects a host, symptoms often develop. Symptoms are the
outward changes in the physical appearance of plants. Symptoms take time to
develop, and thus, disease development may be delayed for several days, weeks,
months, or even years after initial infection occurs. Examples of symptoms include
wilt, leaf spots, cankers, rots, and decline.
Physical evidence of pathogens (called ‘signs’) may also be observed on
diseased tissue. Examples of signs include fungal fruiting bodies, bacterial ooze,
nematode cysts, and fungal mycelia. Both symptoms and signs are utilized in making
disease diagnoses.
Conditions for Disease Development
Disease development is dependent upon three conditions: a susceptible host
plant, a favorable environment, and a viable pathogen. All three of these factors must
be present for disease to occur. Each side of the triangle represents one of these
factors: host plant, environment, or pathogen. When all three sides of the triangle are
complete, disease occurs. If one of the conditions is not present (one side of the
triangle is missing), then disease does not occur. By altering the susceptibility of host
plants, the surrounding environment, and/or the viability of pathogens, the disease
triangle can be broken and disease development prevented.
Host plant genetic makeup determines its susceptibility to disease. This susceptibility
depends upon various physical and biochemical factors within the plant. A plant’s
stature, growth habit, cuticle thickness (a protective outer layer on plant tissues), and
shape of stomata (small openings that allow water, oxygen, and carbon dioxide in
and out of plant tissues) are a few physical factors that influence disease
development. Plants may also produce biochemical compounds that limit or prevent
colonization or infection. Growth stage and ability to deter pathogens can also impact
plant susceptibility to disease. For example, young leaves are often more susceptible
to infection than mature leaves.
Environment plays an important role in disease development. Pathogens generally
require specific environmental conditions for infection and spread. Most plant
pathogens require high humidity and moderate temperatures. Other pathogens, such
as bacteria and water molds, require surface water for spread. In some disease cycles,
environmental conditions influence the development of symptoms. For example,
extreme temperatures or drought can cause plant stress; this loss of vigor can
increase host susceptibility to both infection and disease development. Other
environmental factors affecting disease can include those resulting from planting and
maintenance practices. For example, high density plantings can a have higher relative
humidity, while overhead watering increases leaf surface moisture needed by
pathogens to infect plant leaves.
Pathogens must be present and viable in order to infect plants and cause disease.
Removal of infected plant parts and other remnants of pathogens makes them
unavailable for infection. Fungicides also reduce amounts of inoculum (infective
propagules) available for infection. Many pathogens, however, have developed
specialized structures that ensure survival during adverse conditions. For example,
several water molds and fungi are capable of surviving in soil for many years until
conditions are favorable for infection. Pathogens may also survive winter
temperatures and other harsh conditions in infected plant tissue. If a susceptible host
and favorable environment are not available, some pathogens can assume a dormant
state for many years.
Bacterial Infection in Plants
Bacteria are microscopic organisms typically composed of single cells. About
200 types of bacteria are known to cause plant diseases. Due to their small size, a
high-magnification microscope is required to observe bacteria. Occasionally, when a
large number of cells are present, plants may be observed ‘oozing’ bacteria and other
organic byproducts.
Bacteria are capable of rapid reproduction through a process known as binary
fission. In this process, one cell divides to become two, then two divide to become
four cells, and so on. Within a few hours one bacterial cell can become thousands,
and under ideal conditions, populations can double in as little as 20 minutes.
Unlike fungi and water molds, bacteria are not able to penetrate plant tissue
directly. They must infect via wounds or natural plant openings such as stomata. Free
water is required for infection. Once inside plants, bacteria begin to reproduce
immediately. Some types of bacteria produce toxins or enzymes that degrade plant
tissue, and the tissue is then utilized as a food source. Some bacteria can colonize
vascular systems of plants, which results in restriction of water movement.
Bacteria spread by water/splashing rain, wind, or insects, and then move
across plant tissues in surface water to reach wounds or natural openings. Some can
survive for five or more years in soil, as well as in plant debris and cankers. Common
symptoms caused by bacteria include leaf spots, blights, cankers, galls, wilt, dieback,
and soft rots.
Of the over 15,000 identified species of bacteria most are saprophytic and are
of great benefit in decomposing dead and rotting organisms thereby releasing their
nutrients back into the environment. This is the most important roll that bacteria play
in nature. Plants rely on nitrogen from the soil but cannot directly acquire it from the
gaseous nitrogen in the atmosphere. The primary way nitrogen is supplied to plants is
through the mineralization of organic material in the soil. However, nitrogen fixation
by bacteria such as Rhizobium spp. and Cyanobacteria spp. is almost as important as
mineralization, and is a primary source of nitrogen. As these bacteria metabolize they
convert gaseous nitrogen into nitrates or nitrites that become available to plants.
Most phytopathogenic bacteria are aerobic (live in the presence of oxygen)
and some are facultative anaerobes which can grow with or without oxygen. Some
bacteria have thick, rigid cell walls which will retain dye from a cell staining method
developed by Christian Gram, while other bacteria will not accept this stain. This
method of staining results in the bacteria being classed as Gram-positive or Gram-
negative and is an important factor in identification and classification. Gram-positive
bacteria appear purple and Gram-negative bacteria appear pink under magnification.
Bacteria are also distinguished by the different kinds of enzymes they either can or
cannot use for nourishment and the nutrient media on which they can grow.
Rod shaped bacteria reproduce asexually by the process of binary fission (the
transverse splitting in two of a bacterial cell). This process takes place when the
cytoplasmic membrane grows inward dividing the cytoplasm into two approximately
equal parts. When the cell walls are completely formed the cell splits into two cells.
During this process the nuclear material duplicates itself and becomes distributed
equally between the two cells. Bacteria can reproduce at a very rapid rate; some
species can divide every 20 minutes under ideal conditions. It is conceivable that a
single bacterium could produce one million progeny in less than 24 hours. However,
with limited food supply, environmental conditions and other factors the optimum
conditions rarely occur in nature.
There are around 200 species of phytopathogenic bacteria and almost all of
them are parasites within the plant, on its surface, in plant debris or in the soil as
saprophytes. Dissemination of bacteria can be accomplished by several means. Some
bacteria can survive on inanimate objects, in water or inside insects. It is important to
know the survival characteristics of bacteria for effective management strategy and
intervention in dissemination. Some species have the ability to move short distances
in water on their own power by use of their flagella. Most bacteria, however, are
disseminated by passive agents such as air and insects, water and soil movement, and
to a lesser degree by humans, water and other animals. Infected seeds and transplants
can also be a source of inoculums. Most bacteria require a wound or natural opening
(e.g. stomata, lenticels or hydathodes) to gain entry into the host tissue and also
require warm, moist conditions to establish a colony. Windblown soil and sand will
commonly cause wounds which can facilitate bacterial infections.
Bacteria colonize a host by growing between the cells and absorbing the cells
nutrients that leak into intercellular space or grow within the vascular tissue of the
plant. Depending on the species of bacteria and the tissue infected they produce and
release enzymes that degrade cell walls, growth regulators that alter the plants
normal growth, toxins that degrade cell membranes and complex sugars that plug
water conducting tissue
The following is a general classification (Agrios, 5th Ed, 2005) of
phytopathogenic prokaryotes with the exception of the Division Tenericutes, Class
Mollicutes, which will be addressed in a later section. Genera in bold type are
common plant pathogens.
Kingdom: Procaryotae
Bacteria – Have cell membrane and cell wall and no nuclear membrain.
Division: Bacteria – Gram-positive
Class: Proteabacteria – Mostly single celled bacteria.
Family: Enterobacteriaceae
Genus: Erwinia, causing fire blight of pear and apple, Stewart’s wilt in corn,
and soft rot of fleshy vegetables.
Pantoea, causing wilt of corn.
Serratia, S. marcescens, a phloem-inhabiting bacterium causing
yellow vine disease of cucurbits.
Sphingomonas, causing brown spot of yellow Spanish melon fruit.
Family: Pseudomonadaceae
Genus: Acidovorax, causing leaf spots in corn, orchids and watermelon.
Pseudomonas, causing numerous leaf spots, blights, vascular wilts,
soft rots, cankers, and galls
Ralstonia, causing wilts of solanaceous crops.
Rhizobacter, causing the bacterial gall of carrots.
Rhizomonas, causing the corky root rot of lettuce.
Xanthomonas, causing numerous leaf spots, fruit spots, blights of
annual and perennial plants, vascular wilts and citrus canker.
Xylophilus, causing the bacterial necrosis and canker of grapevines
Family: Rhizobiaceae
Genus: Agrobacterium, the cause of crown gall disease.
Rhizobium, the cause of nitrogen-fixing root nodules in legumes.
Family: still unnamed
Genus: Xylella, xylem-inhabiting, causing leaf scorch and dieback disease on
trees and vines.
Candidatus liberobacter, Phloem inhabiting, causing citrus greening
disease.
Unnamed, laticifer-inhabiting, causing bunchy top disease of papaya.
Division: Firmicutes - Gram-positive bacteria.
Class: Firmibacteria – Mostly single celled bacteria.
Genus: Bacillus, causing rot of tubers, seeds, and seedlings and white
stripe of wheat.
Clostridium, causing rot of stored tubers and leaves and
wetwood of elm and poplar.
Class: Thallobacteria – Branching bacteria
Genus: Arthrobacter, causing bacterial blight of holly, thought to be
the cause of Douglas-fir bacterial gall.
Clavibacter, causing bacterial wilts in alfalfa, potato, and
tomato.
Curtobacterium, causing wilt in beans and other plants.
Leifsonia, causing ratoon stunting of sugarcane.
Rhodococcus, causing fasciation of sweet pea.
Streptomyces, causing common potato scab.
Diagnostics Symptoms of Bacterial Infections
Symptoms of bacterial infection in plants are much like the symptoms in
fungal plant disease. They include leaf spots, blights, wilts, scabs, cankers and soft
rots of roots, storage organs and fruit, and overgrowth.
Bacterial spots: the most common symptom of bacterial disease is leaf spots. Spots
appear on leaves, blossoms, fruits and stems. If the spots appear and advance rapidly
the disease is considered blight. Spots on leaves of dicotyledonous plants often have
a rotten or fishy order, are water soaked and are initially confined between the leaf
veins and will appear angular. In some cases bacterial ooze will be present; this is
diagnostic for bacterial infections. Sometimes a chlorotic halo will surround the
bacterial lesion of an infected leaf. Spots may coalesce causing large areas of
necrotic tissue. Bacterial spots will appear as streaks or stripes on monocotyledonous
plants. Almost all bacterial leaf spots and blights are caused by the genera
Pseudomonas and Xanthomonas.
Cankers: primarily Pseudomonas and Xanthomonas cause canker disease of stone
fruit and pome fruit trees, and canker disease of citrus respectively. Canker
symptoms can appear on Trunks, stems, twigs and branches. The most conspicuous
symptom of a bacterial canker disease in stone and pome fruit trees is the
development of cankers and gum exudation (gummosis). Cankers can be slightly
sunken, dark brown and much longer than broad. The cortical tissue of the canker
can be orange-brown to dark brown. Gum is produced in most cankers and some
branches and twigs. Cankers that do not produce gum may have a sour odor and be
soft, sunken and moist. Cankers that girdle trunks and branches can result in leaf
stress and eventual dieback of the portion of the tree distal to the canker.
Bacterial Galls: bacterial galls can be produced by the genus Agrobacterium and
certain species of Arthrobacter, Pseudomonas, Rhizobacter and Rhodococcus.
Agrobacterium tumefaciens, A. rubi and A. vitis alone are responsible for galls in
over 390 plant genera worldwide. Galls of these genera have been referred to as
crown gall, crown knot, root knot and root gall. Species of these bacteria are thought
to be present in most agriculture soil. A wound in the host is required for the
pathogen to gain entry into the host tissue. Gall tissue is composed of disorganized,
randomly proliferating cells that multiply in the intercellular (between the cells)
spaces in the vicinity of the wound. In the presence of the pathogen rapid and
continuous cell division (hyperplasia and hypertrophy) of the plant tissue persists.
Gall damage can be benign to deadly. Crown gall first appears as small, whitish, soft
round overgrowths typically on the plants crown or at the main root. The color of
galls (tumors) caused by A. tumefaciens can be orange-brown and as it enlarges the
surface can become convoluted and dark brown. This is most often found in
commercial nurseries.
Bacterial Vascular Wilts: Vascular wilts caused by bacteria primarily affect
herbaceous plants such as vegetables, field crops, ornamentals and some tropical
plants. The causal pathogen enters, multiplies in, and moves through the xylem
vessels of the host plant and interferes with the translocation of nutrients and water
by producing gum. The pathogen will often destroy parts of the cell wall of the
xylem vessels resulting in pockets of bacteria, gums and cellular debris. The
symptoms of bacterial wilt disease include wilting and death of the aboveground
parts of the plant. In some cases bacterial ooze seeps out through stomata or cracks
onto the surface of infected leaves. Usually this ooze does not occur until the infected
plant tissue is dead.
Bacterial Soft Rots: Primarily the bacteria that cause soft rots in living plant tissue
include Erwinia spp., Pseudomonas spp., Bacillus spp. and Clostridium spp. Many
soft rots are caused by nonphytopathogenic bacteria which are saprophytes that grow
in tissue that has been killed by pathogenic or environmental causes. Soft rots attack
a large number of hosts and are best known for causing disease in fleshy plant
structures both above and below ground. These bacteria are almost always present
where susceptible plants under stress are in the field or in storage. Soft rot pathogens
enter the host through wounds. After entering the host tissue these bacteria produce
enzymes that break down the middle lamella causing separation of the cells at the site
of the infection. The cells die and disintegrate. Rotting tissue becomes watery and
soft and bacteria will form slimy foul smelling ooze that will ooze out of infected
tissue. Bacterial ooze is diagnostic of soft rot diseases.
Bacterial scabs: bacterial scabs primarily infect belowground parts of plants such as
potatoes. Common scab of potato is caused by Streptomyces scabies which cause
localized scabby lesions on the outer surface of the tuber. Typically corky tissue will
form below and around the lesion. Rot pathogens can gain entrance into the host
tissue through these lesions and further degrade the host.
REFERENCES
Agrios, 5th ed. 2005, Plant Pathology WSU, OSU U of I, 2005, Pacific Northwest Plant
Disease Handbook
American Phytopathological Society Glossary http://www.apsnet.org/edcenter
/illglossary/Pages/ default.aspx
Jim Cooper, Master Gardener WSU County Extension, SJI Edited by Dr. Tom Schultz
Copyright © Mar.21, 2006
Submitting Plant Specimens for Disease Diagnosis http://www2.ca.uky
.edu/agcollege/plantpathology/ ext_files/PPFShtml/PPFS-GEN-09.pdf
University of Kentucky Department of Plant Pathology Publications http://www2.ca.uky.
edu/agcollege/plantpathology/ extension/pubs.html