Plant Pathogens and Disease: General Introduction

G N Agrios, University of Florida, Gainesville, FL, USA

ª 2009 Elsevier Inc. All rights reserved.

Defining Statement Interactions of Pathogens, Plants, and Humans

Introduction: Causes of Disease in Plants Conclusion
Pathogens that Cause Disease in Plants Further Reading

Glossary monopartite A virus whose each particle contains the

disease cycle The chain of events involved in the life total nucleic acid of the virus.
cycle of a pathogen and in the development of the parenchyma Plant tissue consisting of thin-walled cells
disease caused by this pathogen. with intercellular spaces between them and synthesize
effectors Proteins produced by pathogens that act to or store foodstuffs.
suppress the defenses of the plant basal immune system phloem Tubelike cells of the plant conductive system
but in some cases trigger the defenses of the host plant. that carry sugar and other organic molecules from
elicitor A molecule coded by a gene for avirulence of a leaves to other parts of the plant.
pathogen, which, upon contact with the receptor sporulate Produce spores.
molecule of the corresponding plant gene for resistance, transgenic An individual that has been transformed
triggers the defense reaction of the plant that keeps it with and carries genes obtained from other organisms
resistant to the pathogen. and are expressed by these organisms.
haustoria Feeding organs of some fungi and parasitic vector A specific insect, fungus, nematode, and so on,
higher plants that enter the host tissues and absorb that can acquire and transmit a specific pathogen from
nutrients from host cells. an infected to a healthy host plant.
life cycle The stages in the growth and development of virulence Relative ability of a pathogen to cause
an organism that occur between two successive disease on a given host plant.
occurrences of the same stage, for example, sporulation
of the organism.

Abbreviations HR hypersensitive response

DPM Doctor of Plant Medicine TMV tobacco mosaic virus

Defining Statement caused by pathogens (Figure 1). In all organisms, diseases

and injury are caused by internal or external abiotic, that is,
Plant pathogens are mostly microorganisms, such as bac- environmental factors, such as nutritional deficiencies,
teria, viruses, fungi, nematodes, and protozoa, and some freezing temperatures, droughts, floods, pollution, and so
parasitic plants and algae. They attack plants, obtain on. (Figure 2), or by pathogens. Although abiotic factors
nutrients, and cause disease by releasing enzymes, toxins, may cause damage to plants over extensive areas, they differ
and so on. Successful infections depend on the interaction from pathogens in that they (1) do not multiply and (2) do
of appropriate genes present in pathogens and in plants. not spread from plant to plant. As a result, abiotic factors,
Plant diseases cause financial losses, hunger and famines, although they cause damage to plants, do so much less
food poisoning, and extinction of plant species. frequently than plant pathogens, they are unpredictable
and, usually, are easy to diagnose but difficult to control.
Introduction: Causes of Disease in Plants
Pathogens that Cause Disease in Plants
Plants, like humans and animals, are injured or damaged by
abiotic, that is, environmental, factor, and by biotic ones, Pathogens are living entities, mostly certain microorganisms,
such as various pests, for example, insects, and by diseases such as certain fungi, prokaryotes such as bacteria and

613
614 Pathogenesis | Plant Pathogens and Disease: General Introduction

Proteins synthesized

Vitamins and
hormones formed

Shoot blight
Leaf blight

Reproduction and
storage of strach,
protiens, and fats

Transpiration

Fruit spot

Fruit rot

Carbon
dioxide Leaf spot
Light
Canker
Tanslocation
of water
and minerals

Food
translocation Wilt
Vascular wilt

Photosynthesis
(food manufacture) Crown gall

Sugars and nitrogen

form amino acids Root rot

Uptake of water
and minerals
Protein
synthesized
Figure 1 Typical plant of which the left half is showing the basic functions of its main organs, whereas the right half is showing the
various symptoms of infection by pathogens and their interference with the basic functions. Modified from Agrios GN (2005) Plant
Pathology, 5th edn., p. 6. Burlington, MA: Elsevier/Academic Press.

mollicutes, viruses, protozoa, nematodes, and so on, which cause injury in plants closely resembling disease, as does the
can attack other organisms and cause disease. In addition to pressure for food and water, and for space, brought on by
these pathogens, diseases in plants can also be caused by numerous weeds. The vast majority of diseases in plants are
several plants that parasitize other plants, and by some para- caused by the same groups of pathogenic microorganisms as
sitic green algae. However, parasitic plants and parasitic algae those that cause disease in animals and humans.
cause only a few important diseases to plants. Furthermore, Plant pathogens vary considerably in size (Figures 3
many pests, such as aphids, mites, and other microorganisms, and 4), shape, and method of multiplication (Figure 4).
 Pathogenesis | Plant Pathogens and Disease: General Introduction 615

(a) (b)

(c) (d)

(e) (f)

Figure 2 Symptoms of plants and plant organs showing the effect of abiotic factors on plants. (a) Effect of freezing
temperatures on the trunk of apple tree. (b) Young barley plants show yellowing and dieback of leaf tips caused by nitrogen
(N) deficiency. (c) Young pear fruit showing symptoms of boron deficiency. (d) Tomato fruit showing symptoms of calcium
deficiency. (e) Tobacco leaf showing typical symptoms of ozone injury. (f). Citrus seedlings showing yellowing, stunting, and
some have been killed by improper fumigation with pesticides. Photos: (a), (b), and (e) courtesy of USDA; (d), Clemson
University; (f), JH Graham, University of Florida. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington,
MA: Elsevier/Academic Press, (a) p. 362, (b) p. 374, (c) p. 376, (e) p. 370, (f) p. 319.

Like all pathogens, those affecting plants vary consider- and roots; blights, that is, the sudden death of leaves and
ably in host specificity. Some pathogens are able to infect young shoots; cankers, that is, necrotic patches of bark and
all or most plants belonging to one species, to a genus, or under-the-bark tissues of branches and trunks; vascular
to several plant families, whereas other pathogens can wilts due to blockage by the pathogen of xylem vessels;
infect only one or a few varieties within one plant species. rots of roots, stems, and fruit; galls on stems or roots; pro-
Plant pathogens cause disease in plants by entering and liferation, that is, excessive branching of shoots and roots;
invading plant tissues and by absorbing food stuffs from mosaics; stunting; decline; and death of branches or entire
the host cells for their own growth, multiplication, and plants. Symptoms vary considerably depending on the kind
spread. By entering the plant, pathogens disturb the struc- of pathogens that cause them, and they may vary in sever-
tural integrity and the metabolism of plant cells and tissues ity, ranging from insignificant to death of the entire plant. It
through enzymes, toxins, growth regulators, and other has been estimated that plant pathogens are responsible for
active substances they secrete (Figure 5). Some pathogens a 16% loss of the attainable annual world crop production,
may also cause disease by growing and multiplying in the estimated at 1.2–1.3 trillion dollars.
xylem and phloem vessels of plants, thereby blocking the
upward movement of water and minerals and the down-
Fungi
ward translocation of sugars (Figures 1 and 5).
Plants infected with pathogens develop a variety of It is estimated that there are approximately 1.5 million
symptoms, such as necrotic spots on leaves, stems, fruit, species of fungi on earth, of which 70 000 species are
616 Pathogenesis | Plant Pathogens and Disease: General Introduction

5µ
Protozoon
4

2
Head of
nematode
1

Beet yellows virus

Tabacco mosaic virus
Wheat striate mosaic virus Fungus
Cucumber mosaic virus (mycelium)
Tobacco necrosis satellite virus
Hemoglobin molecule

Viroids

Mollicutes

Nucleus
Cell wall
Bacterium
Nucleolus

Figure 3 Morphology and ways of multiplication of some of the groups of plant pathogens. Reproduced from Agrios GN (2005) Plant
Pathology, 5th edn., p. 8. Burlington, MA: Elsevier/Academic Press.

known and have been studied. Fungi comprise the king- only by remaining in constant association with their living
dom Fungi and constitute an independent group of host plants. Other pathogenic fungi are nonobligate para-
organisms of equal rank to that of plants and animals. sites that either require a living host plant for part of their
Most pathogens of plants are fungi. They cause the life cycles, but are able to complete their cycles on dead
majority (approximately 70%) of all plant diseases. More organic matter, or are able to grow and multiply on dead
than 10 000 species of the known 70 000 fungal species can organic matter (necrotrophs) as well as on living plants.
cause disease in plants. Fungi are eukaryotic organisms The vast majority of fungi provide an indispensable
with one or more chromosomes of DNA contained in service to humanity and the world, by decomposing the
well-organized nuclei. Fungal cells, however, contain vast amounts of plant, especially wood, and animal tissue
only one copy of each type of chromosome (haploid), produced and dying each year. A number of plant patho-
whereas in most eukaryotes the nuclei are diploid, con- genic fungi, such as Penicillium, produce penicillin and other
taining two copies of chromosomes. Fungal cell walls antibiotics useful to humans and domestic animals. In addi-
contain chitin and glucans. tion, several fungi, such as Trichoderma, species of Aspergillus,
Some of the plant pathogenic fungi are biotrophs and so on, provide various levels of commercially used
(obligate parasites) because they can grow and multiply biological control of soilborne plant pathogenic fungi.
Figure 4 Schematic diagram, for comparison purposes only, of the shapes and sizes of certain plant pathogens in relation to a plant
cell. Note: Bacteria, mollicutes, and protozoa are not found in nucleated, living plant cells. Reproduced from Agrios GN (2005) Plant
Pathology, 5th edn., p. 7. Burlington, MA: Elsevier/Academic Press.

(a) (c)

(b)

Figure 5 Mechanisms that pathogens use to attack plants. (a) Certain enzymes, such as pectinases and cellulases, which break down
plant tissue. (b) Toxins, which kill plant cells, near or at a distance from the location of the pathogen on or in the plant. (c) Overgrowths
or galls, which results in infected cells producing or drawing nutrients used by the pathogen. Photo: (b) courtesy of RJ MacGovern,
Department of Plant Pathology, University of Florida. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA:
Elsevier/Academic Press, (a) and (b) p. 51, (c) p. 663.
618 Pathogenesis | Plant Pathogens and Disease: General Introduction

Plant pathogenic fungi elimination of several oak species in the Pacific Coast states
Many plant pathogenic fungi are famous for damaging or by the recent outbreak of ‘oak sudden death’ disease caused
eliminating plants from nature. This happened, for exam- by the oomycete Phytophthora ramorum; and others. Certain
ple, with chestnut blight disease (Figures 6(a)–6(c)), caused species of some fungi, for example, special forms of Fusarium
by the fungus Cryphonectria parasitica, which, within about 20 lycopersici, the cause of Fusarium vascular wilt in several
years from the time (1904) the pathogen was brought to crops, for example, F. oxysporum f. sp. cubense, the cause of
North America, destroyed about 4 billion chestnut trees, banana wilt (Panama disease). Once such pathogens are
and almost completely eliminated and threatened with introduced into a field, they become permanent inhabitants
extinction, the American chestnut in North America. A of the field and destroy the plant they infect, in this case,
similar direction of destruction, elimination, and possible banana, for ever after.
extinction of the American elm in North America has been A number of plant pathogenic fungi, such as Aspergillus,
followed by the Dutch elm disease (Figures 6(d)–6(f)), Penicillium, Claviceps, Fusarium, Trichoderma, and so on, pro-
caused by the fungus Ophiostoma nova-ulmi; near elimination duce, in plant seeds infected by these fungi, extremely
of several species of red oak in the Northeastern United poisonous toxins, called mycotoxins (Figure 7), some of
States by the oak wilt fungus, Ceratocystis fagacearum; near which are the most potent carcinogens known. Every year, a

(a) (c)

(d)

(b)

Figure 6 (Continued)
 Pathogenesis | Plant Pathogens and Disease: General Introduction 619

(e)

(f)

Figure 6 Two catastrophic diseases of trees, chestnut blight (Figures 6(a)–6(c)) and Dutch elm disease (Figures 6(d)–6(f)), are
caused by two different fungi that enter the vascular tissues of the trees and inhibit the passage of water to the tree branches.
(a) Chestnut blight disease, caused by the fungus Cryphonectria, has killed nearly all 4 billion American chestnut trees in the plant’s
natural range (a) with its spores spreading from diseased to healthy tree, infecting branches and the trunks of small trees, causing
cankers (b) on tree branches and small trunks, and blocking the passage of water beyond the canker. Multiple cankers on the tree
result in the death of the whole trees (c). Photos: (b) courtesy of WL MacDonald, West Virginia University; (c), RL Anderson, US Forest
Service. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic Press, (a), (b), and (c) p. 33.
(b) Dutch elm disease, caused by the fungus Ophiostoma, has killed almost all American elm trees along streets of cities and towns in
North America and many of those growing in forests. The fungus spores, which spread from tree to tree by two small beetles, grow
inside and clog the xylem vessels of twigs and branches, which wilt (d), then the infected branches die (e) and soon after, whole trees
die (f). Photos: (d) courtesy of RJ Stipes; (e), R.L. Anderson; (f), EL Barnard, Florida Forest Service. Reproduced from Agrios GN
(2005) Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic Press, (d), (e), and (f) p. 34.

number of these adversely affect humans and animals in Reproduction

many parts of the world. On the other hand, many mycor- Fungi reproduce primarily by means of spores (Figures 4
rhizal fungi live symbiotically with the plants they infect and 8), which may consist of one or a few cells. In different
and seem to improve the resistance of the host plants to fungi, spores may form asexually, like buds on a twig, or
other pathogens, whereas the endophytic fungi Epichloe sexually, as the result of a sexual fertilization. Asexual
(Acremonium) and others, infecting many grasses, are poiso- spores in some fungi are produced inside containers called
nous to animals that consume such grasses (Figure 7(c)). sporangia and such spores are called sporangiospores. In
some of these fungi, and in the oomycetes, the sporangios-
pores have flagella with which they can swim and are
Morphology: Shapes and sizes of fungi therefore called zoosporangiospores or zoospores. In
Most fungi have a branching filamentous body called most fungi, the asexual spores are called conidia and are
mycelium (Figures 2, 3, and 8). Mycelium produces produced by the cutting off of terminal or lateral cells from
numerous branches that grow outward in a radial fashion special hyphae called conidiophores. In some fungi, con-
and produces a colony (Figure 2). Each branch of the idia and condiophores are produced naked on the
mycelium, called a hypha, is tubular and generally is of mycelium, whereas in others they are produced inside
uniform thickness (1–5 mm in diameter). In some fungi, thick-walled containers called pycnidia. Sexual reproduc-
the mycelium is more or less a continuous tube containing tion occurs in most but apparently not all fungi. In sexual
many nuclei; in others, the mycelium is partitioned into reproduction, generally, two cells of similar or dissimilar
cells by cross-walls, called septa, with each cell containing size and appearance fuse to produce a zygote. Depending
one or two nuclei. The length of the mycelium in some on the group of fungi, the zygote is produced and under-
fungi is only a few millimeters, whereas in others it may goes meiosis inside a germinating zygospore (in
be several centimeters long. zygomycetes), an ascus (in ascomycetes), or a basidium
620 Pathogenesis | Plant Pathogens and Disease: General Introduction

(a) (b)

(c)

Figure 7 (a) Head of rye plant showing sclerotia (ergots) of the fungus Claviceps purpurea. (b) Cracked corn kernels attacked by fungi
that produce mycotoxins. (c) Fluorescent mycelium of an endophytic fungus in a grass plant in which it produces mycotoxins. Photos:
(a) courtesy of IR Evans, Canada; (b), RW Stack, ND State University; (c), A DeLucca. Reproduced from Agrios GN (2005) Plant
Pathology, 5th edn. Burlington, MA: Elsevier/Academic Press, (a) p. 38, (b) and (c) p. 40.

(in basidiomycetes) (Figure 8). In some fungi, no sexual and 12(c) show the symptoms of stem rust of wheat, caused
spores have been found and these are known as deuter- by the basidiomycete Puccinia graminis.
omycetes or imperfect fungi. Some appear to never
produce any kind of spores and are known as sterile fungi. Classification
As a result of the different types of mycelium, spores, The fungal pathogens of plants include some microorgan-
and spore containers, each group of fungi follows a slightly isms, the Myxomycota, Plasmodiophoromycota, and
different life cycle reflecting these differences (Figures 8 Oomycota, that are now known not to be fungi but to
and 9(a)). When the life cycle also incorporates and shows belong to different kingdoms of organisms. However,
the changes in the host plant, the sequence of events is then these organisms are similar to fungi and they continue
called a disease cycle and is much more informative, to be studied along with the fungi and the diseases they
regarding the interaction of each pathogen with its host, cause. The following is a sketchy classification of fungal
than the life cycle alone. Figure 9(b) shows a generalized and fungal-like pathogens of plants.
disease cycle applicable to any host–pathogen combination, Fungal-like organisms
and lists primarily the events taking place in the fungus and Kingdom: Protozoa
in the plant after the pathogen comes in contact with the Phylum: Myxomycota – produce a plasmodium
host plant. The primary disease cycle after overwintering instead of mycelium; they are the surface slime molds.
includes the sexual as well as asexual spores. In contrast, Cause few plant diseases.
secondary disease cycles develop from pathogens produced Phylum: Plasmodiophoromycota – cause endoparasitic
from the primary inoculum and usually contain only or slime mold diseases. Cause a few diseases of importance,
mostly asexual spores. Figure 10 shows the symptoms and for example, clubroot of cabbage.
most of the stages of the disease cycle of the potato late Kingdom: Chromista
blight disease caused by the oomycete Phytophthora infestans. Phylum: Oomycota – produce mycelium that has no
Figure 11 shows a fairly detailed disease cycle of the apple cross-walls; their cell walls are composed of cellulose and
scab disease caused by the ascomycete Venturia inaequalis, the amino acid hydroxyproline, not chitin; produce oospores
Figure 12(a) shows the disease cycle, and Figures 12(b) and zoospores; cause many root rots, seedling diseases, foliar
 Pathogenesis | Plant Pathogens and Disease: General Introduction 621

Figure 8 Representative spores and fruiting bodies of the fungal-like oomycetes and of the main groups of fungi. Reproduced from
Agrios GN (2005) Plant Pathology, 5th edn., p. 389. Burlington, MA: Elsevier/Academic Press.

blights, and the downy mildews. They include the following Phylum: Ascomycota – Recent, 2007, taxonomic studies
extremely important pathogens: have placed most of the 32 000 species of Ascomycetes in
Pythium, the cause of many root, stem, and fruit rots. the subphylum Pezizomycotina. Under their new umbrella,
Phytophthora, the cause of many blights, root and tuber the species and genera are, of course, similar/identical to
rots, and of cankers, declines, and death of many trees. Ascomycota, but the Pezizomycotina have septate hyphae,
Plasmopara, the cause downy mildews. the single septum having a single pore that divides the
hyphae into hyphal compartments or cells, and also have
Woronin bodies, which are specialized vesicles that seal the
The true fungi septal pore in response to cellular damage. The Woronin
Kingdom: Fungi body consists of HEX-1 protein that self-assembles and
Phylum: Chytridiomycota – have round or limited forms the solid form of the vesicle. The Pezizomycotina,
elongated nonseptate mycelium, restricted to the host like all Ascomycetes, have mycelium that has cross-walls;
plant, and, alone among the fungi, produce motile zoos- produce sexual spores (ascospores) within sacs (asci) (e.g.,
pores and survive as sporangia. Cause few plant diseases, Figure 7) that are either naked or contained in fruiting
for example, wart of potato. structures of different shapes, namely, cleistothecia, peri-
Phylum: Zygomycota – Order: Mucorales: no zoos- thecia, and apothecia; produce asexual spores (conidia) on
pores; produce conidia in sporangia; mycelium nonseptate; naked hyphae or in containers (pycnidia) or other struc-
survive as zygospores; most are saprophytic but a few are tures; and they cause the most plant diseases (leaf, stem, and
weak plant pathogens causing bread molds (Figure 13(b)) fruit spots and blights, root rots, fruit rots, cankers, vascular
and fruit rots (Figures 3(b) and 3(c)) in storage. wilts, seed rots, etc.).
Order: Glomales: Form vascular – arbuscular mycor- The new classification scheme rejects the previous taxa
rhizae within roots of host plants. of Discomycetes – apothecial fungi, Pyrenomycetes –
622 Pathogenesis | Plant Pathogens and Disease: General Introduction

(a)

(b)

Figure 9 (a) Schematic representation of the generalized life cycles of the main groups of plant pathogenic fungi. (b) Stages of
development of a disease cycle. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic
Press, 8(a) p. 402, 8(b) p. 80.

perithecial fungi, Plectomycetes – cleistothecial fungi, and this volume, we will continue to use the already estab-
Loculoascomycetes – ascostromatal fungi. Instead they lished scheme, with the exception of a few names, which
recommend the use of 10, and possibly 12 taxa in place we will use here. So,
of the 4 in the previous scheme. Because of the newness of Pezizomycotina – have mycelium that has cross-
the new terminology and the fact that the literature has so walls; produce sexual spores (ascospores) within sacs
far used the old system, for the purpose of the audience of (asci) (e.g., Figure 11) that are either naked or contained
 Pathogenesis | Plant Pathogens and Disease: General Introduction 623

(a) (b)

(c)

(d)
Sporangium

Sporangium

Germination Sporangium

Oospore

Sporangiophore
on infected
Karyogamy seedling
Oogonium Sporangium Zoospores

Sporangiophore
Germ tube
on infected tuber
Antheridium (in spring)

Sporangiophore Sporangia and

Meiosis on leaf zoospores infect leaf
Oogonium Mycelium
from tuber
infects
seedling
Antheridium

Infected leaf
Infected
tuber
Zoospores
infect tuber
Sexual reproduction,
extremely rare in nature

Infected
plant

Figure 10 Symptoms and the disease cycle of the late blight disease of potato caused by the oomycete Phytophthora infestans.
(a) Infected potato leaf showing fungal mycelium, sporangia, and sporangiophores. (b) Potato plant resistant to late blight looks almost
unaffected (right) compared to a susceptible potato plant which has been killed by the fungus. (c) Rotting of a potato tuber following
infection by the late blight. (d) Disease cycle of late blight of potato caused by the oomycete P. infestans. Photos: (a) and (d)
courtesy of PWeingarten, University of Florida; (b), Cornell University; (c), USDA. Reproduced from Agrios GN (2005) Plant Pathology,
5th edn. Burlington, MA: Elsevier/Academic Press, (a) p. 20, ((b) and (c)) p. 21, (d), p. 425.
Figure 11 Disease cycle of apple scab disease caused by the ascomycete Venturia inaequalis. Reproduced from Agrios GN (2005)
Plant Pathology, 5th edn., p. 506. Burlington, MA: Elsevier/Academic Press.

(a) (b)

(c) (d)

Figure 12 Early (a) and later (b) stages of lettuce infection with the ascomycete Sclerotinia sclerotiorum that resulted in total loss of the
lettuce crop. (c) Strawberry and (d) tomato fruit showing, at first, lesions and later total rotting following infection of the fruit by the
ascomycete Colletotrichum as the fruits approach maturity. Photos: (a) and (b) courtesy of KV Subbarao, University of California,
Salinas; (c), L Legard; (d), RJ MacGovern, both University of Florida. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn.
Burlington, MA: Elsevier/Academic Press, (a) and (b) p. 271, (c) p. 490, (d) p. 489.
 Pathogenesis | Plant Pathogens and Disease: General Introduction 625

(a) (b)

(c)

Figure 13 (a) Bread mold caused by the fungus Penicillium. (b) Strawberries rotted by the fungus Rhizopus. (c) Postharvest rotting
of tomatoes by different fungi. Photos: (b) and (c) courtesy of University of Florida. Reproduced from Agrios GN (2005) Plant Pathology,
5th edn. Burlington, MA: Elsevier/Academic Press, (a) p. 554, (b) p. 13, (c) p. 566.

in fruiting structures of different shapes, namely, cleis- Sclerotinia sclerotiorum, causing the white rot or watery
tothecia, perithecia, and apothecia; produce asexual soft rot of vegetables (Figures 14(a) and 14(b)).
spores (conidia) on naked hyphae or in containers (pyc- Athelia (Sclerotium) and Thanatephorus (Rizoctonia), caus-
nidia) or other structures; cause most plant diseases (leaf, ing root and stem rots of vegetables and fleshy ornamentals
stem, and fruit spots and blights, root rots, fruit rots, and soft rots of fleshy leaves and fruits.
cankers, vascular wilts, seed rots, etc.). Phylum: Basidiomycota – have mycelium, often with
Important Plant Pathogenic Ascomycetes: binucleate cells, sexual spores (basidiospores) produced
Taphrinales, causing peach leaf curl and plum pockets. externally on a clublike structure called a basidium;
Erisyphales, causing powdery mildews. some of them produce several types of spores and
Pyrenomycetes, Ascomycetes with perithecia or spore-bearing structures, namely, basidiospores on basi-
cleistothecia. dia, spermatia in spermagonia; aeciospores in aecia;
Claviceps, causing ergot (Figure 7). uredospores in uredia; and teliospoes in telia; rusts are
Gibberella (foot rot and stem rot, of corn), Epichloe, very serious diseases of grain (Figures 12(a) and 12(b)),
Balansia, Adkinsonella: endophytic on grasses and of beans and soybeans, and other crops, and of trees.
sedges apple (Figure 7(c)). Basidiomycetes also include the smuts of grain crops
Glomerella (Colletotrichum sp.), causing many anthrac- (Figures 12(a) and 12(b)), and the root rots, wood rots,
nose diseases (Figures 14(c) and 14(d)). and decays of trees (Figures 12(c)–12(e)) and timber.
Ophiostoma, causing the Dutch elm disease (Figure 6). Ustilaginales (the smut fungi);
Cryphonectria, causing chestnut blight (Figure 6). Ustilago, causing corn smut and loose smut of grains
Loculoascomycetes, causing Ascostromata. Asci within (Figure 15(a)).
locules (cavities). Tilletia, causing covered smut (Figure 15(b)) or bunt
V. inaequalis, causing apple scab (Figure 11). of wheat, and Karnal bunt of wheat.
Discomycetes, causing Ascomycetes with apothecia Uredinales (the rust fungi)
626 Pathogenesis | Plant Pathogens and Disease: General Introduction

(a)

(b) (c)

Figure 14 (a) Disease cycle of stem rust of wheat caused by the basidiomycete Puccinia graminis. Notice the variety and sequence of
the spores and fruiting bodies, the secondary disease cycle at bottom center, and the need for two alternate hosts, wheat and
barberry. (b) Severe infection of wheat by the wheat stem rust fungus, (c) Empty, poor quality kernel from rust-infected wheat plant
(left), and wheat kernels from healthy plant. Photos: (b) courtesy of CIMMYT; (c), USDA. Reproduced from Agrios GN (2005) Plant
Pathology, 5th edn. Burlington, MA: Elsevier/Academic Press, (a) p. 570, (b) p. 13, (c) p. 566.

Puccinia, causing the devastating rust diseases of cereals, and Prokaryotes: Bacteria and Mollicutes
other plants. Cronartium, the rust of pine trees. Gymnosporangium,
the cedar-apple rust. Hemileia, the coffee rust. Approximately 100 species of bacteria and an unknown
Agaricales: The mushrooms; many are mycorrhizal number of mollicutes cause many severe diseases and
fungi, and many, for example, Armillaria, cause losses of losses (Figures 17 and 18) in plants. Bacteria and molli-
about 1 billion dollars in the United States every year. cutes are prokaryotic organisms, that is, they do not have
Ceratobasidiales, causing root rots and decays of trees. organized nuclei bound by a membrane and their DNA is
Aphyllophorales, causing wood rots and decays organized as a single chromosome present in an area of
(Figure 16). the cytoplasm called nucleoid. They also have circular
 Pathogenesis | Plant Pathogens and Disease: General Introduction 627

(a) (b)

Figure 15 (a). Field symptoms of barley heads infected with loose smut fungus Ustilago. (b) Kernels of wheat infected with and
carrying teliospores of the cover smut fungus Tilletia compared with a few healthy whitish kernels. Photos: (a) courtesy of P Thomas;
(b), PE Lipps, Ohio State University. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic
Press, (a) and (b) p. 12.

(a) (b) (c)

Figure 16 Three stages or types of rotting and decay of trees by wood rotting fungi. Photos: (a) and (c) courtesy of EL Barnard, Florida
Department of Agriculture and Forestry; (b), University of Florida. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn.
Burlington, MA: Elsevier/Academic Press, (a) p. 608, (b) p. 607, (c) p. 609.

pieces of DNA contained in plasmids and mitochondria. Most bacteria attack tissues at or near the surface of a host
Most plant pathogenic bacteria are necrotrophs, that is, plant but the fastidious bacteria grow in the xylem vessels
they are facultative saprophytes, and can be grown on of the plant. The mollicute phytoplasmas and spiroplas-
synthetic nutrient media, but survive and multiply best in mas, as well as some rather fastidious bacteria, grow only
contact with their host plants. Some fastidious vascular in the phloem of the plant.
bacteria, however, survive in nature only inside their host Plant pathogenic mollicutes also survive and multiply
plants, in either the xylem vessels or the phloem sieve in nature only inside the phloem sieve tubes of their
tubes. living hosts (Figure 19). Mollicutes of only one genus,
It is estimated that the plant pathogenic bacteria com- Spiroplasma, can be grown in culture; all others, usually
prise between 30 and 100 species. Most plant pathogenic referred to as phytoplasmas, can be maintained but do not
bacteria are facultative parasites or necrotrophs and are multiply on nutrient media.
easy to grow on artificial media. Several, known as fasti- One rather common plant pathogenic bacterium,
dious bacteria, are difficult to grow on common nutrient Agrobacterium tumefaciens, which causes the crown gall dis-
media but can be cultured on specialized complex media. ease (Figure 4(c)) in many plants but can survive freely in
628 Pathogenesis | Plant Pathogens and Disease: General Introduction

(a) (b)

(c)

(d)

(e)

Figure 17 (a) Typical rod-shaped plant pathogenic bacteria. (b) Bacterium with peritrichous flagella. (c) Mollicutes in a phloem vessel.
(d) A grapevine with scorched leaves caused by the xylem-limited bacterium Xylella. Photos: (a) courtesy of Roos and Hatting, The
Netherlands; (b), Oregon State University; (c), JE Worley, USDA; (d), DL Hopkins, University of Florida; (e), E Alves, Federal University,
Lavras, Brazil. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic Press, (b) p. 645,
(c) p. 9, (d) p. 680, (e) p. 681.
 Pathogenesis | Plant Pathogens and Disease: General Introduction 629

Figure 18 (a) Relative morphology of the most important genera of plant pathogenic bacteria and the symptoms they cause on their
host plants. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn., p. 620. Burlington, MA: Elsevier/Academic Press.

the soil for more than a year after infection, has been Reproduction
proven to be a natural genetic engineer and the bacterium Plant pathogenic bacteria and mollicutes reproduce
and its purified DNA have been used to multiply and to asexually by fission. That is, following the replication of
transfer DNA from some plants and other organisms into the DNA in the bacterium, the cell wall of the bacterium
particular plants. This was and still is one of the most also divides, separating the two halves of the bacterium
important steps and contributions of plant pathology to into two new identical bacteria (Figure 2). Any plasmids
the huge development of genetic engineering of plants present in the bacterium replicate independent of the
and of plant biotechnology. chromosomal DNA, but may code for proteins that play
an important role in the development of disease. Although
Morphology bacteria do not have a typical sexual reproduction,
Most plant pathogenic bacteria are rod-shaped, consider- genetic change in their DNA is introduced by conjugation
ably smaller than fungi (Figures 4, 17, and 19), ranging of two identical or different bacteria, during which seg-
from 0.5–1.0 mm in diameter to 0.6–3.5 mm in length. In ments of the DNA from one bacterium are transferred to
some bacteria, and in older cultures, the bacteria may be the other bacterium; by bacteria-infecting viruses (bacter-
longer and they may be branched. Bacteria of the genus iophage); and most commonly by mutation, that is,
Streptomyces are filamentous. Most species of bacteria have mistakes that occur during replication of the DNA.
one or more flagella that can be found at the polar ends or
cover the entire surface of the bacteria (Figure 19(b)). Classification of plant pathogenic bacteria and
Although bacterial cells are contained by a cell wall, mollicutes
which gives them the typical rod shape, plant pathogenic Because most bacteria lack distinctive morphological
mollicutes lack a cell wall and thereby take on a shape characteristics, their taxonomy and names are less
that is spherical, tubular, or polymorphic (Figures 3, 4, stable than in other organisms. Scientists, however,
and 17). have developed an array of diagnostic techniques for
630 Pathogenesis | Plant Pathogens and Disease: General Introduction

(a)

(d)

(b)

(c)

Figure 19 (a) A young apple orchard destroyed by fire blight caused by the bacterium Erwinia amylovora. (b) Infected apple fruit
exuding droplets of fire blight bacteria. (c). Bacterial soft rot of vegetables, for example, cabbage, caused by at least three species of
bacteria. (d) Symptoms of citrus canker on a young stem. Photos: (a) and (b) courtesy of T Van Der Zwet, USDA; (c), Department of Plant
Pathology, University of Florida; (d), Division of Plant Industry, Florida Department of Agriculture. Reproduced from Agrios GN (2005)
Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic Press, (a) p. 644, (b) p. 645, (c) p. 658, (d) p. 673.

bacteria, including serological and molecular techniques Erwinia – causing fire blight of apples and pears, nur-
that are very effective in bacterial taxonomy and sery stock, vascular wilts, and soft rots of fleshy fruits,
classification. vegetables, and ornamentals (Figure 17(b)).
The most common prokaryotic pathogens of plants Pseudomonas – causing numerous leaf spots, blights,
can be classified approximately as shown below. The wilts, and so on. (Figure 17(a)).
shapes of the bacteria and the kinds of plant symptoms Ralstonia – causes wilt of solanaceous crops.
caused by most important bacteria are shown in Figures Xanthomonas – causes leaf spots, blights, and citrus
5 and 18. canker.
Kingdom: Prokaryotae Family: Rhizobiaceae
Bacteria: Have a cell membrane and a cell wall. Genus: Agrobacterium, A. tumefaciens (Figure 5(c)) caus-
Gram-negative bacteria ing the crown gall disease.
Family: Enterobacteriaceae. Selected important genera. Family: Still unknown;
 Pathogenesis | Plant Pathogens and Disease: General Introduction 631

Genus: Xylella fastidiosa. Xylem - inhabiting, causing the spread of the virus and produce numerous small local
leaf scorch and dieback in trees (Figures 17(c)–17(e). lesions (Figure 21(a)), which, usually, but not always, keep
Gram-positive bacteria the virus limited to the lesions and do not allow it to spread
Genus: Clavibacter – causing bacterial wilts in alfalfa, systemically through the plant and cause systemic symp-
potato, and tomato. toms. When the plant is susceptible to the virus and
Genus: Streptomyces sp. – causing the common scab of becomes infected, the virus may spread internally through
potato. Antibiotics. parts of, or more commonly throughout, the plant and the
Mollicutes: Have cell membrane but not cell wall plant then develops symptoms appearing as leaf mosaic
Family: Still unknown (Figures 21(c) and 21(d)), as necrotic leaves and poor
Genus: Phytoplasma (Figure 17(c)), causes numerous growth of the plant, as internal necrotic veins
yellows, proliferation, and decline diseases in trees and (Figure 21(i)), as yellowing of leaves and severe stunting
some annuals. of the plant (Figures 21(f) and 21(h)), as ringlike or amor-
Family: Spiroplasmataceae phous, discolored, bumpy or necrotic patches on fruit and
Genus: Spiroplasma, causes corn stunt, citrus stubborn seeds (Figure 21(g)), and many others. In some cases,
disease. viruses also produce internal symptoms in the epidermal
cells of the leaves of their hosts, appearing as amorphous or
crystalline inclusions (Figure 21(b)) in the cytoplasm or the
Viruses and Viroids nucleus of the plant cells. The inclusions are characteristic
More than 2000 viruses causing disease in plants have been of the genus of the virus and in many cases are used in the
identified and their properties have been studied. Viruses detection and identification of the virus.
are the smallest and simplest pathogens. Although viroids Although plant viruses are not known to infect humans
are much smaller and simpler than even viruses (Figures 20 and animals – with the exception of some of their insect
and 21), the ability of viroids and viruses to cause disease vectors – the study of plant viruses, and particularly the
and losses from disease is second only to those of fungal tobacco mosaic virus (TMV) (Figures 20(a) and 22(a)),
pathogens. Plant viruses cause a wide variety of symptoms causing the tobacco mosaic disease, has contributed immen-
on the plants they infect. When plants infected with the sely to our knowledge and understanding in several areas of
virus are more or less resistant to the virus, they may limit biology, genetics, plant pathology, plant and animal science,

(a)

(b)

Figure 20 (a) Diagram of shapes and relative sizes of plant viruses. AA&C, flexuous, threadlike virus; NA, nucleic acid; PS, protein
subunit. (a) Rigid rod-shaped virus: B-2 ¼ cross-section of the virus showing the arrangement of the nucleic acid. (b) Viroids (small
circles and equivalent length lines). The viroid that causes the coconut cadang-cadang (dying, dying). Photo (b) courtesy of JW Randles.
Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic Press, (a) p. 729, (b) p. 822.
632 Pathogenesis | Plant Pathogens and Disease: General Introduction

(a)
(b)

(c)
(d)

(e)

Figure 21 The shapes and relative sizes of plant viruses. (a) Rod-shaped virus, (b) filamentous flexuous virus, (c) isometric
polyhedral virus, (d) short, bacillus-like virus, and (e) geminivirus. Photos: (d) courtesy of EMJ Jaspars, The Netherlands; (f), E Hiebert,
Department of Plant Pathology, University of Florida. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA:
Elsevier/Academic Press, (a) p. 759, (c) p. 793, (d) p. 728, (e) p. 806.

medicine, biochemistry, and others. The reason for these shaped particles visible under the electron microscope.
successes of TMV is that this virus was the first to be Soon after, biochemical and genetic studies of TMV
detected and was hardy enough to withstand rough experi- showed that the protein subunits of the virus coat were
mental treatment. At first it appeared to investigators to be attached to the viral RNA and that the size and other
an infectious fluid, then to be or contain elongated particles properties of each protein were determined by the sequence
composed of infectious protein. It was soon shown to con- of the triplets of the four nucleotides along a stretch of the
tain not only protein but also nucleic acid (RNA). It was viral RNA containing three times as many nucleotides as
subsequently shown that infectivity resided in the nucleic the number of amino acids in the protein subunits. This
acid only while the protein played primarily a protective suggested that each amino acid of the protein subunit
role for the nucleic acid. Subsequently, it was studies of corresponded to three nucleotides of the RNA. The study
TMV that showed that the virus could be measured of the sequence of nucleotide triplets on the RNA to the
through counting the local lesions produced by a virus slight changes of amino acids in the coat protein subunits of
preparation and that the virus consisted of cylindrical rod- the various viral strains showed that each change in one or
 Pathogenesis | Plant Pathogens and Disease: General Introduction 633

more of the three nucleotides corresponded to always the survive in plant debris or outside plant cells but even they
same changes in amino acids of the protein subunits. This do not multiply there. Viruses and viroids are too small to
information, gained from the study of TMV, led to the be seen even with the compound microscope. Therefore,
discovery of the genetic code in viruses, which was later TMV was the first virus to be detected and identified by
shown to be the same in all organisms. several indirect techniques, which include kind of symp-
toms on infected plants, presence of characteristic inclusion
bodies in infected young tissues, transmission of symptoms
Characteristics of plant viruses and viroids to healthy plants by sap, grafting, specific insect and other
Viruses consist of one or a few molecules of nucleic acid vectors, and using serological tests or nucleic acid probe
(RNA or DNA but never of both being present in the same tests against known viruses. In sap from infected cells and in
virus), whereas viroids have only a small naked RNA purified preparations, most viruses and viroids can be seen
(Figure 20(b)). Plants are also affected by about 40–50 under the electron microscope.
viroids that have only an RNA that is about 10 times
smaller than that of viruses, is free or naked, and not con-
tained in a protein coat. Both, viruses and viroids can infect Morphology
plants, replicate themselves, and cause disease. Both viruses Nearly half of the known viruses are rigid rodlike,
and viroids are intracellular parasites. Only a few of them 15  300 nm (nanometers) (Figure 20(a)), or flexuous

(a) (b)

(c) (d)

(e)

Figure 22 (Continued)
634 Pathogenesis | Plant Pathogens and Disease: General Introduction

(f)
(h)

(g)

(i)

Figure 22 (a) Local lesions on leaf of Chenopodium (lamb’s quarters) plant hand-inoculated with sap from Potato virus Y-infected
plant. (b) Cellular inclusions produced by cells of some plant inoculated with certain viruses. (c) Cowpea leaf showing typical mosaic
symptoms. (d) Mosaic on tobacco mosaic virus-infected tobacco. (e) Mosaic and foliar malformation on pepper leaves infected with
Potato virus Y. (f) Rice plants infected with the grassy rice stunt virus and showing severe stunting, yellowing, and excessive tillering.
(g) Tomato fruit showing ringlike and other malformations caused by the Tomato spotted wilt virus. (h) Barley yellow dwarf symptoms of
varying severity on barley plants. (i) Potato tuber showing vein necrosis caused by the potato leaf roll virus. Photos: (b) courtesy of
R. Christi; (d), (e), and (g) courtesy of Department of Plant Pathology, University of Florida; (f), H Hibino; (h), WF Rochow, Cornell
University; (i), Department of Plant Pathology, University of Idaho. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn.
Burlington, MA: Elsevier/Academic Press, (a) p. 736, (b) p. 726, (c) p. 725, (d) p. 759, (e) p. 765, (f) p. 800, (g) p. 796, (h) p. 782, (i) p. 783.

threads about 11 nm in diameter by lengths varying from similar or different sized particles made of the same
480 to 2000 nm (Figure 20(b)). Almost as many viruses protein molecules (protein subunits). The surface of
are spherical, actually isometric or polyhedral, about viruses consists of a definite number of protein subunits,
17–100 nm in diameter (Figure 20(c)), and a few are which are spirally arranged in the elongated viruses and
bacillus-like rods 52  75 nm in diameter and packed on the sides of polyhedral viruses. In cross-sec-
300–380 nm long (Figure 20(d)) and some are monopar- tions, the elongated viruses appear as hollow tubes of
tite or bipartite geminiviruses (Figure 20(f)). Many spirally arranged protein subunits, with the nucleic acid
viruses have split genomes, that is, they consist of two or strand embedded between successive spirals of protein
more distinct nucleic acid strands, each encapsidated in subunits. In spherical viruses, the visible shell consists of
 Pathogenesis | Plant Pathogens and Disease: General Introduction 635

protein subunits, whereas the nucleic acid is inside the Single isometric particles – for example, Nanovirus
shell but it is not known how it is arranged. Viruses are (banana bunchy top virus)
not cells and have neither the cytoplasm nor the nucleus, Taxonomy (Grouping) of Viroids. (Figure 21(b)) –
but the rhabdoviruses and a few spherical viruses have an small, single-stranded, circular RNAs that replicate and
outer membrane composed of lipoprotein. cause disease in plants. There are no known viroids
infecting humans or animals.
Reproduction (replication)
Plant viruses and viroids replicate only within living plant
cells or protoplasts. The mechanisms of replication are quite
Parasitic Higher Plants
complex and the complexity varies depending on whether
the nucleic acid of the virus is RNA or DNA, single- or More than 2500 species of higher plants live parasitically
double-stranded, and positive or negative strand. Basically, on other higher plants. Relatively few of these plants,
in the replication of viruses and viroids, the infected plant however, affect and cause disease on cultivated higher
cell provides all the structural materials (nucleic acid and plants or on forest trees of commercial significance. The
proteins), the energy, and the machinery needed to make parasitic plants produce flowers and seeds like all plants.
more viruses, whereas the virus or viroids supply only the They belong to widely separated botanical families and
blueprint in the form of their nucleic acid. Virus (and viroid) vary greatly in dependence on their host plants. Some
replication is analogous to making copies of an original page parasitic plants (e.g., mistletoes) have chlorophyll but no
(the virus) with the help of a modern copier (the plant cell). roots; therefore, they depend on their hosts only for
Once new viral nucleic acid and viral protein are water and inorganic nutrients. Others (e.g., dodder)
produced in the cell, they are assembled into virus particles have little or no chlorophyll and no true roots; therefore,
(Figure 20). The virus particles, in turn, either accumulate in addition to water and inorganic nutrients, they also
in the cell singly or as parts of inclusion bodies, or they move depend on their hosts for photosynthetic products.
to other adjacent plant cells and, possibly through the Parasitic higher plants obtain water and nutrients from
phloem sieve elements, throughout the plant. their hosts by producing and sinking into the vascular
Classification
system of their host stems or roots food-absorbing organs
The main characteristics considered in the classification called haustoria. The following are the most important
of viruses are the nature and number of their nucleic acids parasitic higher plants and the botanical families to
and the shape and size of their particle(s). A very sketchy which they belong.
classification of plant viruses is as follows: Cuscutaceae – Cuscuta sp., the dodders (Figure 23)
Kingdom: Viruses Viscaceae – Arceuthobium, the dwarf mistletoes of the
RNA Viruses conifers
Single-stranded positive RNA Phoradendron, – the American true mistletoe of broad-
Rod-shaped particles – for example, Tobamovirus (TMV). leaved trees
Filamentous particles – for example, Potyvirus (potato Viscum – the European true mistletoes
virus Y). Orobanchaceae – Orobanche, the broomrapes of tobacco
Isometric particles – for example, Luteovirus (barley Scrophulariaceae – Striga, the witchweeds of many
yellow dwarf virus). monocotyledonous plants.
Single-stranded negative RNA – for example, potato Parasitic higher plants vary in size from a few millimeters
yellow dwarf virus. to a few centimeters in diameter and from 1 cm tall to
Buniaviridae – for example, Tospovirus (tomato spotted upright green plants more than 1 cm tall. Some, however,
wilt virus). are yellow or orange leafless vine strands that may grow to
Double-stranded RNA several meters in length and entwine around the stems of
Isometric viruses – for example, Reoviridae, many adjacent host plants. Parasitic higher plants reproduce
Phytoreovirus (wound tumor virus) by seeds. Seeds are disseminated to where host plants grow
DNA Viruses by wind, runoff water, birds, and cultivating equipment.
Double-stranded DNA Seeds of some parasitic plants are forcibly expelled to sig-
Isometric – for example, Caulimovirus (Cauliflower nificant distances (10 m or more). Parasitic higher plants
mosaic virus). overwinter on perennial hosts or as seeds on the hosts or
Nonenveloped bacilliform – for example, Badnavirus on the ground. In spring, the seeds germinate and the seed-
(rice tungro bacilliform virus). ling infects a new host plant. Control or management of
Single-stranded DNA parasitic higher plants depends on removing infected plants
Geminate (twin) particles – for example, Geminivirus carrying the parasites and avoiding bringing seeds of para-
(bean golden mosaic virus). sitic plants into new areas.
636 Pathogenesis | Plant Pathogens and Disease: General Introduction

(a) (b)

Figure 23 (a) Parasitic higher plant, dodder, of the species Cuscuta, has entwined itself around a pepper plant, which has been
overcome. (b) A small field of geraniums attacked by dodder. Photo courtesy of Department of Plant Pathology, University of Florida.
Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic Press, (a) p. 707.

Plants growing in the wild often suffer damage caused by (Figure 24(c)); devitalized root tips; excessive root stunt-
other plants growing over the first plants and covering them ing or branching; distortion of above-ground plant organs
so completely that they cannot get any sunlight. As a result and death of the plant (Figure 24(d)); and, in some cases
such plants decline and die prematurely (Figure 23). (Figures 25(a) and 25(b)), death of branches and whole
trees. The annual worldwide losses of crops caused by
nematodes on various crops are more than 80 billion
Parasitic Green Algae dollars.
Algae are the organisms, often microorganisms, other than
typical land plants, that can carry on photosynthesis. Algae
are sometimes considered as protists with chloroplasts. Morphology
Some algae, the so-called blue-green cyanobacteria, belong Plant parasitic nematodes are small and eel-shaped but
to the kingdom Prokaryotes but most of them, that is, the only 300–1000 mm long, with some up to 4 mm long 
rest, belong to the kingdom Chromista. Algae are the main 15–35 mm wide (Figure 26). The females of some of
producers of photosynthetic materials in aquatic ecosys- these nematodes become swollen at maturity and have
tems, including unstable systems such as muds, sands, and pear-shaped or spherical bodies. Nematode bodies are
intertidal aquatic habitats. Green algae are single-celled transparent, unsegmented, and have no legs or other
organisms that form colonies, or multicellular free-living appendages. Plant parasitic nematodes have a hollow
organisms, all of which have chlorophyll b. stylet or spear that is used to puncture holes in plant
Several algae are pathogenic of other organisms. For cells through which they withdraw nutrients from the
example, cyanobacteria cause the black band disease plant.
that leads to the bleaching and death of coral symbionts
of the algae. Many red algae are parasites on other, Reproduction
mostly related red algae. Colorless green algae of the Nematodes have well-developed reproductive systems
genus Prototheca cause skin infections in humans. Most that distinguish them as female and male nematodes. The
of the green algae live as endophytes of many hydro- females lay eggs, usually after fertilization by males but in
phytes to which they seem to cause little or no damage. some cases without fertilization. Many species lack males.
A few genera of green algae are parasitic on higher Nematode eggs hatch into juveniles that resemble the adult
plans. nematodes but are smaller. Juveniles grow in size and each
juvenile stage is terminated by a molt. All nematodes have
four juvenile stages, with the first molt usually occurring in
Nematodes
the egg. After the final molt, the nematodes differentiate
Nematodes are usually microscopic animals that are into males and females, and the new females can produce
wormlike in appearance but quite different taxonomically fertile eggs in the presence or absence of males. One life
from true worms. Most nematode species live freely in cycle from egg to egg may be completed within 2–4 weeks
fresh- or saltwater but numerous species attack and cause in favorable weather, longer in cooler temperatures. In
disease on humans and animals and several hundred spe- some nematodes, only the second-stage juvenile can infect
cies feed primarily on roots of plants and cause disease in a host plant, whereas in others all but the first juvenile and
them. Symptoms of infected plants include the following adult can infect. When the infective stages are produced,
(Figure 24): root lesions (Figure 24(a)) or root galls they must feed on a susceptible host plant or they will
 Pathogenesis | Plant Pathogens and Disease: General Introduction 637

Figure 24 Diagram of shapes and sizes of most of the important plant nematodes. Reproduced from Agrios GN (2005) Plant
Pathology, 5th edn., p. 828. Burlington, MA: Elsevier/Academic Press.

starve to death. In some species, however, some juveniles protozoan diseases of plants, although rare, are severe on
may dry up and remain quiescent, or the eggs may remain the infected plants, generally resulting in the collapse and
dormant in the soil for years. death of the plants. The plant pathogenic protozoa
invade and multiply in the phloem of diseased plants
Classification (Figure 27). They are mostly one-celled microscopic
The plant pathogenic nematodes can be classified as organisms that have flagella and typical nuclei.
follows (only a few important genera are listed): Kingdom: Protozoa: (see also Figures 2 and 3)
Kingdom: Animalia Phylum: Euglenozoa, Order: Kinetoplastidae, Family:
Order: Tylenchida Trypanosomatidae, Genus: Phytomonas
Genus: Anguina, seed-gall nematode; Ditylenchus, bulb Plant pathogenic biflagellate protozoa are transmitted
and stem n. from tree to tree by grafting and by insect vectors of the
Pratylenchus, lesion n. Radopholus, burrowing n. families Pentatomidae, Lygaeidae, and Coreidae.
Globodera, round cyst n. Heterodera, cyst.
Meloidogyne, root knot n.
Aphelenchoides, foliar n. Bursaphelenchus, pine wilt n.
Interactions of Pathogens, Plants,
Order: Dorylaemida
and Humans
Genus: Longidorus, needle n. Xiphinema, dagger n.
Paratrichodorus, Trichodorus; stubby root nematodes.
Pathogens interact with plants and humans in a variety of
ways. The main ways by which pathogens manage to
survive, infect plants, and through the damage they cause
Biflagellate Protozoa
to also affect humans, are also discussed briefly below.
To date, only a few diseases have been shown to be
caused by plant pathogenic biflagellate protozoa. All of
Ecology, Dissemination, and Epidemiology
these diseases have been found to occur in the tropics of
of Plant Pathogens
Central America. They include phloem necrosis of coffee,
heart rot of coconut palm, sudden wilt or Marchitez Some plant pathogens (viruses, viroids, mollicutes, proto-
sorpresiva of oil palm, wilt and decay of red ginger, and zoa, parasitic higher plants, most nematodes, some obligate
empty root of cassava. All these diseases seem to be biotrophic oomycetes, the powdery mildew and the rust
caused by related protozoa of the genus Phytomonas. The fungi, and the phloem- and xylem-inhabiting bacteria)
638 Pathogenesis | Plant Pathogens and Disease: General Introduction

(a) (c)

(d)

(b)

Figure 25 (a) Damage of root of tobacco plant by the lesion nematode Pratylenchus. (b) The disease cycle of the lesion nematode.
(c) Galls or root knots on roots of tomato plant infected with the root knot nematode (Meloidogyne sp.). (d) Sugar beet field in which
a large area of sugar beet plants have been severely damaged or killed by the sugar beet cyst nematode, Heterodera schachtii. Photos:
(a) and (c) courtesy of DW Dickson; (d), RJ Howard. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA:
Elsevier/Academic Press, (a) p. 850, (b) p. 852, (c) p. 838, (d) p. 847.

spend all of their lives on their host plants. Most plant introduction into their hosts. Fungi produce different
pathogenic fungi and bacteria live most of their lives in or kinds of spores that can be carried by mild or strong
in contact with their host plants, and only during periods winds to nearby or distant places. Most plant pathogenic
of unfavorable weather they live in soil or in plant debris. bacteria do not produce spores and are carried as such by
Pathogens that live all of their lives in their hosts depend splashing rain, and so on. Almost all pathogens are carried
entirely on specific animal vectors, primarily certain in infected seed, transplants, tubers, and so on. When
insects, for movement from plant to plant, and for pathogens reach or exist in their hosts, they usually grow
 Pathogenesis | Plant Pathogens and Disease: General Introduction 639

(a) Plant pathogens cause disease in plants when a virulent

pathogen lands on a susceptible host plant, and the envir-
onmental conditions, primarily temperature and
moisture, are favorable for the pathogen to be activated,
germinate if it has to, and grow. Generally, high moisture
favors most pathogens, especially bacteria and fungi, the
latter because germination of their spores depends on the
availability of moisture. Favorable temperatures and high
moisture speed up infection, sporulation, spore release
and new, secondary infections and, therefore, the number
of disease cycles and severity of the disease during the
season.
(b)

How Pathogens Infect Plants

Pathogens have evolved so that there is wide variability as
to which kind of plant it can infect and which it cannot. Of
the many kinds of plants, for example, corn, tomatoes,
apple trees, each of them may be attacked by a hundred or
more kinds of pathogens. For example, tomatoes may be
attacked by 60 fungal pathogens, 15 viruses, 10 bacteria
and mollicutes, and 15 nematodes, whereas corn may be
attacked by 130 different pathogens, including 85 fungi,
8 bacteria and mollicutes, 20 viruses, and 17 nematodes.
Figure 26 (a) Typical, although long, plant nematode, Perhaps none of the pathogens of corn are the same as the
Bursaphelenchus xylophilus, causing wilt of pine trees. (b) Dead pathogens of tomato, or 2–3 pathogens, for example, some
pine trees in a forest in Japan infected with the pine wilt
nematodes, may infect both plants. Hence, each pathogen
nematode. Photos: (a) courtesy of RP Esser; (b), JJ Witcosky.
Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. is limited as to which plants can provide it with food.
Burlington, MA: Elsevier/Academic Press, (a) and (b) p. 871). However, even if the pathogen (inoculum) has reached
the plant (inoculation), there are still a number of steps
that the pathogen must go through before the plant allows
it to feed and to multiply.

Infection by fungi and bacteria

The mechanisms of infection of plants by pathogens and
of defense of plants to infection are much better under-
stood in diseases caused by fungi and some bacteria than
in any of the other pathogens.
Once a pathogen comes in contact with a susceptible
plant, and provided that there is enough moisture and the
temperature is favorable, the sequence of the many active
phases of the plant disease cycles begins. The pathogen
must be able to adhere to the host, which the pathogen
does by excreting sticky substances. If the pathogen is a
fungal spore, or a bacterium, the latter and the spore must
Figure 27 Phytomonas protozoa in a phloem sieve tube of root get ready to penetrate the host surface. Bacteria, and
of oil palm tree affected with sudden wilt disease. W, plant cell
wall. Photo courtesy of W deSousa. Reproduced from Agrios GN
many fungi, enter plant tissues through wounds or
(2005) Plant Pathology, 5th edn., p. 876. Burlington, MA: through natural openings. Many of the fungi, however,
Elsevier/Academic Press. can also enter the host plant through direct penetration of
the plant epidermis. For these fungi to penetrate, the
inside host cells, but many fungi grow between parench- spore must germinate, so an entire new machinery must
yma cells. The spores of fungi, nematodes, and some go to work to mobilize the inert spore into an active
bacteria may come out of their hosts and be carried by inoculum. The germinating spore produces a short germ
air, water, or other means to other plans, whereas those tube on the surface of the plant and at the tip of the germ
living inside plants must be carried by their vectors. tube the fungus forms an appressorium, that is, a
640 Pathogenesis | Plant Pathogens and Disease: General Introduction

specialized feeding organ present in some organisms. The from infected plants and introduce it into healthy plants
appressorium is oppressed and sticks to the plant surface through the wounds they make while feeding, or they are
on which it produces a penetration peg that exerts tre- brought into the host plant by pollen. Once a virus (or
mendous pressure on the epidermal cell it tries to viroid) enters a plant cell, the virus sheds its protein coat
penetrate. Finally, a thin infection hypha grows through and the naked nucleic acid (RNA or DNA) associates
the opening created by the penetration peg and the fine with appropriate cell organelles, primarily ribosomes,
hypha resumes its normal diameter and absorbs nutrients and also membranes, enzymes, and so on, which replicate
from the plant. the nucleic acid into hundreds of thousands or millions of
Up to that moment of penetration by the infection copies. Soon, translation of part of the nucleic acid strand
hypha, the fungus spore provided genes for producing results in the production of molecules that are the sub-
adhesive substances, enzymes, cytokinins, and so on, for units of the viral coat protein and these assemble with the
the activation and germination of the spore, for producing nucleic acid strands and form the complete virus particle
the appressorium and then the penetration peg, which (a virion). In the process of virus replication and accumu-
enabled the fungus to reach the food-laden contents of lation, the metabolism of the host plant is affected.
the plant cell. If the plant is susceptible to this race of the The defense mechanisms of the infected plant are
pathogen, the pathogen will produce enzymes for break- activated, and if the defenses are effective, that is, if the
ing down the substances that make up the plant cell plant is resistant, virus replication and movement may be
wall, such as chitinases for chitin, cutinases for cutin, stopped, the infected and a few surrounding cells may die,
cellulases, pectinases, ligninases, and so on, to break and no further infection develops beyond a tiny visible or
down cellulose, pectins, and lignin, and enzymes for invisible local lesion. If, however, the host defenses are
breaking down cell content substances, such as amylase ineffective or inadequate, the virus moves from cell to
for starch, proteases for proteins, and lipases for lipids and cell, replicating in each of them, and finally reaches the
fats. This, of course, allows the pathogen to grow and phloem sieve elements that make up part of the phloem
multiply and, eventually, to produce spores on the plant
veins. Once in the phloem, the virus is distributed
surface.
throughout the plant and from the phloem cells it moves
In the meantime, through their enzymes and other
into adjacent parenchyma cells and to the other living
toxic substances that they produce in the plant, and
cells in which it again replicates. Depending on the age
depending on the plant organs they infect, the pathogens
and size of the plant, viruses can spread throughout the
may cause smaller necrotic (dead) spots or larger patches
plant within a few days to a few weeks or months. Infected
of plant tissue to die and rot, to shrink, or to show other
plants are often stunted, their leaves, shoots, and fruit
symptoms. Many fungal and bacterial plant pathogens
become smaller, discolored, or malformed, yields are
may also produce toxins that can kill tissues directly, or
growth regulators that cause infected plant tissues to grow reduced, and with some virus–host combinations some
and produce galls rather than be destroyed and die. plant parts or whole plants may be killed.
On the other hand, if the plant is resistant to the Nematodes: When an infective nematode juvenile or
pathogen, as soon as some molecule or factor (elicitor) adult reaches the surface of a plant, it pierces the outer
of the pathogen is recognized by the plant, the defenses of epidermal cell wall with its stylet, and secretes saliva
the plant, both structural and chemical, preexisting or into the cell, the contents of which are liquefied and
induced, are mobilized and the pathogen may become absorbed through the stylet. Some nematodes are rapid
neutralized, and this leads to the death of the pathogen feeders and feed only on epidermal cells, moving their
and to no plant infection. stylet from one cell to the next without ever entering the
Bacteria infect plants in a manner similar to that plant. Others enter the plant parts and feed slower while
employed by fungi, except that bacteria enter plants inside the plant. Some of the latter become attached to
through wounds or natural openings and, therefore, do one area of the plant and do not move. Some of the
not need or produce a germ tube or haustoria, but multi- nematodes enter the plant, feed internally for varying
ply and move about the host rather than grow into them. lengths of time, and then exit the plant and move about
freely. Depending on where the female of a particulate
nematode species is feeding, she lays her eggs inside or
Infection by pathogens surviving only in living outside the plant. When the eggs hatch, the new infec-
cells tive juveniles either cause further infections inside the
Plant viruses, viroids, mollicutes, xylem- or phloem- plant or infect new plants. The mechanical injury caused
limited bacteria, protozoa, and nematodes, with just a by nematodes in the infected area, as well as the removal
few exceptions, enter plant cells either with the help of of nutrients from plants by nematodes, certainly has a
a specific vector such as a specific insect, mite, fungus, or detrimental effect on the plant. It is thought, however,
nematode, which, with their mouth parts, pick up virus that much greater damage by nematodes on plants is
 Pathogenesis | Plant Pathogens and Disease: General Introduction 641

caused by the enzymes, growth regulators, and toxic antimicrobial substances known as pathogenesis-related
compounds contained in the secretions of the nematodes proteins, and a variety of secondary metabolic com-
into the plant and by the ports of entry for other patho- pounds, such as chlorogenic acid, caffeic acid, and
gens (fungi and bacteria) created by nematodes on plant ferulic acid. Some plants, upon infection with a pathogen,
roots. produce phytoalexins, which are toxic antimicrobial sub-
stances produced in appreciable amounts in plants only
after stimulation of the plant by various types of patho-
How Plants Defend Themselves against
genic microorganisms or by chemical and mechanical
Pathogens
injury. Besides, although the plant cells produce antimi-
Because each pathogen can attack and infect only a few crobial compounds aimed against the pathogen, the
kinds of plants, that is, its own hosts, we presume that plants, in some diseases, defend themselves by detoxifying
nonhost plants are resistant, indeed immune, to these the toxins of the pathogen.
pathogens. Also it is likely that all plants normally pro- If the defense reactions are quick and effective, further
duce structures such as thick-walled epidermal cells, growth and activities by a biotrophic pathogen are limited
fewer or smaller stomates, and chemicals such as phenolic or stopped, and the infected cell collapses and dies.
compounds. These structures or chemicals, by their pre- Because the pathogens can no longer attack and obtain
sence in or around plant cells, act as the first mechanisms food from such plants, such a plant is considered resistant
of plant defense against any pathogen that must overcome to the pathogen. This type of defensive response by the
them to reach the interior of the cells for the food in it. In plant, following recognition of a pathogen elicitor by the
addition to such preexisting structural and chemical receptor of a plant cell, is known as the hypersensitive
defenses, plants seem to be induced by the presence of response (HR). Frequently, however, the term ‘HR’ is
the pathogen, to produce induced structural defenses such used to indicate that the host plant reacts to a particular
as cork layers, abscission layers, gums and tyloses, and pathogen by producing small local lesions and therefore is
biochemical defenses consisting of numerous biochemical considered as resistant. Plant defenses against pathogens
reactions taking place in infected cells and tissues. may be in the form of single or multiple, structural or
Indeed, the induced biochemical reactions play a chemical barriers, in the form of programmed plant cell
major role in plant defense against pathogens. In the death that stops further growth of biotrophic pathogens,
host–pathogen combinations that have been studied, as or in activation of defense responses regulated by the
the pathogen comes in contact with a host plant, some of salicylic acid-dependent pathway, or, more commonly,
the molecules secreted by the pathogen, or released from in a combination of various mechanisms of defense.
host plant tissues by enzymes of the pathogen, react with Necrotrophic pathogens, which actually benefit from
receptor molecules present in host cells. The plant recep- host cell death, are not limited by cell death and the
tor molecules recognize such existing or induced plant associated salicylic acid-dependent defenses but rather
molecules (elicitors) as harbingers of a pathogen attack by a different set of responses activated by jasmonic acid
and quickly trigger a cascade of defense reactions in the and ethylene signaling. If no defense reactions occur, or if
attacked and the surrounding cells. they are ineffective or too slow, the plant becomes
One of the first products of defense reactions in the infected, and such a plant is considered susceptible to
infected cell is the rapid and transient generation of the pathogen. Generally, resistance in commercial crop
activated oxygen species, which, in turn, triggers the plants can be, and usually is, the result of one or more
hyperoxidation of plant cell membranes, which produce resistance genes (R gene). In almost all wild plants, and in
molecules toxic to the plant cell and to the pathogen. most cultivated ones, however, resistance in a plant to
Such cells become dysfunctional and lead to cell collapse infection by one or more pathogens is the result of the
and death. The activated oxygen species are also involved expression of many plant genes controlling various defen-
in host defense reactions by activating the phenoloxidase sive structural, chemical, and physiological characteristics
enzymes that oxidize phenolic compounds into more of the plant.
toxic quinones. In addition, oxidation of membrane lipids
produces several biologically active molecules, such as
Role of Genetics in Disease Initiation
jasmonic acid, which is the precursor of the wound hor-
and Development
mone traumatin, and, along with salicylic acid, appears to
induce numerous protein changes and acts as a signal When a pathogen attacks a plant, it may not be able to
transducer of the defense reaction in plant–pathogen infect it because the plant belongs to a taxonomic group
interactions. The affected plant cells continue to defend outside the range of the pathogen and has genes for
themselves by producing cell wall strengthening materials resistance for which the pathogen has no corresponding
such as callose, glycoproteins such as extrensin, phenolic genes for virulence. This is called nonhost resistance. In
compounds, and so on. They further produce a variety of another kind of resistance, the host may actually be
642 Pathogenesis | Plant Pathogens and Disease: General Introduction

susceptible to the pathogen but because it possesses genes old gene for resistance (A) and has all the plants with that
that allow it to grow earlier or later than the time that the gene to itself, without any competition from other patho-
pathogen is present (disease escape), or allow it to grow gens. As a result, the new variant multiplies unimpeded
and produce a crop in spite of it being infected with the and produces a new race of the pathogen, race (B) that can
pathogen (tolerance), this type of resistance is called infect all plants of variety (A). The plant breeders, there-
apparent resistance. In most cases, however, plants are fore, must produce a new plant that will contain a gene for
resistant to a pathogen because they possess genes for resistance (B) that will stop the new pathogen. The easiest
resistance (R genes) directed against avirulence genes of and best way to do that is by collecting individuals of the
the pathogen (true resistance or race-specific, cultivar- pathogen race (B) and inoculating as many new varieties
specific, or gene-for-gene resistance). or germ lines they have and hoping to find one or more
Plant pathogens have genes for pathogenicity, which plants that do not become infected with this race. This
make them pathogens, and genes for virulence or aviru- new variety or individual is propagated to produce the
lence, corresponding to genes for susceptibility or new variety (B) of the crop that will replace variety (A) in
resistance in the host. This statement is probably the field.
the most straightforward statement about the role of
genetics in disease and calls to mind that, for each gene
Effect of Pathogens on Plants, Crops,
for resistance in a plant, there is a gene for virulence in the
and Humans
pathogen and vice versa. Pathogens are pathogenic on
some varieties of the host plant but not pathogenic on Pathogens affect plants in some general ways regardless of
other varieties of the same host. Plants have genes some specific differences depending on the kind of plant,
for resistance, which produce the innumerable structural the kind of pathogen, and the prevailing weather condi-
and biochemical substances that help them defend them- tions during the initiation and development of disease.
selves, whereas pathogens have virulence genes that The first and main effect of pathogens on plants they
enable them to infect susceptible plants. Pathogens also infect is that they change the appearance of plants and
have avirulence genes thought to code for molecules that, produce the symptoms characteristic for each disease.
when recognized by specific receptor molecules, pro- Symptoms are almost always accompanied by reduced
duced by resistance genes in the plant, act as elicitors of growth, productivity, and quality of the plants. It is the
the host resistance response. Recognition of the aviru- amount of yield that goes to the heart of the reasons plants
lence gene product (elicitor) of the pathogen by the are cultivated by humans. The extent of reduction is
plant cell-produced defense response gene product important not only because it reduces the expected
(receptor molecule) sets in motion the production of a income or profit but also because yield reduction as a
variety of structural compounds and biochemical reac- result of disease results in hunger and starvation of
tions that lead to cell disruption and eventually to the humans and animals affected by the loss. Yield losses
death of the attacked plant cell and of the attacking may be insignificant (2–3%), low (10–30%), large
pathogen. The death of the two combatants (attacking (40–60%), or complete (100%). Such losses are generally
pathogen and cell of attacked plant) stops further devel- avoided and absent in areas and countries with advanced
opment of infection and disease. In addition to the education and economies and where humans have the
virulence/avirulence genes, several other types of genes knowledge and the resources to buy foodstuffs from coun-
seem to play important roles in the development and tries in which the disease was not as destructive that year.
expression of disease. In advanced countries, severe disease losses may be of
When a resistant variety (A) is widely distributed for little concern to a few people, who may have to go to the
cultivation, within 2–4 years the variety loses its resis- bank and borrow money to buy from other places quan-
tance to pathogen (a) and becomes susceptible. That tities of the crop needed for consumption and for seed for
variety, therefore, must be replaced with a new variety the next year. In poor areas and poor, underdeveloped
that contains a new gene for resistance (B) against the new countries, however, severe losses of a staple crop may
gene for virulence (A) of the pathogen. Actually, the affect the survival of humans, animals, and industries
variety (A) did not lose any resistance. Instead, the patho- that may exist and flourish in a large area or country. In
gen, indeed all races of it, because they were excluded poor countries, people have no knowledge of the cause of
from the resistant new crop (A), was placed under a plant disease and how to control it, no access to resistant
extreme survival pressure and this led to the selection or varieties or to fungicides, if available, or collateral to
appearance by mutation of a new variant of the pathogen borrow money from a bank and to find quantities of the
individual(s) that carried the gene for virulence that crop in other countries. Also, because of lack of knowl-
enabled the pathogen to overcome or bypass the product edge, a catastrophic epidemic of a plant disease occurring
of the gene for resistance (A). This new individual, being 1 year is likely to be repeated in subsequent years because
the only one with virulence gene (B), is unaffected by the of the large amount of pathogen inoculum that survives in
 Pathogenesis | Plant Pathogens and Disease: General Introduction 643

the field and that can cause disease again the following before or after they are placed in storage. Such infections
year. Therefore, undernourishment, extreme suffering produce in the seeds a variety of toxic substances, called
from hunger, and devastating famines caused by plant mycotoxins, although they do not produce any character-
diseases are commonplace in poor, little-developed coun- istic structures as do plants infected with the ergot fungus.
tries, and some of them, such as the Irish famine of 1845 Peanuts and corn seem to be the seeds affected most
and 1846, caused by the destruction of the potato crop by frequently and most severely by these fungi, especially
the late blight oomycete P. infestans, and the Bengal fam- if they are damaged and when the weather is very humid.
ine caused by the destruction of rice by the leaf spot Mycotoxins are extremely toxic and cause severe hemor-
fungus Cochliobolus miyabeanus. rhage, serious diseases of the nervous and circulatory
When plants are attacked by pathogens frequently and systems, damage to one or several of the internal organs,
repeatedly until they can no longer survive the attacks of vomiting, rejection of food, and others. The most common
these pathogens, there may be effects on whole ecosys- fungi that produce mycotoxins in infected plant seeds and
tems. For example, repeated infections of wheat and of products are Aspergillus, which produces aflatoxins,
forage grasses by the rust, smut, or ergot fungi reduce the ochratoxins, and so on; Fusarium, which produces
amounts of food not only to humans but also for wild trichothecins (vomitoxin), zearalenone, and fumonicins;
animals dependent on such plants for their survival. Penicillium, which produces patulin, roquefortin, and so
Although such repeated attacks occur on plants of all on, and several others.
shapes and sizes, we are most aware of those occurring A few fungal pathogens of grasses grow inside the
on trees because they are easier to see and to follow the plants (endophytes), and although these pathogens do
happenings to them by a particular pathogen. The best not seem to cause apparent diseases on their hosts, they
known of these are the destruction and near extinction of produce toxic compounds that cause severe diseases in
the American chestnut by the chestnut blight disease, the wild and domestic animals that eat the plants.
caused by the fungus Cryphonectria parasitica, in the All plant diseases cause some financial losses because
Northeastern United States and Canada; the destruction of reduction in the quantity and quality of the plants and
and near extinction of the American elm by the Dutch plant products they infect. Such losses become increas-
elm disease caused by the fungus Ophiostoma nova-ulmi; the ingly more important as the amounts of losses increases
near extinction of the coconut palm tree in Florida and and as the size of the area affected by a disease increases.
the Caribbean basin by a phytoplasma mollicute that still The amount or percentage of losses is, of course, particu-
has not been named or classified, and so on. In many cases, larly important to the growers whose crop was affected by
the survival of some plants is impossible in areas that have the disease – and some growers, who were fortunate for
been invaded by a pathogen, which then becomes a per- their crop to escape the disease, may actually profit by the
manent inhabitant of the area, as happened in Central higher price they get as a result of reduced supplies and
American banana fields showing infection with the increased prices for the crop. In spite of the few who may
Panama disease, caused by the fungus Fusarium lycopersici profit from a plant disease, the vast majority of times,
if. [Link] that became unfit for the production of bana- everybody, growers and consumers alike, lose from it.
nas after the field became contaminated and thoroughly The reasons for this is that the costs for growing such a
infested with the pathogen. This effect reduces the crop will necessarily increase for every grower in subse-
amount of land available for each crop that is susceptible quent years because they may have to replace the
to that pathogen and forces growers to plant crops other susceptible variety with a resistant one that may not be
than the one that is more productive and profitable. as productive or as desirable, may not be as profitable, and
Plants infected with certain pathogens, for example, may be susceptible to another pathogen present in the
rye or wheat, infected with the ergot fungus, Claviceps area; besides, it will take time for all growers to acquire
purpurea, become unfit for human or animal consumption seed of the new variety. Also, the grower may need to
because the fungus produces fruiting structures, called change methods and equipment for cultivation, harvest-
ergots, that replace several of the seeds in each head. ing, and storage and transportation of the new variety or
Ergots (Figure 7(a)) contain a large number of toxic new crop, may need to build refrigeration, and so on.
alkaloids and other compounds, and humans or animals Finally, the grower may need to use pesticides to manage
eating even a small amount of ergots become severely or control the disease, and this will add more costs for
diseased because the contents of the ergots damage inter- purchase, storage, and application of pesticides, plus the
nal organs, such as the kidneys, the nervous system, and environmental costs of applying pesticides to the crop.
the circulatory systems, and result in severe painful dis- Besides, for many diseases, such as those caused by
eases characterized by gangrene of the extremities of viruses, phloem- and xylem-limited fungi, bacteria, mol-
affected individuals. licutes, and so on, no pesticides are available, and
Certain fungi infect mature, harvested seeds of grains therefore this method of control with pesticides often is
and legumes and also bread, hay, or plant products such as not available.
644 Pathogenesis | Plant Pathogens and Disease: General Introduction

Detection and Identification of Plant Pathogens Management and Control of Plant Diseases
Although some pathogens, such as the parasitic higher Some of the most common and most effective measures
plants, are big and can be identified by their structure, for management or control of plant diseases, regardless of
most pathogens are small and must be looked under the the kind of pathogen, include the following:
microscope (nematodes, fungi, bacteria), for example, or
even underthe electron microscope (viruses, bacteria, 1. Exclusion of the pathogen from a field by planting
protozoa), to even begin getting close to a correct identi- pathogen-free seed, seedlings, tubers, and so on.
fication. Generally speaking, pathogens are difficult to 2. Eradication of infected parts or entire plants, and of
identify. possible alternate hosts of the pathogen.
For detection and identification of plant pathogens it is 3. Improving cultural practices so that they support better
fastest when one reexamines the symptoms, amount growth of the host while impeding growth of the patho-
Attempt to culture pathogens. Different and modern tech- gen. These include appropriate fertilization, watering,
niques are used. Although microscope use is routine, and pruning, thinning, temperature manipulation, and so on.
access to electron microscope is increasing, the use of 4. Planting resistant varieties whenever possible and
modern serological tests and DNA/RNA can provide a available. One should always plant varieties resistant
faster, more accurate, and more dependable result. to the particular pathogen, and preferably to more than
Most fungi and nematodes can be identified by examina- one pathogen. As many varieties are resistant to only
tion of whole nematodes under the microscope and fungi by one gene for resistance, one should expect that new
examination of the kinds of their spores and fruiting bodies. races of the pathogen will soon bypass or overcome the
Viruses and viroids are identified by their symptoms resistance gene and therefore must continually pro-
and by inoculating other plants and comparing them with duce varieties resistant to each of the pathogen races
the symptoms shown by other inoculated plants; checking that carry different genes for resistance. Breeding vari-
epidermal cells for characteristic cellular inclusions; eties resistant to disease has received a tremendous
exposing viruses to different virus vectors; and, wherever impetus by employing the tools and methodology of
available, most certainly serological tests and tests invol- genetic engineering and biotechnology.
ving DNA and RNA. 5. Management and control with chemicals. Plant patho-
The value of quick identification of a pathogen that genic fungi and bacteria are sensitive to numerous
may cause a potentially serious disease has increased chemicals (fungi to fungicides, bacteria to bacteri-
greatly in recent years both because it has been proven cides). Fungicides (and bactericides), applied on the
repeatedly that plant diseases, like human and animal surface of plant foliage, blossoms, stems, and fruits,
diseases, cause disproportionately more damage to crops protect, to a certain extent, these organs from infection.
when detection and diagnosis are late and slow and Some fungicides are absorbed by the plant roots, and
because of the political realities and threats to the secur- are distributed systemically through the plant, stop-
ity of our food supply. Extreme prolonged droughts, ping infections wherever they may be initiated. There
attack of plants by diseases, insects, and so on; political are no fungicides or other chemicals effective against
interference over large agricultural areas in the past few viruses, viroids, or any of the other pathogens limited
years have already resulted in frequent hunger and to the xylem or phloem of the plant.
several famines affecting the poor people inhabiting 6. Biological control. Some plant pathogenic fungi and
these areas. bacteria are inhibited in their growth and ability to
The need for rapid and correct diagnoses of the causes cause disease when exposed to certain antagonistic or
of plant diseases, insects, and so on; has lead the parasitic microorganisms present in the vicinity
University of Florida, at Gainesville, FL, to create a new (Figure 28). This is called biological control.
program of graduate study called the Doctor of Plant Unfortunately, for only about 1% of the diseases
Medicine (DPM) Program, The DPM program leads to have materials and methods been developed, but
a professional Doctorate degree. The main purpose of the even with these, control is not satisfactory.
DPM is to train students at the graduate/doctoral level,
on how to diagnose and how to control any kind of In addition, inoculating plants with mild strains of a virus
disease, injury or damage caused by any biotic (insects, protects them from infection by severe strains of the same
mites, weeds, plant pathogens, birds, etc.) or abiotic fac- virus. In the past several years, successful control of plant
tors (such as nutritional deficiencies, cold temperatures, viruses has been achieved by genetically engineering
air and soil pollutants). The 4-year program accepted its plants to carry and express the coat protein and other
first 15 students in 2000 and by 2007; it had graduated genes of the virus, which makes the plants resistant to
nearly 40 Doctorates in Plant Medicine. subsequent infection by that virus.
 Pathogenesis | Plant Pathogens and Disease: General Introduction 645

(a) (b)

Figure 28 Biological control of certain plant pathogenic oomycetes and fungi with (a) other oomycetes or fungi that penetrate and
feed on the first. (b) A plant pathogenic fungus (Botrytis sp.) is attacked by a yeast fungus (not pathogenic on plants). Photos: (a)
courtesy of R Baker; (b), P Wisniwski. Reproduced from Agrios GN (2005) Plant Pathology, 5th edn. Burlington, MA: Elsevier/Academic
Press, (a) and (b) p. 306.

Nematodes also benefit from the above, but when See also: Fungi: Plant Pathogenic; Mycoplasma and
nematodes are present in a field, they can be managed Spiroplasma; Plant Pathogens and Disease: Newly
primarily by planting resistant plant varieties and by Emerging Diseases; Plant Pathogens Minor
applying pre- or postplant nematicides. Several cultural (Phytoplasmas); Plant Disease Resistance: Breeding and
practices, such as plowing the soil before planting, expose Transgenic Approaches; Plant Disease Resistance:
the soil to the sun and dries it up and causes reduction of Natural, Non-Host Innate or Inducible; Plant Pathogens,
the numbers of nematodes present. Bacterial; Virus Infection

Conclusion Further Reading

Agrios GN (2005) Plant Pathology, 5th edn. San Diego: Academic
Plant pathogens, with a couple of minor exceptions, are Press/Elsevier.
microorganisms that belong to the same taxonomic Alexopoulos CJ, Mims CW, and Blackwell M (1996) Introductory
Mycology, 4th edn. New York: Wiley.
groups, that is, bacteria, viruses, fungi, protozoa, and American Phytopathological Society Numerous Volumes on Plant
nematodes, which include the pathogens that cause dis- Diseases and Their Control, in a Series of ‘Plant Disease Compendia’
ease in humans and animals. Each species of plants for Specific Plants. St. Paul, MN: APS.
Bos L (1999) Plant Viruses, Unique and Intriguing Pathogens. Leiden,
appears to be attacked by about 100 kinds of pathogens. The Netherlands: Backhuys Publishers.
Plant pathogenic fungi and bacteria live most of the time Brandl MT (2006) Fitness of human enteric pathogens on plants and
within their plant hosts and the rest of the time in the implications for food safety. Annual Review of Phytopathology
44: 367–392.
soil. The other pathogens live only in their plant hosts. Burdon JF, Thrall PH, and Lars Ericson (2006) The current and future
Plant pathogens cause disease in plants and cause losses dynamics of disease in plant communities. Annual Review of
Phytopathology 44: 19–39.
in food and other necessary items. The losses may be DeBoer SH (ed.) (2001) Plant Pathogenic Bacteria. Dordrecht, The
light or very severe, sometimes destroying all the plants Netherlands: Kluwer Academic Publisher.
and causing hunger, starvation, and famines, whereas in Dollet M (1984) Plant diseases caused by flagellated protozoa
(Phytomonas). Annual Review of Phytopathology 22: 115–132.
other cases they result in extinction of entire species of Fuchs M and Gonsales D (2007) Safety of virus-resistant plants
plants. two decades after their introduction: Lessons from realistic field
646 Pathogenesis | Plant Pathogens and Disease: General Introduction

risk assessment studies. Annual Review of Phytopathology Schumann GL and D’Arcy CJ (2006) Essential Plant Pathology. St. Paul,
45: 173–202. MN: APS Press.
Hull R (2002) Matthews’ Plant Virology, 4th edn. New York: Academic Sinclair WA and Lyon HH (2005) Diseases of Trees and Shrubs, 2nd
Press. edn. Ithaca, NY: Cornell University Press.
Maloy OC (1993) Plant Disease Control: Principles and Practice. Spetafora J (2007) Pezizomycotina. The Tree of Life Web Project.
New York: Wiley and Sons. Strange RN and Scott P (2005) Plant disease: A threat to global
Nickle WE (ed.) (1991) Manual of Agricultural Nematology. New York: food security. Annual Review of Phytopathology 43: 83–116.
Dekker. Van Loon LC, Rep M, and Pieter CMJ (2006) Significance of inducible
Press MC and Graves JD (1995) Parasitic Plants. London: Chapman defense-related proteins in infected plants. Annual Review of
and Hall. Phytopathology 44: 135–162.