PLANT PATHOLOGY HANDOUT

1. INTRODUCTION
History of Plant Diseases
Plant diseases had profound effects on mankind through the centuries as evidenced by Biblical
references to the blasting and mildew of plants. The Greek philosopher Theophrastus (370-286
B.C.) was the first to describe maladies of trees, cereals, and legumes that we currently classify
as leaf scorch, rots, scab, and cereal rust. The Romans were also aware of rust diseases of their
grain crops.
With the invention of the microscope in the 17th century, fungi and bacteria associated with
plants were investigated. In 1665, Robert Hooke published the first illustration of rust on a rose
leaf. Advances in the study of diseases were hampered by the widely held belief in the theory of
spontaneous generation. This theory, held by most people in the mid-18th century, considered
pathogenic or disease causing microorganisms as products of disease rather than causes of
disease. Epidemics of late blight of potato devastated Ireland in 1845 and 1846. These epidemics
dramatized the effect of plant diseases on mankind. Tragically, these epidemics caused famine
and death for over a million people. Between 1845 and 1860, death and migration accounted for
the loss of nearly one-third of Ireland’s population. In 1861, a German botanist, Anton De Bary,
proved that a fungus (Phytophthora infestans) was the causal agent of late blight of potato. This
was a milestone in the study of plant diseases since it showed that a fungus was indeed the cause
of a plant disease rather than an organism simply associated with the disease. Two years later,
Louis Pasteur proposed his germ theory of disease that finally disproved the theory of
spontaneous generation and changed the way modern science investigated the diseases of all
living organisms.

Significance of Plant Diseases in the United States

A few examples of plant disease epidemics that have resulted in devastating plant losses in the
United States include: chestnut blight, introduced in 1904, virtually eliminated chestnut trees

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from North America; citrus canker, introduced in 1910, and a closely related bacterium called
citrus bacterial spot discovered in 1984, resulted in the destruction of millions of citrus trees;
white pine blister rust, introduced in 1912, caused large economic losses in the timber industry;
and Dutch elm disease, introduced in 1930, continues to destroy large numbers of elm trees from
the East Coast to the Pacific Northwest

Plant Diseases:
• Prevent the cultivation and growth of food plants in some areas;
• Destroy parts or all of the plants, and
• Reduce much of the crop produce/food, before they can be harvested.

Plants and Disease

Plants make up the majority of the earth’s living environment. They make up all the food on
which humans and all animals depend. Plants are the only higher organisms that can convert the
energy of sunlight into stored, usable chemical energy (Carbohydrates, Proteins and fats). All
animals, including humans, depend on these plant substances for survival. Plants, however, also
get sick. Sick plants grow & produce poorly; they exhibit various types of symptoms. Often,
parts of plants or whole plants die. It is not known whether diseased plants feel pain or
discomfort. The agents that cause disease in plants are the same to those causing disease in
humans and animals.

Types of Plant Diseases

• Several thousands of diseases affect cultivated and wild plants.
• Each kind of crop plant can be affected by a hundred/more plant diseases.
• Some pathogens affect only one variety of a plant.
• Other pathogens affect several hundreds of species of plants.

Plant pathology (phytopathology) is the scientific study of plant diseases caused by pathogens
(infectious diseases) and environmental conditions (physiological factors). Organisms that cause

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infectious disease include fungi, oomycetes, bacteria, viruses, viroids, virus-like organisms,
phytoplasmas, protozoa, nematodes and parasitic higher plants.
The term pathology comes from two Greek words (Pathos - refers to suffering; logos - the
study/to speak). Plant pathology is therefore;
• The “study of the suffering plants.”
• The study of microorganisms and environmental factors that cause disease in plants, the
mechanisms by which disease occurs, the interaction between these causal agents and the
plant, and the methods of managing or controlling plant disease.

Disease in Plants:
Definitions
Disease: any malfunctioning of host cells and tissues that result from continuous irritation by a
pathogenic agent or environmental factor that leads to development of symptoms.
Pathogen: any entry that can incite disease; any living or non-living entity that brings about or
incites a disease by its persistent association with the host or plant.
Parasite: an organism living on or in another living organism (host) and obtaining its food from
the latter.
Host: a plant that is invaded by a parasite & from which the parasite obtains its nutrients.
Pathogenicity: the capability of a pathogen to cause disease.
Virulence: the degree of the pathogenicity of a given pathogen.
Symptom: the external or internal reactions of a plant as a result of a disease.
Sign - the pathogen or its parts or products seen on a host plant.

1.2. Plant Pathology and Its Role

Plant pathology (Phytopathology) is a science that has a practical and noble goal of protecting
the available food for humans and animals. It is an integrative science and profession that uses
and combines the basic knowledge from other scientific fields such as botany,
nematology, mycology, bacteriology, virology, genetics, biochemistry, horticulture, agronomy,
physics, chemistry, forestry, molecular biology and genetic engineering, plant physiology, plant
anatomy, meteorology and many other branches of science.
As a science, plant pathology tries to:
• Increase our knowledge about plant diseases

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 • Develop methods, equipments, and materials through which plant diseases can be
avoided or controlled.
• Also involve the study of pathogen identification, disease cycles, economic impact, plant
disease epidemiology, plant disease resistance, pathosystem genetics, and management of
plant diseases.
Uncontrolled plant diseases may result in less food and higher food prices or in food of poor
quality. Some plant diseases may wipe out entire plant species and many affect the beauty and
landscape of our environment. Controlling plant disease results in more food of better quality
and a more aesthetically pleasing environment. It is estimated that diseases, insects & weeds
together annually interfere with the production or destroy between 31 & 42% of all crops
produced world wide. The losses are usually lower in the more developed countries & higher in
the developing countries. It has been estimated that of the 36.5% average of total losses (14.1%
by diseases, 10.2% insects and 12.2% by weeds). The total annual worldwide crop loss from
plant diseases is about $ 220 billion.

2. CLASSIFICATION OF PLANT DISEASES

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2.1. Criteria for Disease Classification:

2.1.1. Host plant (crop category/taxonomic approach):

• Cereal, legume, forest, etc. diseases
2.1.2. Based on organ of the crop affected:
• Root, stem, leave, floral, fruit and seed diseases
2.1.3. Based on physiology of diseased plants (physiological approach):
• Storage of food: cabbage soft rot, seed decays, etc.
• Hydrolysis and utilization of stored food: damping off, seedling blight, etc.
• Absorption and accumulation of water and minerals: diebacks, etc.
• Growth and reproduction: galls and smuts (instead of normal seed)
• Conduction of water and transpiration: vascular wilts
• Photosynthesis: leaf diseases (spots, mildews, blights, rusts and anthracnose affect the
photosynthetic areas.
• Translocation of elaborated food materials: Tumors
2.1.4. Based on symptoms (symptomological approach):
• Major focus on the host plant (rots, wilts, blights, etc.)
2.1.5. Based on the occurrence of the disease (epidemiological approach):
Endemic disease: regularly present from season to season
Epidemic/epiphytotic disease: a wide spread & destructive outbreak of a disease of plants
Sporadic disease: an occasional occurrence of infectious disease
Pandemic disease: epidemic occurrence in multiple countries that results in shortage of food
and death. e.g., late blight of potato.
Plant diseases were first studied because of the yield losses they cause.

2.1.6. Based on source of Inoculum spread:

• Air-borne, seed-borne, soil-borne, animals & man dispersed

2.1.7. Taxonomy of the pathogen (living and environmental factor prone to disease):

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On this base diseases are classified as:
• Infectious or transmissible plant diseases
Diseases caused by fungi, bacteria, nematodes, viruses and viroids and parasitic higher
protozoa.
• Noninfectious or physiological disorders
Too low or too high temperature, lack or excess of soil moisture, light, lack of oxygen, air
pollution, nutrient deficiencies, mineral toxicity, soil acidity or alkalinity (pH), toxicity of
pesticides and improper agricultural practices.
Non-pathogenic/non-infectious/non-transmissible diseases:-
The most useful criterion for classification of a disease is the type of pathogen that causes the
disease.

2.2. Taxonomy/Classification of Plant Pathogenic Fungi

The fungi that cause diseases on plants are diverse group. Some fungi, often referred to as the
lower fungi, are now considered to belong to the kingdom Protozoa (e.g. the mycomycotes and
plasmodiophoromycetes) or the kingdom chromista (e.g. the Oomycetes). The true fungi,
however, that is chytridiomycetes, zygomycetes, ascomycotes, basidiomycetes and
deuteromycetes belong to the kindom Fungi (earlier called Eumycotina or Mycetae).
Fungilike Organisms or Pseudofungi
KINGDOM PROTOZOA:
Microorganisms that may be unicellular, plasmodial, colonial, very simple multi-cells, or
phagotrophic, i.e. feeding by engulfing their food. The kingdom contains many microorganisms
in addition to fungi-like organisms Myxomycetes and Plasmodiophoromycetes.
Phylum: MYXOMYCOTA - Produce a plasmodium like structure.
CLASS: MYXOMYCETES (slime moulds) – Their body is naked, amorphous plasmodium.
They produce zoospores (swarm cells). May grow on and over parts of low-lying plants but do
not infect plants.
Order: Physarales - Saprophytic plasmodium that gives rise to crusty fructifications containing
spores. They produce zoospores that have flagella.
Genus: Fuligo, Mucilago, and Physarum cause slime molds on low-lying plants.

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Phylum: PLASMODIOPHOROMYCOTINA (theplasmodiophoromycetes endoparasitic slime
moulds).
Order: Plasmodiophorales – Plasmodia produced within cells of roots and stems of plants. They
produce zoospores that have two flagella. Obligate parasites.
Genus: Plasmodiophora, P. brassicae causing club-root of crucifer Polymyxa, P. graminis
parasitic in wheat and other cereals. Can transmit plant viruses. Spongospora, S. subterranean
causing powdery scab of potato tubers.

KINGDOM: CHROMISTA
Unicellular or multicellular, filamentous or colonial, primarily phototrophic (micro-organisms),
some with tubular flagellar appendages or with chloroplasts inside the rough endoplasmic
reticulum, or both. Contains the brown algae, diatoms, oomycetes, and some other similar
organisms.
Phylum: OOMYCOTA – Have biflagellate zoospores, with longer tinsel flagellum directed
forward and a shorter whiplash flagellum directed backward. Diploid thallus, with meiosis
occurring in the developing gametangia. Gamentagial contact produces thick-walled
sexual oospores. Cell walls composed of glucans and small amounts of hydroxyproline and
cellulose.
CLASS: Oomycetes (the water moulds, white rusts, and downy mildews) – Have non-
septate elongate mycelium. Produce zoospores in zoosporangia. Zoospores have two
flagella. Sexual resting spores (oospores) produced by the union of morphologically
different gametangia called antheridia (male) and oogonia (female).
Order: Saprolegniales – Have well-developed mycelium. Zoospores produced in long,
cylindrical zoosporangia attached to mycelium. Usually several oospores in an
oogonium.

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Genus: Aphanomyces, A. euteiches causing root rot of peas.
Order: Peronosporales – Mycelium well-developesd, non-septate, branching, inter- or
intracellular, often with haustoria. Zoosporangia oval or lemon-shaped, borne on
ordinary mycelium or on sporangiophores. Sporangia in most species germinate by
producing zoospores, but in some they germinate directly and produce a germ tube.
Sexual reproduction is by characteristic oogonia and antheridia that fuse and produce
an oospore.
Oospores germinate by giving rise to zoospores or to a germ tube which soon
produce a sporangium, depending on the species
Family: pythiaceae - sporangia, usually zoosporangia, produced along somatic hyphae or
at tips of hyphae of indeterminate growth and set free. Oogonia thin-walled.
Facultative parasite.
Genus: pythium, causing damping-off of seedlings, seed decay, root rots, and cottony
blight of turf grasses.
Phytophthora, P. infestans causing late blight of potato, others causes mostly root rots.
Family: peronosporaceae (the downy mildews) - sporangia borne on sporangiophores of
determinate growth. Sporangia are wind-borne. Obligate parasite.
Genus: plasmopara, P. viticola causing downy mildew of grape.
Peronospora, P. tabacina causing downy mildew (blue mould) of tobacco.
Bremia, B. lactucae causing downy mildew of lettuce.
Pseudoperonospora, P. cubensis causing downy mildew of cucurbits.
Peronosclerospora causing downy mildew of corn (P. philippinensis), of sugarcane
and corn (P. sacchari), of sorghum (P. sorghi), and others.
Sclerophthora causing the crazy top downy mildew of corn.
Sclerospora causing downy mildew of pearl millet and many other grasses.
Family: Albuginaceae (the white rusts) – sporangia borne in chains.
Albugo, A. candida causing white rust of crucifers.

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THE TRUE FUNGI (KINGDOM FUNGI)
Produce mycelium, the walls of which contain glucans and chitin. Lack chloroplasts.
Phylum: CHYTRIDIOMYCOTA produce zoospores that have a single posterior flagellum.
CLASS: CHYTRIDIOMYCETES- have round or elongated mycelium that lacks cross walls.
Genus: Olpium, O. brassicae being parasitic in roots of cabbage and other plants.
Can transmit plant viruses. Physoderma, P. maydis causing brown spot of corn.
Synchytrium, S. endobioticum causing potato wart. Urophlyctis, U. alfalfae causing
crown wart of alfalfa
Phylum: ZYGOMYCOTA – produce nonmotile asexual spores in sporangia. No zoospores.
The resting spore is a zygospore, produced by the fusion of two morphologically similar
gametes.
CLASS: ZYGOMYCETES (the bread moulds) – Saprophytic or parasites of plants,
humans and animals.
Order: Mucorales – Nonmotile asexual spores formed in terminal sporangia.
Genus: Rhizopus, causing bread moulds and soft rots of fruits and vegetables.
Choanephora, C. Cucurbitarum causing soft rot of squash.
Mucor, causing bread mould and storage rots of fruits and vegetables.
Order: Glomales – Fungi causing vesicular-arbuscular mycorrhizae, also known as
endomycorrhizae. Arbuscules procured in host roots. Chlamydosporelike spores
produced singly in soil, in roots, or in sporocarps. Sexual reproduction rate.
Genus: Glomus, Acaulospora, Gigaspora, Scutelloslpora.
Phylum: ASCOMYCOTA (Ascomycetes, the sac fungi) - Most have a sexual stage
(teleomorph) and an asexual stage (anamorph). Produce sexual spores, called ascospores,
generally in groups of eight within an ascus. Produce asexual spores (conidia) on free
hyphae or in asexual fruiting structures (pycnidia, acervuli, etc.).
A. CLASS: ARCHIASCOMYCETES – A group of diverse fungi, difficult to characterize.
Order: Taphrinales – Asci arisng from binucleate ascogenous cells.
Taphrina, causing peach leaf curl, plum pocket, oak leaf blister, etc.
B. CLASS: SACCHAROMYCETES (the yeast fungi) – Asci naked, no ascocarps produced.
Mostly unicellular fungi that reproduce by budding.
Genus: Galactomyces, causing citrus sour rot.

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 Saccharomyces, S. cervisiae, the bread yeast

FILAMENTOUS ASCOMYCETES
Order: Erysiphales (the powdery mildew fungi) – Asci in fruiting bodies completely closed
(cleistothecia). Mycelium, conidia, and cleistothecia on surface of host plant.

Obligate parasites:
Genus: Blumeria, causing powdery mildew of cereals and grasses.
Erysiphe, causing powdery mildews of many herbaceous plants.
Leveillula, causing powdery mildew of tomato and pepper.
Microsphaeria, one species causing powdery mildew of apple.
Sphaerotheca, S. pannosa causing powdery mildew of roses and peach.
Uncinula, U. necator causing powdery mildew of grape.

A.PYRENOMYCETES: ASCOMYCETES WITH PERITHECIA - Perithecia or,

occasionally, cleistothecia in a stroma, immersed in a loose hyphal mat, or free.
Asci have one wall.
Order: Hypocreales – Stromata pale to blue, purple or brightly colored. Asci ovoid to
cylindrical with apical pore. Conidia produced from phialidic conidiophores. Some
produce substances toxic humans and animals. Some produce growth regulators. Some
are antagonistic or parasitic on other fungi and some are systemic parasites
(entophytes) of many grain crops and make them poisonous to grazing animals.
Genus: Hypocrea, some species of which produce anamorphs.
Trichoderma and Gliocladium, which are used as biocontrol agents against several
plant pathogenic fungi.
Melanoslpora, of which its anamorphs Phialophora and Gonatobotrys parasitize
the mycelium of many fungi, including the important plant pathogens.
Ophiostoma, Ceratocystis, Fusarium, and Verticillium.
Nectria, causing twig and stem cankers of trees.
Gibberella, causing foot or stalk rot of corn and small grains.
Claviceps, C. purpurea causing ergot of rye, which is poisonous to humans and animals.

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 Epichloe, endophytic in grasses and sedges.
Atkinsonella, endophytic in grasses and sedges.
Myriogoeenospora, endophytic in grasses and sedges
Order: Microascales- Lack stromata. Mpst have perihecia but some have cleistothecia.
Asci are globoid or ovoid, disintegrating. Ascospores one celled.
Ginus: Ceratocystis, causing oak wilt (C. fagacearum); cankers in stone fruit and
other trees and root rot of sweet potato (C. fimbriata); butt rot of pineapple
(C. paradoxa); sapstain or blue stain of cut wood surfaces (C. coerulescens and others).
Order: Phyllachorales – perithecia in stroma, asci oblong to cylindrical, with pores at their
tips. Ascospores of varying shapes, hyaline or dark.
Genus: Glomerella, G. cingulata causing many anthracnose diseases and bitter rot
of apples; its anamorphic stage is Colletotrichum gloeosporioides
Phyllachora, P. graminis causing lead spots on grasses.
Order: Ophiostomatales - Perithecia without plaraphyses. Asci globose to ovoid,
disintegrating. Several species are dispersed by beetles. Some species cause
sapstain (blue stain) in wood.
Genus: Ophiostroma, O. ulmi ( formely Ceratocystis ), causing the Dutch elm
disease, O. novo-ulmi, causing a more severe form of Dutch elm disease, is
replacing O. ulmi in nature ( anamorphs are Sporothrix and Graphium ).
Order: Diaporthales – Perithecia in a substrate of either fungal and substrate tissue, or of
hyphae on substrate. Asci cylindrical with pores. Ascospores have one to several
septa and may be hyaline to brown.
Genus: Diaporthe, causing citrus melanose (D.citri), eggplant fruit rot (D.vexans),
soybean lpod and stem rot (D. Phaseolorum), their anamorphs are species of
phomopsis.
Gnomonia, causing anthracnose and leaf spot diseases.
Gaeumannmyces, G. graminis causing the take-all disease of grain crops
Magnaporthe, M. grisea causing the very important rice blast disease, its anamorph
is pyricularia oryzae.
Cryphonectria (formerly Endothia), C. parasitica causing the chestnut blight disease.
Leucostoma (formerly Valsa ), causing canker diseases of peach and other trees.

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Order: Xylariales – Perithecia dark, leathery, hard, sometimes embedded in a stroma. Asci
cylindrical to subglobose. Ascospores one to a few-celled, hyaline or dark.
Genus: Hypoxylon, H. mammatum causing a severecandker on poplars.
Roselinia, R. necatrix causing root diseases of fruit trees and vines.
Xylaria, causing tree cankers and wood decay.
Eeutypa, E. armeniacae causing serious canker diseases of fruit trees and vines.
B.LOCULOASCOMYCETES: ASCOMYCETES WITH ASCOSTROMATA – Produce
asci within locules (cavities) preformed in stroma. Ascostroma may be monolocular
(pseudothecium ) or multilocular. Asci have a double wall.
Order: Dothideales – Locules lack sterile hyphae and open by an apical pore, Asci ovoid,
cylindrical, in fasciles. Ascospores one to several celled, hyaline to brown.
Genus: Mycosphaerella, causing leaf spots on many plants, such as the Sigatoka
diseases of banana (M. musicola and M. fijiensis), and leaf spot of strawberry
(M. fragariae), its anamorphs may be Cercospora, Septoria, and others.
Elsinoe, causing citrus scab (E. faucetii), grape anthracnose (E. ampelina), and
raspberry anthracnose ( E. veneta).
Order: Capnodiales – Ascocarps superficial, produced in a loose mat of dark hyphae.
Genus: Capnodium, being one of several fungi causing sooty moulds on plants.
Order: Pleosporales – Asci surrounded by pseudoparaphyses. Ascostroma variable.
Genus: Cochliobolus, whose anamorphs are Bipolaris or Curvularia, causes leaf spots
and root rots on grain crops and grasses.
Pyrenophora, whose anamorph is Drechslera, causing leaf spots on cereals and
grasses.
Septosphaera (anamorph is Exserohilum), causing leaf spots on cereals and grasses.
Pleospora (anamorph is Stemphylium), causing black mould rot of tomato.
Leptosphaeria (anamorph is Phoma), causing black and foot rot of cabbage.
Venturia (anmorph is Spilocaea), causing black rot of grapes.
Dibotryon , D. morbosum causing black rot of cherries and plums.
C. DIASCOMYCETES: ASCOMYCETES WITH APOTHECIA – Ascocarps shaped as
cups, saucers, or cushions and called apothecia. Asci cylindrical to ovoid, often
interspersed with paraphyses. Ascospores discharged forcibly.

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Order: Rhytismales – Ascocarps are black, spherical, discoid, or elongate, and are
produced in stromata. Asci variable. Ascospores hyaline or brown, ovoid to filiform.
Genus: Hypoderma, causing pine leaf spot (needle cast) diseases.
Lophoderminum , causing pine needle blight.
Rhabdocline, causing pine cast disease.
Rhytisma, R. ascerinum causing tar spot of maple leaves.
Order. Helotiales – Apothecia cup - or disk-shaped. Asci with only slightly thicket apices.
Ascospores are spherical, elongate, to filiform, and have one to several septa.
Genus: Monilinia, causing the brown rot disease of stone fruits.
Sclerotinia, S. sclerotiorum causing the white mould or watery soft rot of vegetables.
Stromatinia, S. gladioli causing corm rot of gladiolus.
Pseudopeziza, P. trifolii causing alfalfa leaf spot
Diplocarpon, D. maculatum causing black spot of quince and pear, and black spot of
roses (D. rosae).
Sclerotium , S. cepivorum causing the white rot of onions.
D. DEUTEROMYCETES (imperfect or asexual fungi) - Mycelium well-developed, septate,
branched. Sexual reproduction and structures rare, lacking, or unknown. Asexual
spores (conidia) formed on conidiophores existing singly, grouped in specialized
structures such as sporodochia and synemata, or produced in structures known as
pycnidia and ascervuli.

Anamorphic stage Certain/likely teleomorphic

group
Genus : Geotrichum , G. candidum causing sour
rot of fruits and vegetables. Saccharomycetales
Penicillium, causing blue mould rot of fruits Cleistothecial Ascomycetes
Aspergillus, causing bread mould and seed decays Talaromyces
Paecilomyces, used as biological control agent Eurotium
against white flies Byssochlamys
Oidium, causing the powdery mildews Erysiphe, Etc.
Chalara, causing oak wilt, tree cankers Perithecial Ascomycetes
Acremonium, endophytic in grasses Ceratocystis

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 Sporothrix and Graphium, causing the Dutch elm disease Epichloe
Trichoderma, used as biocontrol agent against other fungi Ophiostoma
Verticillium , causing vascular wilts in many plants Hypocrea
Fusarium , causing vascular wilts, root rots, stem rots, Hypocrea
seed infections Gibberella
Colletotrichum , causing anthracnose in many plants Glomerella
Cercospora, causing Sigatoka disease of bananas Loculoascomycetes
Septoria, causing leaf spots on many crops Mycosphaerella
Phyllosticta, causing black rot of grape Mycosphaerella
Alternaria, causing many leaf spots, blights Gulgnardia
Stemphylium, causing fruit rots on tomato Lewia
Bipolaris, causing leaf spots and root rots in grasses Pleospora
Drechslera, causing leaf spots on grasses Cochliobolus
Exserohilum, causing leaf spots on grasses Pyrenophora
Curvularia, causing leaf spots on grasses Setosphaera
Cladosporium, causing leaf moulds on tomato (C.fulvum), Cochliobolus
and scab of peach Fulvia, Venturia and almond Sphaeropsis
(C. carpophilum) causing black rot on apple. Physalospora
Botrytis, B. cinerea causing gray mould rots on many Apothecial Ascomycetes
plants Botryotinia
Monilia, causing the brown rot of stone fruits Monilinia
Marssonina, causing the black spot of rose Diplocarpon
Entomosporium, causing a leaf and fruit spot on pear Diplocarpon
Cylindrosporium, causing leaf spots on many knids of Mycosphaerella
plants Greeneria
Melanconium, causing the bitter rot of grape
Rhizoctonia, R. solani causing root and stem rots Basidiomycetes
Rhizoctonia binucleate forms Thanatephorus
Sclerotium, S. rolfsii causing southern blight of many Ceratobasidiales
crops Aethalium

Phylum: BASIDIOMYCOTA (Basidiomycetes, the club and mushroom fungi)- Sexual

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 spores, called Basidiospores, are produced externally on a club-like, one- or four –
celled spore - producing structure called a basidium.
Order: Ustilaginales (the smut fungi) - Basidium has cross-walls or is non-septate. It is the
promycelium of the teliospore. Teliospores single or united into clusters or columns,
remaining in host tissue or bursting through the epidermis. Fertilization by union of
compatible spores, hyphae, etc. Only teliospores and basidiospores are produced.
Genus: Ustilago, causing smut of corn (U. maydis), loose smuts of oats (U. avenae), of
barley
(U. nuda) and wheat (U. tritici). Tilletia, causing covered smut or bunt of wheat (T.
caries), and kernel bunt (Partial bunt) of wheat (T. indica). Urocystis, U. cepulae
causing smut of onion. Sporisporium, causing covered kernel smut of sorghum
(S. sorghi) and loose smut sorghum smut (S. cruentum). Sphacelotheca, causing head
smut of sorghum (S. relianum).
Order: Uredinales (the rust fungi) - Basidium with cross-walls. Sperm cells called spermatia
fertilize special receptive hyphae in spermogonia. Produce two to several types of
spores: telioslpores, basidiospores, aeciospores, and uredospores. Uredospores can be
repeating spores. Obligate parasites.
Genus: cronartium, several species causing stem rusts of pines.
Gymnosporangium, G. Juniperi-virginianae causing cedar-apple rust.
Hemeleia, H. vastatrix causing coffee rust disease.
Melampsora, M. lini causing rust of flax.
Phakopsora, P. pachyrrhizi causing rust of soybeans.
Phragmidium, one species causing rust of roses
Puccinia, several species causing severe rust diseases of cereals and other plants.
Uromyces, U. appendiculatus causing rust of beans.
Order: Exobasidiales - Basidiocarp lacking: basidia produced on surface of parasitized tissue.
Genus: Exobasidium, causing leaf, flower, and stem galls on several ornamentals.
Order: Ceratobasidiales – Basidiocarp is web like, inconspicuous. Basidium without cross-
walls, with four prominent sterigmata.
Genus: Thenatephorus, T. cucumeris is the teleomorph of Rhizoctonia solani, causing
root and stem rots, damping-off and fruit rots in many plants. Typhula, causing

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 typhula blight (snow mould) of turf grasses.
Order: Agaricales (the mushrooms) – Basidium without cross walls, produced on radiating
gills or lamellae. Many are mycorrhizal fungi.
Genus: Armillaria, A. mellea causing root rots of forest and fruit trees.
Crinipellis, C. pernicious causing witches’-broom or cocoa in Central and South
America. Marasmius, causing the fairy ring disease of turf grasses.
Pleurotus, causing white rot on logs, tree stumps, and living trees.
Pholiota, causing brown wood rot in deciduous forest trees.
Order: Aphylloophorales (the polypores) - Basidia without cross walls produced on hymenia-
forming hyphae and lining the surfaces of small pores or tubes.
Genus: Aethalium (Sclerotium), causing root and stem rots of many plants.
Chondrostereum, C. purpureum causing the silver leaf diseases of trees
Corticium, one species causing the red thread disease of turf grasses.
Heterobasidion, H. annosum causing heart rot of many trees.
Ganoderma, causing root and basal stem rots in many trees.
Inonotus, causing a heart rot of living trees and rot of dead trees and logs
Postia, causing wood and root rots of forest trees.
Phellinus, (Poria) causing tree root rots and cubical rots in buildings.
Peniophora, causing decay in coniferous logs and pulpwood.
Polyporus, causing heart rot of living trees and rot of dead trees or logs.

3. INFECTIOUS CAUSES OF PLANT DISEASES/ETIOLOGY

3.1. Fungi as Plant Pathogens
Living things are organized for study into large, basic groups called kingdoms. Fungi were listed
in the Plant Kingdom for many years. Then scientists learned that fungi show a closer relation to
animals, but are unique and separate life forms. Fungi are not plants. Now, Fungi are placed in

16
their own Kingdom. They are grouped under the kingdom Mycota (fungal kingdom). The living
body of the fungus is a mycelium made up of a web of tiny filaments called hyphae. The
mycelium is usually hidden in the soil, in wood, or another food source.

Fungi comprise about 100,000 species and more than 10,000 species cause plant diseases. Fungi
are small, generally microscopic, eukaryotic (= having a nucleus), usually filamentous, branched,
spore-bearing organisms that lack chlorophyll. 80% of plants diseases are caused by fungi while
the remaining 20% by bacteria, viruses, nematodes, etc. Fungi could be non-obligate parasites or
saprophytes, which can grow and multiply on dead organic matter as well as on living plants.
Some fungi, known as obligate parasites or biotrophs, which can grow and only multiply in
association with their host plants during their entire life. Others, known as non-obligate parasites
can be facultative saprophytes or facultative parasites depending on whether they are primarily
parasites or primarily saprophytes. Fungi feed by absorbing nutrients from the organic material
in which they live. They must digest their food before it can pass through the cell wall into the
hyphae. Hyphae secrete acids and enzymes that break the surrounding organic material down
into simple molecules they can easily absorb.

Fungi have evolved to use a lot of different items for food. Some are decomposers living on dead
organic material like leaves. Some fungi cause diseases by using living organisms for food.
These fungi infect plants, animals and even other fungi. Athlete’s foot and ringworm are two
fungal diseases in humans. The mycorrhizal fungi live as partners with plants. They provide
mineral nutrients to the plant in exchange for carbohydrates or other chemicals fungi cannot
manufacture. People eat mushrooms of all shapes, sizes and colors. Yeasts are used in making
bread, wine, beer and solvents. Drugs made from fungi cure diseases. Fungi are also grown in
large vats to produce flavorings for cooking, vitamins and enzymes for removing stains.

The main characteristics of fungi:

Nutrition: Heterotrophic and absorptive; ingestion of food is rare
Thallus: Unicellular, filamentous (branched mycelium)
Cell wall: Made up of chitin (cellulose only in class Oomycetes)
Nucleus: Eukaryotic organisms

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Life cycle: Simple to complex
Reproduction: Asexual and sexual spores
Distribution: Cosmopolitan
Structure or Morphology of Fungi
• A filamentous vegetative body in most fungi (mycelium)
• Some lower fungi lack true mycelium

Habitat of fungi
• Air, water, soil and dead organic matter. Suitable temperature is 0 to 40oC.
Reproduction
Fungi reproduce chiefly by means of spores
Spores are reproductive bodies consisting of one or a few cells
Spores can be classified into two.
• Asexual spores which include:-
Zoospores: have a flagellum (can swim)
: produced inside a sac (sporangium)
Conidia: produced on conidiophores, either directly on the mycelium or inside walled
structures (conidiomata)
Pycnidium: a distinctive form of flask-shaped conidiomata
Chlamydospores: round, a thick wall asexual spore

• Sexual spores: produced by Ascomycetes, Basidiomycetes & Oomycetes

Ascospores: usually eight in number, produced within a sac-like zygote cell (ascus)
Basidiospores: produced on the outside of a club-like zygote (basidium)
Oospores: gametangia of unequal size fuse to form zygotes

Spores! THE agents of fungal dispersal

Structures for long distance dispersal: Sexual and asexual spores of fungi

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Ecology of fungi
Almost all plant pathogenic fungi spend part of their lives on their host plants, in the soil or in
plant debris on the soil. Some fungi are strictly biotrophs (obligate), i.e., they spend all of their
lives on the host, and only the spores may land on the soil, where they die or remain inactive).

Dissemination
Fungi are disseminated primarily in the form of spores. Agents of dissemination: wind, water,
birds, insects, other animals and humans. Wind is probably the most important disseminating

19
agent of spores of most fungi; water or insects may play a much more important role. Spore
dissemination in almost all fungi is passive

3.2. Bacteria as Pathogens

• About 1,600 bacterial species are known.

• Most are strictly saprophytic (= lacking an organized nucleus), i.e., Archaea (preexisting
ancient prokaryotes) and Bacteria (unicellular prokaryotes)
• Beneficial to humans because they help decompose the enormous quantities of organic
matter produced yearly by humans, animals, & factories as waste products or by the death
of plants and animals
• About 100 species of bacteria cause diseases in plants
• Most plant pathogenic bacteria are facultative saprophytes and can be grown artificially
on nutrient media (culture media)
• Bacteria may be rod shaped, spherical, spiral, or filamentous (threadlike)
• Some bacteria can move through liquid media by means of flagella

Some Important Bacterial Diseases of Plants and their Causal Agents

Pathogen Disease
1. Pseudomonas (aerobic, Gram –ve rods)
P. solonacearum ‘Ring-rot’ (wilt) of potato
P. gladicola (= P. allicola) Onion rot
P. syringe ‘Fire-blight’ of potato

2. Xanthomonas (aerobic, Gram –ve rods)

X. citri Citrus canker
X. malavacearum Angular leaf-spot of cotton
3. Agrobacterium (aerobic, Gram –ve rods)
A. tumefaciens ‘Crown’ gall of dicot plants
A. rhizogenes ‘Hairy root’ disease of plants
4. Erwinia (Gram –ve, facultative anaerobic rods)
E. carotovora Soft rot of carrot

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 E. atroseptica ‘Black-leg’ of potato
E. amylovora ‘Fire blight’ of apple
5. Corynebacterium (none-spore forming, Gram
+ve rods)
C. sepedonicum ‘Ring-rot’ (wilt) of potato
C. tritici ‘Gumming’ disease of wheat
6. Streptomyces (Gram +ve, filamentous)
S. scabies ‘Scab’ of potato

The filamentous bacteria Streptomyces can produce spores called conidia. Bacteria can multiply/
produce tremendous numbers of cells in a short period of time). Bacterial diseases are
particularly common and severe in the humid tropics.

Characteristics of Plant Pathogenic Bacteria

Morphology
Most plant pathogenic bacteria are rod shaped except Streptomyces (filamentous). The cell walls
of bacteria of most species are enveloped by a viscous, gummy material. When a single
bacterium is allowed to grow (multiply) on the surface of or in a solid medium, its progeny soon
produces a visible mass called a colony. Colonies of most species are whitish/grayish, but some
are yellow. Some bacteria pathogens (Erwinia amylovora) produce their populations in the plant
host, while in the soil their numbers decline rapidly. Some other bacterial pathogens
(Agrobacterium tumefaciens, Rolstonia solanacearum) cause the bacterial wilt of solonaceous
crops, and particularly Streptomyces scabies are rather typical soil inhabitants.
When in soil, bacteria live mostly on plant material. Less often they live freely or
saprophytically; may also survive in or on seeds, other plant parts, or insects found in the soil.
The dissemination of plant pathogenic bacteria from one plant to another is carried out primarily
by water, insects, other animals, and humans. Insects not only carry bacteria to plants, but they
inoculate the plants with bacteria by introducing them into the particular sites in plants where
they can almost surely develop.
Bacteria have thin, relatively tough, rigid cell walls and an inner cytoplasmic membrane. The
cell wall allows the inward passage of nutrients and the outward passage of waste matter and

21
digestive enzymes. Bacteria also have single or multiple copies of additional, smaller circular
genetic material called plasmids.
Reproduction
Rod-shaped phytopathogenic bacteria reproduce by the asexual process (binary fission or
fission).
Ecology and Spread
Almost all plant pathogenic bacteria develop as:
• Parasites ( in the host plant)
• Epiphytes (on the plant surface) and
• Saprophytes (in plant debris/in the soil)
Identification of Bacteria
The main characteristics of some of the most common plant pathogenic genera of bacteria are as
follows.
Agrobacterium
• Motile by means of one to four peritrichous flagella
• The colonies are non-pigmented and usually smooth
• They are rhizosphere and soil inhabitants.
Clavibacter (Corynebacterium)
• Straight to slightly curved rods and generally non-motile
Erwinia
• Motile by means of many peritrichous flagella
• The only plant pathogenic bacteria that are facultative anaerobes

Pseudomonas
• Motile (one or many polar flagella)
• Many species are common inhabitants of soil, fresh water and marine environments
• Produce yellow-green, diffusible, fluorescent pigments in a medium of low iron content
Xanthomonas
• Motile by means of a polar flagellum
• Growth on agar media is usually yellow

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 • All species are plant pathogens and are found only in association with plants or plant
materials
Streptomyces (Bacteria)
• Have the shape of slender, branched hyphae without cross walls
• The aerial mycelium forms chains of three to many spores
• Many species and strains produce a wide variety of pigments that color the mycelium,
viruses, protozoa, or tumor tissues
• All species are soil inhabitants and they are gram positive
Symptoms Caused by Bacteria
Plant pathogenic bacteria induce as many kinds of symptoms on the plants they infect as do
fungi. They cause leaf spots & blights, soft rots of fruits, roots, & storage organs, wilts,
overgrowths, scabs, and cankers. Species of Agrobacterium can cause only overgrowths or
proliferation of organs.

3.3. Viruses and Viroids as Pathogens

Definition
“Viruses are entities whose genome is a nucleic acid, either DNA or RNA, which reproduce
inside living cells and use their synthetic machinery to direct the synthesis of specialized
particles-the virions, which contain the viral genome and transfer it to other cells. Viron is a
technical term for the virus.

In 1898, Friedrich Loeffler and Paul Frosch found evidence that the cause of foot-and-mouth
disease in livestock was an infectious particle smaller than any bacteria. This was the first clue to
the nature of viruses, genetic entities that lie somewhere between living and non-living states.
Viruses depend on the host cells that they infect to reproduce. When found outside of host cells,
viruses exist as a protein coat or capsid, sometimes enclosed within a membrane. While outside
the cell, the virus is metabolically inert.

When it comes into contact with a host cell, a virus can insert its genetic material into its host,
literally taking over the host's functions. An infected cell produces more viral protein and genetic
material instead of its usual products.

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 • Tobacco mosaic virus in tobacco plants B) virus that attacks bacteria (bacteriophage)

Viruses:
• Nucleoprotein that multiplies only in living cells and have the ability to cause disease
• The total number of viruses known to date exceeds 2,000
• Too small to be seen individually with a light microscope
• All parasitize cells and cause a multitude of diseases in all forms of living organisms
including fungi and bacteria
• A plant may sometimes be infected by more than one kind of virus at the same time
• Behave like microorganisms in that they have genetic functions (able to reproduce, and
cause disease); they also behave as chemical molecules.
• Consist of nucleic acid (5 to 40%) and protein (60 to 95%), with a protective coat
(capsid) around the nucleic acid.
• Have no cytoplasm, ribosome, nuclear membrane, mitochondria, cell membrane, and
other organelles
• Multiply/replicate/duplicate intra cellular (within the cell)
• Have no enzyme system

The capsid (protein coat) surrounding the internal nucleoid has the following functions:
• It protects the nucleic acid from unfavorable extra-cellular environment

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 • It facilitates nucleic acid entry into the host cells
• It is antigenic

They are mostly rod shaped, polyhedral, or variants of these two basic structures. The nucleic
acid is called nucleoid (which may be DNA or RNA, but never both) and forms the genome.
Viruses do not divide and do not produce any kind of specialized reproductive structures such as
spores; instead, they multiply by inducing host cells to make more viruses. Viruses cause disease
not by consuming cells/killing them with toxins, but by utilizing cellular substances during
multiplication, taking up space in cells, and disrupting cellular processes. Because of their small
size and the fact that they are transparent, viruses generally cannot be viewed and detected by the
methods used for other pathogens. Cell inclusions consisting of virus particles (virus aggregates)
however, are visible by light microscopy. Viruses are not cells nor do they consist of cells.

Morphology
Plant viruses have different shapes and sizes:
• elongate (rigid rods or flexuous threads)
• many are spherical (isometric or polyhedral),
• Cylindrical bacillus-like rods
Classification
• Division one - DNA viruses
• Division two - RNA viruses

Naming of viruses are remarkable based on their symptoms.

e.g., TMV - Tobacco mosaic virus
PLR - Potato leaf roll virus
Chlorotic or necrotic ring spots

Most other symptoms caused by viruses resemble those caused by mutations, nutrient
deficiencies or toxicities, insect or mite feeding damage, other pathogens, and other factors.
Almost all viral diseases seem to cause some degree of dwarfing or stunting of the entire plant
and reduction in total yield. Viruses usually shorten the length of life of virus-infected plants,

25
although they rarely kill plants on infection. Viral diseases of plants may be systemic (spread
throughout the host plant) or, local (restricted to certain areas). Mostly the symptoms appear on
leaves.

The important symptoms of viral plant diseases:

• Mosaic: Mixed light green & yellow patches appear on leaves. This is caused by systemic
infection (Tobacco mosaic, mosaic of potato, mosaic of sugarcane, etc.).

• Yellows: Uniform chlorosis of the leaves (Rice yellows).

• Vein clearing (Vein banding and Vein thickening): yellowing of the veins & the veinlets.
The rest of the leaf remains green.
• Leaf curling or Leaf rolling: a very common symptom (e.g., Leaf curl of papaya, tomato
and potato).
• Stunting: General retardation of growth results in stunting of plants (e. g., Bunchy top of
banana).
• Breaking & greening of plants: Changes in color of petals give them a variegated
appearance. When petals become green due to virus infection, it is called virescence.

26
 • Tumors: Tumors are outgrowths produced by uncontrolled division & growth of cells.
These appear on leaves or roots.
• Witches broom: The leaves become reduced in size. The internodes are shortened and the
abnormal growth of leaves results in a densely packed structure called witches broom.
Usually many viral symptoms appear simultaneously in the same plant.

Transmission of Plant Viruses

Viruses being non-motile are carried passively from one place to other. The following are the
main means of virus transmission:
Transmission by contact: The most important being the TMV. Sometimes, the sap of the infected
plants comes in contact with healthy plants through implements. Grafting, artificial or natural
(cuscuta or dodder causes natural grafting by carrying the virus from one plant to others), is
another important method of contact transmission.
Transmission through seeds & vegetative organs: Seeds produced by diseased plants carry virus
particles in their seed coats, endosperm or embryo. Organs of vegetative propagation like the
tubers, bulbs, rhizomes, stem cuttings, etc. derived from infected plants, give rise to infected
plants.

Transmission through nematodes and fungi: The nematodes and soil fungi are important carriers
(vectors) of viruses. These vectors are carried to long distances through irrigation water; and
along with them, the viruses, that they harbor, reach far off places. In fungi, the unicellular water
molds (Chytrids) are vectors of several viruses. The viruses contained in plant debris also reach
long distances through water.

Transmission through insects: No other group of diseases is so much dependent on insects for
transmission as are the viruses. Aphids, leaf-hoppers, thrips, beetles and mites transmit large
number of viruses. Aphids are most important among all insect vectors.

Viroids
• At least 40 plant diseases have been shown to be caused by viroids.

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 • Viroids are small, low molecular weight ribonucleic acids that can infect plant cells,
replicate themselves, and cause disease.
• Viroids differ from viruses in at least two main characteristics:
• The size of RNA in viroids (250 to 370 bases) is much smaller compared to that in
viruses (4 to 20 kilo bases)
• Viroids lack a protein coat and apparently exist as naked RNA

Diseases caused by some viroids

• Potato spindle tuber viroid
• Tomato apical stunt viroid
• Citrus exocortis viroid
• Tomato chlorotic dwarf viroid

How viroids replicate themselves is still not known. How viroids cause disease is also not
known. Viroid diseases show a variety of symptoms that resemble those caused by virus
infections. Viroids are spread from diseased to healthy plants primarily by mechanical means
(through sap carried on hands or tools during propagation or cultural practices and by vegetative
propagation). Viroids apparently survive in nature outside the host or indeed plant matter for
periods of time varying from a few minutes to a few months. Generally, they seem to overwinter
and oversummer in perennial hosts. The control of viroids is based on the use of viroid-free
propagating stock, removal and destruction of viroid-infected plants, and washing of hands or
sterilizing of tools after handling viroid infected plants before moving on to healthy plants.

3.4. Nematodes

• Belong to the kingdom Animalia

• Wormlike in appearance but quite distinct taxonomically from the true worms
• Most of the several thousand species live freely in fresh or salt waters or in the soil
• Several hundred species are known to feed on living plants, obtaining their food with
spears or stylets
• Cause a variety of plant diseases worldwide which include all grains and legumes,
banana, cassava, coconut, potato, sugar beet, sugarcane, sweet potato, and yam (11%)

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 • losses for most other economically important crop (vegetables and fruits) (about 14%, i.
e., over $80 billion annually)

Morphology and anatomy

Plant-parasitic nematodes are small, 300 to 1,000 micrometers, with some up to 4 mm long.
They can be observed easily under the microscope. In general, eel-shaped and round in cross
section, with smooth, unsegmented bodies, without legs or other appendages. Most plant
parasitic nematodes have hollow stylets or spear (e.g., Meloidogyne-root-knot nematode of
almost all crops). The stylet is used to puncture holes in plant cells and through which to
withdraw nutrients from the cells.

Reproduction and Life cycle

The female can produce fertile eggs either after mating with a male or, in the absence of males,

parthenogenetically. Egg ↔ Larvae (4 juvenile stages) ↔ Adult

Ecology and Spread

Nematodes spread through the soil slowly under their own power. In addition to their own
movement, nematodes can be spread easily by anything that moves & can carry particles of soil.
Farm equipment, irrigation, flood or drainage water, animal feet, birds, and dust storms spread
nematodes in local areas, whereas over long distances nematodes are spread primarily with farm
produce and nursery plants. In terms of habitat, pathogenic nematodes are either ectoparasites
(species that do not normally enter root tissue but feed only from the outside on the cells near the
root surfaces) or endoparasites (species that enter the host and feed from within). Both of these
can be either migratory (they live freely in the soil and feed on plants without becoming
attached) or move around inside the plant, or sedentary (species that, once within a root, do not
move about). Control is usually attempted through cultural practices (use of clean planting stock,
crop rotation, fallow, and cover crops; resistant varieties and natural or genetically engineered
antagonistic or parasitic bacteria.

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3.5. Mycoplasma-Like Organisms (MLO) and Related Pathogens

Mycoplasmas are prokaryotic organisms that have no cell walls. They are Members of the class
Mollicutes (genus Mycoplasma, Acholeplasma, and Spiroplasma). As they lack a true cell wall,
mycoplasmas are bounded only by a “unit” membrane. Mycoplasmas are small, sometimes
ultramicroscopic cells containing cytoplasm, randomly distributed ribosomes, and strands of
nuclear material. They reproduce by budding and by binary transverse fission of cells.
Mycoplasmas have no flagella, produce no spores, and are gram negative. Phytoplasmas (and
spiroplasmas) are generally present in the sap of a small number of phloem sieve tubes. Most
plant mollicutes (Mycoplasma, Acholeplasma, & Spiroplasma) are transmitted from plant to
plant by leafhoppers, phyllods and plant hoppers. More insects become vectors when feeding on
young leaves and stems of infected plants than on older ones.

Rikettsia-like Organisms (RLO) Or Xylem-inhabiting Fastidious Vascular Bacteria

Generally rod-shaped cells (0.2 to 0.5 Mm in diameter by 1 to 4 Mm in length) and Gram-
negative. They are bounded by a cell membrane and a cell wall with no flagella. Several such
xylem-limited bacteria have been placed in the genus Xylella. The symptoms of diseases often
consist of marginal necrosis of leaves, stunting, and general decline and reduced yields.

3.6. Parasitic Higher Plants

More than 2500 species of higher plants are known to live parasitically on other plants. Their
main common characteristic is that these parasites are vascular plants that have developed
specialized organs (haustorium) which penetrate the tissues of other (host) vascular plants and
absorb nutrients from them. These parasitic plants produce flowers & seeds & belong to several
widely separated botanical families. The most common & serious parasites belong to the
following botanical families and genera.

3.6.1. Cuscutaceae
Genus: Cuscuta, the dodders of alfalfa, onion, potato, and numerous other plants. Some species
prefer legumes, whereas others attack many other broadleaved plants as well as legumes.

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Dodder is a slender, leafless, twining parasitic seed plant with little or no chlorophyll. The stem
is usually yellowish or orange in color, sometimes it is almost white. The haustoria penetrate the
stem or leaf & reach into the vascular tissues, from which they absorb foodstuffs & water. The
infected host plants become weakened; their vigor declines and they produce poor yields (many
are even killed). The seed may be spread to nearby areas by animals, water, & equipment, &
over long distances by contaminated crop seed.

3.6.2. Orobanchaceae
Genus: Orobanche, the broomrapes of legumes, solanaceous, & other plants. Orobanche sp., is a
whitish to yellowish-brown annual plant 15 to 50 centimeters tall. Broomrapes overwinter as
seeds, which may survive in the soil for more than 10 years. Seeds germinate only when roots of
certain susceptible host plants grow near them. Plants affected by broomrapes usually occur in
small patches & may be stunted to various degrees.

3.6.3. Scrophulariaceae
Genus: Striga, the witch weeds of many monocotyledonous and some legume plants. Witch
weed is a serious, well-known semi-root parasite in Africa, Asia and Australia, parasitizing
important economic plants (corn, sugarcane, rice, cowpea, tobacco and some other small grains).
Affected plants remain stunted, wilt, and turn yellowish. Infected roots bear a large number of
witch weed haustoria, which are attached to the root & feed on it.

4. NON-INFECTIOUS PLANT DISEASES

The common characteristic of abiotic diseases is that they are caused by the lack or excess of
something that supports life. Noninfectious diseases occur in the absence of pathogens and
cannot, therefore, be transmitted from diseased to healthy plants. Noninfectious diseases may

31
affect plants in all stages of their lives (seed, seedling, mature plant, or fruit), and they may cause
damage in the field, in storage, or at the market. The symptoms caused by noninfectious
diseases vary in kind and severity with the particular environmental factor involved and with the
degree of deviation of this factor from its normal. The symptoms of several noninfectious
diseases are too indistinct & closely resemble those caused by several viruses, mollicutes, and
many root pathogens.

4.1. Unfavorable Meteorological Factors

Temperature Effects
Most kinds of plants growing best between 15 and 30°C. The minimum & maximum
temperatures at which plants can still produce normal growth vary greatly with the plant species
& with the stage of growth of the plant.
• Low temperature effects
Damage to crops caused by low temperature is far greater than that by high temperatures. Thus,
older, hardened plants are more resistant to low temperatures than young seedlings. Also,
different tissues or organs on the same plant may vary greatly in their sensitivity to the same low
temperature. Buds are more sensitive than twigs, flowers and newly formed fruits are more
susceptible to low temperatures than leaves; and so on.

Low temperature affects plants in two ways:

• Reduction of the rate of metabolism
• Death of plants due to freezing
Plants such as cabbage, alfalfa, and most perennials of the temperate zone can withstand
temperatures below freezing.

• High-Temperature Effects
Too high a temperature rarely occurs in nature. High temperature causes its effects in
conjunction with other environmental factors, particularly excessive light, drought, lack of
oxygen, or high winds accompanied by low relative humidity. High temperatures are usually
responsible for sunscald injuries appearing on the sun-exposed sides of fleshy fruits and

32
vegetables such as peppers, apples, tomatoes, onion bulbs, and potato tubers. High temperature
increases evapo-transpiration that results in discoloration, a water-soaked appearance, blistering,
and desiccation of the tissues beneath the skin, which leads to sunken areas on the fruit surface.
High temperatures may inactivate or accelerate certain enzyme systems & thus leading to
abnormal biochemical reactions and cell death. Wind is a means of transmission of pathogens.
Strong wind results in breakage and dropping of fruits. Low humidity increases evapo-
transpiration that results in wilting.

Status of Moisture Contents

• Low soil moisture effects

Plants suffering from lack of sufficient soil moisture usually remain stunted, pale green to light
yellow, have few, small and drooping leaves, if the drought continues, wilt and die. Even
perennial plants and trees are damaged by prolonged periods of drought and produce less growth,
small, scorched leaves and short twig, dieback, defoliation, and finally wilting and death. Plants
weakened by drought are also more susceptible to certain pathogens and insects.

• High soil moisture effects

Excessive soil moisture due to poor drainage of planted fields, gardens, or potted plants may
result in serious damage or death of plants. Poor drainage results in plants that lack vigor, wilt
frequently, and have leaves that are pale green or yellowish green. As a result of excessive soil
moisture caused by flooding or by poor drainage, the fibrous roots of plants decay, probably due
to the reduced supply of oxygen to the roots. Once parts of roots are killed, more damage is
done by facultative parasites that may be favored by the new environment. Thus, wilting of the
plants, which soon follows flooding, is probably the result of lack of water in the aboveground
parts due to death of the roots.

Low oxygen conditions in nature are generally associated with high soil moisture or high
temperatures. Lack of oxygen may cause the desiccation of roots of different kinds of plants in
water logged soils. A combination of high soil moisture and high soil or air temperature causes
root collapse in plants. Lack of sufficient light retards chlorophyll formation and promotes
slender growth with long internodes, thus leading to pale green leaves, spindly growth, and

33
premature drop of leaves and flowers (etiolation). Etiolated plants are usually thin and tall and
are susceptible to lodging.

4.2. Nutritional Disorders

Plants require several mineral elements for normal growth. Both major and trace elements are
essential to the plant. When essential elements are present in the plant in amounts smaller than
the minimum levels required for normal plant growth, the plants becomes diseased and exhibits
various external and internal symptoms. The symptoms may appear on any or all organs of the
plant, including leaves, stems, roots, flowers, fruits, and seeds. The kinds of symptoms produced
by deficiency of a certain nutrient depend primarily on the functions of that particular element in
the plant. When the deficiency is greater than a certain critical level, the plants develop acute or
chronic symptoms and may even die. Of the essential elements, those required by plants in large
amounts (nitrogen and potassium) are usually much less toxic when present in excess than are
elements required only in trace amounts (manganese, zinc, and boron).

The injury occurring from the excess of an element may interfere with the absorption or function
of another element and thereby lead to symptoms of a deficiency of the element being interfered
with. Excessive amounts of sodium salts, especially sodium chloride, sodium sulfate, and sodium
carbonate, raise the pH of the soil and cause what is known as alkali injury. The injury varies in
different plants and may range from chlorosis to stunting, leaf burn, wilting, and killing of
seedlings and young plants. Plants like wheat and apple are sensitive to alkali injury.

Herbicide Injury
Some of the most frequent plant disorders seem to be the result of the extensive use of herbicides
(weed killers). Most herbicides are safe as long as they are used to control weeds among the right
crop plants, at the right time, at the correct dosage, and when the correct environmental
conditions prevail. Affected plants show various degrees of distortion or yellowing of leaves,
browning, drying and shedding of leaves, stunting and even death of the plant. Much of this
damage is caused by too high doses of herbicides or by applications made too early in the season
or on too cold or too hot a day or when dust or spray droplets of an herbicide are carried by the
wind to nearby sensitive plants.

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4.3. Atmospheric Impurities

Almost all air pollutants causing plant injury are gases, but some particulate matter or dusts may
also affect vegetation. More serious and widespread damage is caused to plants by chemicals
such as ozone, sulfur dioxide, hydrogen fluoride, nitrogen dioxide, peroxyacyl nitrates, and
particulates. More frequently, most air pollutants are transported downwind from the urban or
industrial centers to areas that are several miles, often hundreds of miles and sometimes
thousands of miles, from the source. High concentrations of or long exposure to these chemicals
cause visible and sometimes characteristic symptoms (such as necrosis) on the affected plants.
The air at the earth’s surface consists primarily of nitrogen and oxygen (78 and 21%,
respectively) and the remaining 1% is water vapor and carbon dioxide. Plant injury by air
pollutants generally increases with increased light intensity, increased soil moisture and air
relative humidity, and increased temperature and with the presence of other air pollutants. Ozone
injures the leaves of plants exposed for even a few hours at concentrations of 0.1 to 0.3 ppm.
Many crop plants (alfalfa, bean, citrus, grape, potato, soybean, tobacco and wheat), and many
ornamentals and trees are quite sensitive to ozone, whereas some other crops (cabbage, peas,

35
peanuts, and pepper), are of intermediate sensitivity, and some (beets, cotton, lettuce, strawberry,
and apricot), are tolerant. Acid rain (below pH 5.6) is the result of human activities, primarily the
combustion of fossil fuels (oil, coal, and natural gas) and the smelting of sulfide ores. Prolonged
exposure to air pollutants seems to weaken plants and to predispose them to attack by insects, by
some pathogens, and by other environmental factors. The adverse effects of acid rain on the
microorganisms, plants, and fishes of rivers and lakes have been well documented.

4.4. Improper Cultural Practices

Almost every agricultural practice can cause damage when applied the wrong way, at the wrong
time, or with the wrong materials. Most commonly, however, losses result from the application
of chemicals, such as fungicides, insecticides, nematicides, and fertilizer, at too high
concentrations on plants sensitive to them. Spray injury resulting in leaf burn or spotting or
rosseting of fruit is common on many crop plants. Inadequate or excessive watering may cause
wilting or any of the symptoms described earlier. Potatoes stored next to hot water pipes under
the kitchen sink often develop black heart. Trees frequently grow poorly & their leaves are
chlorotic, curled/reddened because their trunk is girdled by fence wire. The roots of plants potted
in pots that are too small for their size are often badly distorted & twisted & the whole plant
grows poorly. Excessive/too deep cultivation between rows of growing plants may be more
harmful than useful because it cuts pulls many of the plants’ roots. When plants are adversely
affected by an environmental factor (low moisture, nutrient deficiency, air pollution, or freezing),
they are generally and concurrently weakened and predisposed to infection by one or more
weakly parasitic pathogens. For example, all the conditions mentioned earlier predispose annual
plants to infection by the fungus Alternaria and many perennial plants to infection by canker-
causing fungi. Herbicide injury is likely to be followed by root rots caused by Fusarium and
Rhizoctonia. Flooding injury is often followed by Pythium root infections.

5. KOCK’S POSTULATES AND PLANT DISEASE SYMPTOMS

5.1. Koch's Postulates

36
Robert Koch (1843–1910) was a medical doctor and a bacteriologist. He was the first to show a
disease of sheep and other animals was caused by a bacterium that he called Bacillus anthracis.
Koch set out the four criteria that must be satisfied before a microorganism isolated from a
diseased human, animal, or plant can be considered as the cause of the disease. These
steps/rules/criteria are known as “Koch’s postulates.”
• The suspected causal agent (bacterium or other MO) must be present in every diseased
organism (e.g., a plant) examined.
• The suspected causal agent (bacterium, etc.) must be isolated from the diseased host
organism and grown in pure culture.
• When a pure culture of the suspected causal agent is inoculated into a healthy susceptible
host plant, the host must reproduce the specific disease.
• The same causal agent must be recovered again from the experimentally inoculated and
infected host (the recovered agent must have the same characteristics as the organism in
step 2).

Koch’s rules are possible to implement, although not always easy to carry out, with fungi,
bacteria, parasitic higher plants, nematodes, most viruses and viroids, & the spiroplasmas. These
organisms can be isolated & cultured, or can be purified, & they can then be introduced into the
plant to see if they cause the disease. With the other pathogens, (some viruses, phytoplasmas,
fastidious phloem-inhabiting bacteria, protozoa, & even some plant pathogenic fungi that are
obligate parasites of plants), (the powdery mildew, downy mildew, and rust fungi), culture or
purification of the pathogen is not yet possible and the pathogen often cannot be reintroduced
into the plant to reproduce the disease. Despite the difficulties of carrying out Koch’s postulates
with some causal agents, they have been and continue to be applied, sometimes with certain
modifications, in all cases of disease.

5.2. Morphological Symptoms of Plant Diseases

In diseased plants the harmful physiological alterations inside the plant have their counterpart as
visible symptoms expressed externally on the plant. Thus diseased plants show a number of

37
visible symptoms, known collectively as the syndrome. The three general categories of
symptoms are as follows.

5.2.1. Necrotic symptoms (death of the infected tissue)

Those symptoms that result from cessation of function leading to death. Some of the necrosis
symptoms are:
Blight: means burnt appearance, rapid killing of foliage blossoms and twigs. It is caused by a
wide range of pathogens (e.g., leaf blights of potatoes)
Blotch: large irregular lesions on leaves, shoots, and stems
Rot or Decay: disintegration of dead tissues involves the necrosis of large areas of tissue often
complete organs.
Spot: lesions, usually defined, circular, oval in shape with a central necrotic area surrounded by
various colored zones.
Wilt: leaves or shoots loose their turgidity and drop.
Damping off: Seedlings collapse and die.
Dieback: a disease affects the youngest tissues first & progresses down the stem.
Gummosis: lesions or swelling associated with the exudation of gum (e.g., gummosis of citrus).

5.2.2. Hypoplsia
Those symptoms that result from under development or retardation of function.
Chlorosis/Discoloration: failure of chlorophyll development in normally green tissues.
Dwarfing: stunting growth in an entire plant or some of its parts.

5.2.3. Hyperplasia
Those symptoms that result from over development of or acceleration of function. Enlargement
of tissues owing to excessive production of cells.
Gall formation: a swelling or outgrowth commonly having globes, elongated or irregular shape.
Hairy root: numerous fine fibrous roots are produced which are abnormal.
Callus: overgrowth of tissues of the margins of wounds and diseased tissues (smut fungi in
maize cause tumor like swelling).

38
5.3. Effects of Plant Pathogens on Plant Physiology

While pathogens infect plants in the course of their obtaining food for themselves, they interfere
with the different physiological function(s) of the plant and lead to the development of different
symptoms.

Effect of Pathogens on Photosynthesis

Photosynthesis is the ultimate source of nearly all energy used in all living cells, plant or animal.
Those pathogens do interfere with photosynthesis cause chlorosis, necrotic lesions or large
necrotic areas on green plant parts, and reduced growth and fruits on many infected plants. In
leaf spot, blight, & other kinds of diseases in which there is destruction of leaf tissue,
photosynthesis is reduced because the photosynthetic surface of the plant is lessened. In plants
infected by many vascular pathogens, stomata remain partially closed, chlorophyll is reduced,
and photosynthesis stops even the plant eventually wilts. In advanced stages of disease, the rate
of photosynthesis is no more than one-fourth the normal rate.

Effect of Pathogens on Translocation of Water & Nutrients in the Host Plant

All living plant cells require an abundance of water and an adequate amount of organic and
inorganic nutrients in order to live & to carry out their physiological functions. When a pathogen
interferes with the upward movement of inorganic nutrients & water or with the downward
movement of organic substances, diseased conditions result in the parts of the plant denied these
materials. The diseased parts, in turn, will be unable to carry out their own functions & will deny
the rest of the plant their services or their products, thus causing disease of the entire plant. For
example, if water movement to the leaves is inhibited, the leaves can’t function properly,
photosynthesis is reduced or stopped, and few or no nutrients are available to move to the roots,
which in turn become starved & diseased and may die.

Interference with Upward Translocation of Water & Inorganic Nutrients

Many plant pathogens interfere in one or more ways with the translocation of water and
inorganic nutrients through plants. Some pathogens affect the integrity/function of the roots,

39
causing them to absorb less water; other pathogens, by growing in the xylem vessels, interfere
with the translocation of water through the stem.

Effect on Absorption of Water by Roots

Many pathogens (damping-off fungi, root-rotting fungi & bacteria, most nematodes, and some
viruses), cause an extensive destruction of the roots before any symptoms appears on the
aboveground parts of the plant. Root injury affects the amount of functioning roots directly and
decreases proportionately the amount of water absorbed by the roots.

Effect on Translocation of Water through the Xylem

Fungal and bacterial pathogens that cause damping off, stem rots and cankers may reach the
xylem vessels in the area of the infection may cause their destruction and collapse. In many
cases, affected vessels may be filled with the bodies of the pathogen and with substances
secreted by the pathogen or by the host in response to the pathogen & may become clogged.
Whether destroyed or clogged, the affected vessels cease to function properly and allow little or
no water to pass through them.

Effect on Transpiration
In plant diseases in which the pathogen infects the leaves, transpiration is usually increased due
to destruction of the cuticle, an increase in the permeability of leaf cells, and the disfunction of
stomata. If water absorption & translocation cannot keep up with the excessive loss of water, loss
of turgor and wilting of leaves follow. Obligate fungal parasites, such as rust & mildew fungi,
cause an accumulation of photosynthetic products, as well as inorganic nutrients, in the areas
invaded by the pathogen. In these diseases, the infected areas are characterized by reduced
photosynthesis and increased respiration. In some virus diseases, particularly the leaf-curling
type and some yellows diseases, starch accumulation in the leaves is mainly the result of
degeneration (necrosis) of the phloem of infected plants, which is one of the first symptoms.

Effect of Pathogens on Plant Growth

It is easily understood and expected that pathogens that destroy part of the photosynthetic area of
plants and cause significantly reduced photosynthetic output often result in smaller growth of

40
these plants and smaller yields. Similarly, pathogens that destroy part of the roots of a plant or
clog their xylem or phloem elements, thereby severely interfering with the translocation of water
and of inorganic or organic nutrients in these plants, often cause a reduction in size and yields by
these plants and, sometimes, their death.

Effect of Pathogens on Plant Reproduction

Pathogens that attack various organs and tissues of plants weaken and often kill these organs or
tissues, thereby weakening the plants. In several plant diseases, especially in grain crops, the
pathogen interferes directly with the reproduction of the plant by killing the embryo, that would
have produced the seed, and replacing the contents of the seed with its own fruiting structure or
its own spores. Examples of such diseases are ergot of grains, caused by the fungus Claviceps
purpurea; corn smut; and the covered & loose smuts of the various cereals.

5.4. Diagnosis of Plant Diseases

Diagnose: determine the nature of a disease. For diagnosis of plant disease it is prudent to first
determine whether the disease is caused by a pathogen (biotic) or an environmental (abiotic)
factor.
To diagnose a plant disease successfully you need to be familiar with:
• The basic classification of plants & how the healthy plant looks like to be able to
recognize when it is diseased.
• The characteristics of the organisms that cause disease (fungi, bacteria, etc.,).
• The symptoms & signs associated with the major types of diseases. Plant disease
diagnosis is designed to recognize the primary disease-causing agents.

Sample collection:
Obtain good samples of diseased plants for effective diagnosis. It is difficult, and often
impossible to diagnose a plant disease with a poor sample. A plant in a plastic bag for a week
may be so deteriorated and overgrown with microorganisms are useless. Sample that doesn’t
show all the symptoms may also be useless. Always attempt to collect fresh samples in various
stages of disease development & entire plants including the roots & the soil adhering to the roots
should be taken to the laboratory. Collecting your own samples is usually better than having

41
some one else collect them. It is often helpful to collect several healthy plants for comparison
during your examination. Plant samples should be protected from deterioration due to excessive
heat, cold, drying, or moisture. Plants should be maintained in a moist environment to prevent
drying of tissues, but excess moisture should be avoided. If this can’t be done, press leaves,
stems and roots between cardboard using a plant press.

Isolation: The separation of the pathogen from its host & its culture on a nutrient medium.
Isolation and identification of the pathogen is necessary because of the following reasons.
• There are many fungal & bacterial diseases that have similar symptoms & can’t be
distinguished visually from one another.
• In many cases, the pathogen can’t be identified because it is mixed with one or more
contaminants.
• The same disease could be caused by more than one similar looking pathogen and
perhaps by some environmental factor.
• Occasionally a disease is caused by a new, previously unknown pathogen that must be
isolated & studied.

Microscopic observation
A microscopic examination (using wet mount) is necessary for identification of many diseases.
Prepare microscopic mounts by suitable method such as by cutting thin tissue sections with a
razor blade or by making preparation of fungal structures, either removed from the tissue with a
scalpel or needle. Look for fungal mycelium, spores, & spore producing structures, and for
masses of bacterial cells streaming from tissue when placed in a drop of water. Remember that
diseased tissues are usually rapidly colonized by secondary MOs which can confuse the
diagnosis. This is one reason why fresh sample are necessary. Microscopes are obviously very
useful in disease diagnosis but aren’t always necessary. Don’t forget to examine below ground
parts of the plants, because many disease symptoms expressed above ground are due to a
problem in the roots or base of the stem.
Comparative symptomatology of infectious and noninfectious diseases (as a summary)

No Noninfectious Diseases (Disorder) Infectious Diseases

42
 1 All maize plants similarly and evenly
yellowed over an area of saline soil.
2 Symptoms are relatively simple possibly
limited to one or only primary.
3 Lesion borders are sharply non-expanding.
4 Symptoms on individual plants are highly
uniform in nature.
5 Usually no signs.
6 Relatively wide range of plants species
affected.
Plants in an affected area may vary in their
extent of disease development.
Symptoms tend to be complex; possibly
both primary and secondary symptoms.
Lesion borders are expanding.
Symptoms variable in size, pattern and
occurrence.
Signs may be present.
Only certain species affected.

6. HOST-PATHOGEN RELATIONS

6.1. Disease Development

Disease Triangle or Disease Pyramid

It is common to recognize three important components of disease, i.e., a susceptible host,
virulent pathogen, and a favorable environment. The interactions of the three components of
disease have often been visualized as a triangle, generally referred to as the “disease triangle”.
However, very recently, scientists have recognized a fourth factor (time), as an equally important

43
dimension of plant disease. Adding time to the disease-triangle as a fourth dimension gives a
disease-pyramid.

A disease pyramid showing the interrelationships of the different components of disease.

Reducing any component of disease pyramid through management (the host susceptibility, the
virulence of pathogen, degree of favorableness of time or of environment) will decrease the
amount of disease in a plant. Disease development in cultivated plants is also greatly influenced
by a fifth component, humans.

Humans affect disease development by affecting the kind of plants grown in a given area, their
level of resistance, the number of crops planted, time of planting, & density of plants.

6.2. Plant Disease Cycle

Disease cycle: The chain of events involved in disease development, including the stages of
development of the pathogen and the effect of the disease on the host. The main events of a

44
disease cycle are inoculation, penetration, establishment of infection, colonization (invasion),
growth and reproduction of the pathogen, dissemination of the pathogen, and survival of the
pathogen in the absence of the host (over-seasoning) of the pathogen.

6.2.1. Inoculum and inoculation

Inoculation is the initial contact of a pathogen with a site of plant where infection is possible.
The pathogen that lands on or is otherwise brought into contact with the plant is called the
inoculum. The inoculum is any part of the pathogen that can initiate infection. Thus, in fungi
the inoculum may be spores, sclerotia or fragments of mycelium.
In bacteria, mollicutes, protozoa, viruses, & viroids, the inoculum is always whole individuals. In
nematodes, the inoculum may be adult, juveniles, or eggs. In parasitic higher plants, the
inoculum may be plant fragments or seeds. One unit of inoculum of any pathogen is called a
propagule.

6.2.2. Pre-penetration/Penetration process

Pre-penetration
This phase includes all the events prior to actual entry of the pathogen such as germination of
spores & seeds, hatching of eggs (nematodes), attachment of pathogen to host, and recognition
between host and pathogen. Almost all fungi, bacteria, and parasitic higher plants are first
brought into contact with the external surface of plant organs.

Penetration process
This is the actual entry of the pathogens into their host plants. Pathogens penetrate plant surfaces
by direct penetration of cell walls, through natural openings, or through wounds. Some fungi
penetrate tissues in only one of these ways, others in more than one. Bacteria enter plants mostly
through wounds, less frequently through natural openings, and never directly through unbroken
cell walls. Viruses, viroids, mollicutes, fastidious bacteria, and protozoa enter through wounds
made by vectors, although some viruses and viroids may also enter through wounds made by
tools and other means. Parasitic higher plants enter their hosts by direct penetration. Nematodes
enter plants by direct penetration and, sometimes, through natural openings. Penetration does not

45
always lead to infection. Many organisms actually penetrate cells of plants that are not
susceptible to these organisms & that do not become diseased. Direct penetration through intact
plant surfaces is probably the most common type of penetration by fungi (Oomycetes) and
nematodes & the only type of penetration by parasitic higher plants. None of the other pathogens
can enter plants by direct penetration.

6.2.3. Infection/Invasion and establishment

Infection is the process by which a pathogen establishes contact with host cells or tissues &
procures nutrients from them. During infection pathogens grow or multiply or both, within the
plant tissues & invade & colonize the plant. Successful infection results in the appearance of
symptoms on the host plant. For a successful infection to occur, the contact of the pathogen with
the host plant, favorable environmental factors and the inoculum potential (capable of infecting
the host) should be recognized in the infection process.
Invasion refers to the spread of the pathogen in the host.
Incubation period is the time interval b/n inoculation and appearance of disease symptoms.

6.2.4. Reproduction
Plant pathogens reproduce in different ways. Fungi reproduce by means of spores; parasitic
higher plants by seeds; bacteria, mollicutes & protozoa by fission; viruses & viroids inside host
cells as virions; & nematodes by eggs. The rate of reproduction varies considerably among the
various kinds of pathogens. A few pathogens can produce tremendous numbers of individuals
within one growing season (polycyclic).
Polycyclic: Completes many (life or disease) cycles in one year (e.g., rust, chocolate spot, late
blight, etc.).

Monocyclic: Having one cycle per year (e.g., smut)

Under optimum nutritional and environmental conditions (in culture), bacteria divide (double
their numbers) every 20 to 30 minutes. Presumably, bacteria multiply just as fast in a susceptible
plant as long as nutrients & space are available and the temperature is favorable. Virus particles
accumulate within the infected living cell until as many as 100,000 to 10,000,000 particles may
be present in a single cell. Viruses & viroids infect & multiply in most or all living cells of their

46
hosts. Nematode females’ lay about 300 to 500 eggs, about half of which produce females that
again lay 300 to 600 eggs each.

6.2.5. Dissemination
Almost all dissemination of pathogens is carried out passively by such agents as air and insects.
To a lesser extent, water, certain other animals, and humans may be involved. The spores of
many fungi are successfully disseminated through the air for only a few hundred or a few
thousand meters. The spores of other fungi, however, are often carried over distances of several
kilometers, even hundreds of kilometers (the cereal rusts), and in favorable weather may cause
widespread epidemics. Some fungi can spread into new areas quite rapidly and may cause severe
epidemics over large areas, including entire continents, within a few years. Bacteria &
nematodes present in the soil may be blown away along with plant debris or soil particles in the
dust. Wind also helps in the dissemination of bacteria, fungal spores, and nematodes by blowing
away rain splash droplets containing these pathogens. Wind causes adjacent plants or plant parts
to rub against one another, which may help the spread by contact of bacteria, fungi, some viruses
and viroids, and possibly some nematodes.

Although water is less important than air in the long-distance transport of pathogens, the water
dissemination of pathogens is more efficient for nearby infections, as the pathogens land on an
already wet surface & can move or germinate immediately. Insects, particularly aphids,
leafhoppers, and whiteflies, are by far the most important vectors of viruses. Specific insects
also transmit certain fungal, bacteria, and nematode pathogens. Almost all animals that move
among plants and touch the plants along the way can disseminate pathogens such as fungal
spores, bacteria, seeds of parasitic plants, nematodes, and perhaps some viruses and viroids.
Most of these pathogens adhere to the feet or the body of the animals, but some may be carried in
contaminated mouthparts.

Humans disseminate all kinds of pathogens over short & long distances in a variety of ways:
• Within a field through the successive handling of diseased & healthy plants;
• By transporting contaminated soil on their feet or equipment, & using infected
transplants, seed, nursery stock, & bud-wood;

47
 • By importing new varieties or food items into an area that may carry harmful pathogens.

6.2.6. Survival of pathogens

In the absence of their hosts, the pathogens under go over-seasoning, & they have evolved great
variety of means of survival. On perennial plants fungi survive as dormant mycelium in infected
tissues. Fungi pathogens infecting aerial parts of deciduous plants usually survive as mycelia or
spores on fallen, infected leaves or fruits or on the bud scales. Fungi affecting annual plants
usually survive the winter or summer as mycelium in infected plant debris, as resting spores & as
sclerotia (hard masses of mycelium) in infected plant debris or in the soil, & in or on seeds &
other propagative organs, such as tubers. Some rust fungi infect alternately an annual & a
perennial host, & the fungus goes from one to the other host & over-winters in the perennial
host.

Bacteria overwinter & oversummer in essentially the same ways as described for fungi, i.e., in
infected plants, seeds, & tubers, in infected plant debris, &, for some, in the soil. Viruses,
viroids, mollicutes, fastidious bacteria, & protozoa survive only in living plant tissues such as the
tops & roots of perennial plants. Nematodes usually overwinter or oversummer as eggs in the
soil & as eggs or nematodes in plant roots or in plant debris. Parasitic higher plants survive
either as seeds, usually in the soil, or as their infective vegetative form on their host.

7. DEFENSE MECHANISMS OF PLANTS AGAINST PATHOGENS

7.1. Mechanisms of Attack by Pathogens

Pathogens attack plants because during their evolutionary development they have acquired the
ability to live off the substances manufactured by the host plants.

How pathogens attack plants?

48
 • Destruction of food reserves
• Prevention of seedling metabolism/respiration
• Interference with procurement of nutrients/absorption
• ,, ,, upward movement/transport of both water & nutrients (transpiration)
• Interference with photosynthesis
• ,, ,, translocation (food transfer)
• Interference with reproduction

Many substances are contained in the protoplast of the plant cells. If pathogens are to gain access
to these substances, they must first penetrate the outer barriers formed by the cuticle and/or cell
walls. Penetration and invasion seem to be aided by mechanical force exerted by certain
pathogens on the cell walls of the plant.

DEFENSE MECHANISMS:
• Structural defense mechanisms (physical barriers)
• Biochemical reactions that produce toxic substances
• Combination of both mechanisms

7.2. Structural Defense Mechanisms

7.2.1. Pre-existing defense structures (passive defense)

• The amount & quality of wax & cuticle that cover the epidermal cells (waxes on leaf &
fruit surfaces form a water-repellent)
• Cuticle thickness may increase resistance to infection
• Thickness and toughness of the outer wall of epidermal cells
• Location, size & shape of stomata & lenticels & time of stomata opening.

7.2.2. Post-infection defense structures (active defense)

This includes:
• Histological defense structures (via formation of tissues ahead of the pathogen)
• Cellular defense structures (involves morphological changes in the cell wall/derived from
the cell wall)

49
 • Cytoplasmic defense reaction
• Necrotic defense (defense through hypersensitivity)

HISTOLOGICAL DEFENSE STRUCTURES

Formation of Cork Layers: formation of several layers of cork cells following infection by
fungi or bacteria & even by some viruses & nematodes.
Formation of Abscission Layers: formed on young, active leaves of stone fruit trees following
infection by several fungi, bacteria, or viruses. E.g., Mango, Avocado, Peach, etc.
Formation of Tyloses: Tyloses form in xylem vessels of most plants under various conditions of
stress & during invasion by most of the vascular pathogens.

Schematic formation of Abscission layer & Tyloses

• Deposition of Gums: Gums produced & deposited in two places intercellular (outside the
cell) spaces deposited in the cell (intra-cellularly)

CELLULAR DEFENSE STRUCTURES

• Swelling of the outer layer of the cell walls of parenchyma cells.
• Thickening of the cell wall

CYTOPLASMIC DEFENSE REACTION

• The cytoplasm moves toward the enemy/pathogen
• The cytoplasm becomes granular & dense
50
NECROTIC DEFENSE
• The nucleus moves toward the pathogen & soon disintegrates, & brown, resin-like
granules form in the cytoplasm.
• The hyphae don’t grow & further invasion is stopped.

7.3. Biochemical Defense Mechanisms

The resistance of a plant against pathogen attacks depends not so much on its structural barriers
as on the substances produced in its cells preceding or following infection.

7.3.1. Pre-existing biochemical defense (passive mechanism)

• Inhibitors (fungi-toxic substances/exudates) released by the plant in its environment
e.g., Onion with white scale – susceptible to onion smudges.
Onion ,, red scale – resistance to onion smudges.
• Inhibitors present in the plant cells before infection

Several phenolic compounds are present in the plant cells inhibit infection but when the plant
gets older they will be reduced.
• Lack of essential factors:
• Lack of host receptors and sensitive sites for toxins.
• Lack of low concentration of essential nutrients for the pathogen
• Plants lacking growth regulators/essential substances
• Lack of recognition b/n the host & the pathogen - plants may lack recognition
factors, i.e., specific molecules or structures. e.g., Oligosaccharides,
polysaccharides, proteins, glycoproteins.

7.3.2. Induced biochemical mechanisms/active mechanisms

• Biochemical substances (inhibitors) produced in plants due to infection by – pathogens,
chemical treatment agents & physical treatments. e.g., Fungitoxic substances produced
around the site of injury like chlorogenic and phytoalexins.

51
Phytoalexins: toxic to fungi and bacteria and produced upon infection (during infection).
Phytoalexins are produced only by resistance varieties. e.g., pisatin production by pea pods
(Ascochyta pisi), gossypol in cotton, capsidol in pepper, etc.
Elicitors: produced by pathogens stimulate the production of phytoalexins in host plants.

Phenol-oxidizing enzymes in disease resistance

Their activity is generally higher in infected resistant varieties than in the infected susceptible or
uninfected healthy plants.
Example: Quinones are more toxic to pathogens than the original phenols.

Induced synthesis of proteins and enzymes

Pathogen attacks appear to induce protein or enzyme synthesis in the plant that can lead to the
development of a local resistant or immune layer around infection sites.

Formation of substances resisting the enzymes of the pathogen

Pectins, proteins and polyvalent such as Ca, Mg & Ca produced by resistant plants inhibit tissue
maceration and confine the pathogen to lesions of limited size.

Inactivation of pathogen enzymes

Polyphenols inhibit the action of pathogenic enzymes rather than the pathogen itself and
contribute to the resistance of the plant.
Detoxification of pathogen toxins
Detoxification of some toxins, e.g., fusaric acid, pyricularin, etc., is known to occur in plants and
to play a role in disease resistance. E.g., Helminthosporium & Alternaria.

Hypersensitive reaction (HR)

Following infection, loss of turgor, browning and death of infected cells occur rapidly in resistant
varieties. Fungal and bacterial pathogens within the area of operation of the hypersensitive
reaction are isolated by necrotic tissue and quickly die. In virus diseases, the HR always results
in formation of the so-called local lesions in which the virus may survive for considerable time
but is, generally, found in low concentrations and its spread beyond the lesion is checked.

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 8. GENETICS OF PLANT DISEASES

8.1. Variability of Pathogens and Mechanisms

All individuals produced as a result of a sexual process are expected to be different from one
another and from their parents in a number of characteristics, although they retain most
similarities with them and belong to the same species. When individuals are produced asexually,
the frequency and degree of variability among the progeny are reduced greatly, but even then
certain individuals among the progeny will show different characteristics.
Sexual reproduction
Fungi are sexually reproduced through Oospores, Ascospores, Basidiospores & Zygospores.
Nematodes are reproduced through eggs and parasitic higher plants through seeds.
Asexual reproduction
Fungi reproduce through Conidia, Zoospores, and Sclerotia. Bacteria divide by binary fission
MLOs divide by binary fission
Mechanisms of Variability
It is categorized into two:
• General mechanisms: work for all organisms
• Specialized mechanisms: work for specific organisms

8.1.1. General mechanisms

A. Mutation: works for all organisms
It is a more or less abrupt change in the genetic material of an organism, which is then
transmitted in a hereditary fashion to the progeny. Mutations represent changes in the sequence
of bases (A=Adenine, G=Guanine, C=Cytocine, T=Thymine) in the DNA either through:
• Substitution of one base for another
• Addition or deletion of one or many base pairs
• Amplification of particular segments of DNA to multiple copies
• Insertion of a movable DNA segment into a coding gene or

53
 • Inversion of the DNA segments

Mutations occur spontaneously in nature in all living organisms: those that produce only sexually
or only asexually and those that reproduce both sexually and asexually. Mutations in single-
celled organisms (bacteria, in fungi with a haploid mycelium, and in viruses), may be expressed
immediately after their occurrence, i. e., faster in lower organisms than higher organisms. Most
mutations, however, are usually recessive; therefore, mutations can remain unexpressed until
they are brought together in a homozygous condition. Mutations occur intra-nucleus (in the
nucleus) and inter-nucleus (outside the nucleus).
Causes of mutation:
• Radiation (ultra violet rays, x-rays)
• High temperature
• Chemical treatment
Factors favoring/facilitating mutations:
The production of great number of progenicies by pathogens due to monocultural cropping
system and planting a few genetically homogeneous varieties of each crop continuously over
enormous land expenses (for many years).
B. Recombination
Recombination occurs primarily during the sexual reproduction of plants, fungi, and nematodes
whenever two haploid (1N) nuclei, containing genetic material unite to form a diploid (2N)
nucleus, called a zygote.
1N + 1N (Haploid) = 2N (Diploid or Zygote)
The zygote, sooner or later, divides meiotically and produces new haploid cells (gametes, spores,
and mycelium).
Variability: The property or ability of an organism to change its characteristics from one
generation to the other.

8.1.2. Specialized Mechanisms of Variability in Pathogens

Sexual-like process in fungi
A. Heterokaryosis

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Anastomosis/overlapping of fungal hyphae or parts of hyphae containing two or more nuclei
that is genetically different.
Dikaryotic state – when two gametes present in the same cytoplasm without mating is called
dikaryotic. In the Basidiomycetes, the dikaryotic state may differ drastically from the haploid
mycelium and spores of the fungus. Example, wheat stem rust fungus Puccinia graminis f. sp.
tritici, which in the haploid state infects barberry while in the diploid state it infects wheat. In
general, pathogens undergoing any type of recombination pose a greater threat than pathogens
that undergo little or no recombination.
B. Parasexualism
The process by which genetic recombination comes about by the occasional fusion of the two
nuclei and formation of a diploid nucleus.
C. Heteroploidy
The existence of one, two, three, or more extra chromosomes or missing one or more
chromosomes from the normal. In several studies, spores of the same fungus were found to
contain nuclei with chromosome numbers ranging from 2 to 12 per nucleus and also diploids and
polyploidy. Heteroploidy has been observed repeatedly in fungi and has been shown to affect the
growth rate, spore size and rate of spore production, hyphal color, enzyme activities, and
pathogenicity.

Sexual-like Processes in Bacteria

• Conjugation: occurs when two compatible bacteria come in contact with one another and
a small portion of the chromosome or plasmid (non-chromosome) from one bacterium is
transferred to the other.
• Transformation: bacterial cells are transformed genetically by absorbing and
incorporating in their own cells genetic material secreted or released by other compatible
bacteria.
• Transduction: a bacterial virus (phage) transfers genetic material from the bacterium in
which the phage was produced to the bacterium it infects next.

Genetic Recombination in Viruses

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When two strains of the same virus are inoculated into the same host plant, one or more new
virus strains are recovered that are known as recombinants or new mutants. Their properties
(virulence, symptomatology, and so on) are different from the original strains introduced into the
host.

8.1.3. Stages of variation in pathogens

The entire population of a particular organism on the earth, e.g., a fungal pathogen, has certain
morphological & other phenotypic characteristics in common and makes up the species of
pathogen.
Species: The entire population of a particular pathogen with certain morphological
characteristics in common. e.g., Puccinia graminis
Varieties/special forms: Some individuals or groups of the species that attack only certain plant
spp. (such as only wheat, barley, or oats, etc.).
e.g., P. graminis f. sp. tritici (formae specialis)
P. graminis f. sp. hordei, and
P. graminis f. sp. avenae.

Physiological race: group of individuals within each formae specialis that attack some of the
crop varieties but not others, while some others attack another set of varieties. E.g., there are
more than 200 races of P. graminis f. sp. tritici (race 1, race 15, race 59 and so on).
Differential varieties: are used to distinguish physiological races that exist in a locality or
country.
Variant: one of the offspring of a race (physiological race) that can suddenly attack a new
variety or can cause severe symptoms on a variety that it could barely infect before
Biotype: The identical individuals produced asexually by the variant. Each race consists of one
or several biotypes. E.g., (race 15A, 15B, and so on).

8.2. Genetics of Virulence and Resistance

Infectious plant diseases are the result of the interaction of at least two organisms, the host plant
and the pathogen. The properties of each of these two organisms are governed by their genetic
material, the DNA, which is organized in numerous segments making up the genes. DNA is

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usually present in or on the chromosome. Genetic information is encoded in the DNAs. In
viruses and viroids genetic information is encoded in the RNA. In Eukaryotes (and all other
organisms), there are several chromosomes in the nucleus. Similarly, all cells of Eukaryotes
carry their DNAs in their mitochondria. Plant cells, in addition to nuclear and mitochondrial
DNA, also carry DNA in their chloroplasts. In Prokaryotes (bacteria and mycoplasmas) there is
only one chromosome in the cytoplasm. Many prokaryotes and some of the lower eukaryotes
carry smaller circular molecules of DNA called plasmids in the cytoplasm. Plasmid DNA also
carries genetic information but multiplies and moves independently of the chromosomal DNA.

The structure of the DNA (deoxyribose nucleic acid)

a) Introns: non-coding stretch of DNA.
b) Promoters: act as signals for production of DNA and proteins.
c) Terminators: act as receptors and terminators of gene expression.
Genetic information in DNA is encoded in a linear fashion in the order of the four bases (A,
adenine; C, cytosine; G, guanine; and T, thymine). Each triplet of adjacent bases codes for a
particular amino acid. Attachment sequence: A-T, G-C, C-G, and T-A.
Gene: A linear portion of the chromosome that determines or conditions one or more hereditary
characters; the smallest functioning unit of the genetic material. In many cases of host–pathogen
interaction, genes in the one organism are triggered to be expressed by a substance produced by
the other organism. For example, genes for cell wall-degrading enzymes in the pathogen are
apparently induced by the presence of monomers or oligomers of host cell wall macromolecules
that are substrates for these enzymes. Also, genes for defense reactions in the host, e.g., the
production of phytoalexins, apparently are triggered to expression by certain signal compounds
activated by inducer molecules (elicitors) produced by the pathogen.

The gene(s) for virulence in a pathogen is usually specific for one or a few related kinds of plants
that are hosts to the pathogen. Also, the genes and gene combinations that make a plant
susceptible, i.e., a host to a particular pathogen, are present only in that one kind of plant and
possibly a few related kinds of plants.
Why are all the plants not attacked by their pathogens, and why are those that are attacked not
usually killed by the pathogens? The answer is complex, but basically it happens because plants,

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through evolution or through systematic breeding, have acquired, in addition to the genes that
make them susceptible to a pathogen, one or usually numerous genes for resistance that protect
the plants from infection or from severe disease.

With numerous host–pathogen combinations, that, after a new gene for resistance to a pathogen
is introduced into a crop variety and that variety is planted in the fields, a new population (race)
of the pathogen appears that contains a new gene for virulence that enables the pathogen to
attack the crop plants containing the new gene for resistance.
How did this new population of pathogens acquire the new gene for virulence? New genes can
arise randomly and suddenly through mutations, or by rearrangement of the genetic material of
the pathogens through the ever-ongoing events of genetic variability in organisms.
The concurrent occurrence and interaction of specific genes for virulence in the pathogen and of
specific genes for susceptibility in the host determine the initiation and development of disease.
Avirulence: Lacking virulence, lose of capacity to attack a host.

8.3. The Gene-for-Gene Concept

The coexistence of host plants and their pathogens side by side in nature indicates that the two
have been evolving together. “For each gene that confers virulence to the pathogen there is a
corresponding gene in the host that confers resistance to the host and vice versa”. The gene-for-
gene concept was first proved in the case of flax and flax rust, but it has been shown to operate in
many other fungi diseases and in several diseases caused by bacteria, viruses, parasitic higher
plants, nematodes, and even insects.

Quadratic check of gene combinations and disease reaction types in a host–pathogen

system in which the gene-for -gene concept for one gene operatesa.
Virulence or Avirulence Resistance or susceptibility genes in the plant
Virulence or avirulence R (resistant) r (susceptible)
genes in the pathogen dominant recessive

A (avirulent) dominant AR (-) Ar (+)

a (virulent) recessive aR (+) ar (+)
a
Minus signs indicate incompatible (resistant) reactions and therefore no infection. Plus signs
indicate compatible (susceptible) reactions and therefore infection develops.

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A=Avirulence & a=Virulence for pathogen R=Resistance & r=Susceptible for host
Gene combination: A1A2, A1a2, a1A2, a1a2 R1R2, R1r2, r1R2, r1r2
Genes of pathogen Genes of host Disease reaction
A1A2 ---------------------------R1R2------------------------Resistant
A1A2----------------------------R1r2-------------------------Resistant
A1A2----------------------------r1R2-------------------------Resistant
A1A2----------------------------r1r2--------------------------Susceptible
A1a2----------------------------R1R2-------------------------Resistant
A1a2----------------------------R1r2--------------------------Resistant
A1a2----------------------------r1R2--------------------------Susceptible
A1a2----------------------------r1r2---------------------------Susceptible
a1A2----------------------------R1R2-------------------------Resistant
a1A2----------------------------R1r2--------------------------Susceptible
a1A2----------------------------r1R2--------------------------Resistant
a1A2----------------------------r1r2---------------------------Susceptible
a1a2----------------------------R1R2--------------------------Susceptible
a1a2----------------------------R1r2---------------------------Susceptible
a1a2----------------------------r1R2---------------------------Susceptible
a1a2----------------------------r1r2----------------------------Susceptible

Generally, R1-A1- and -A2-R2 = Resistance

8.4. Types of Plant Resistance to Pathogens

Whatever the kind of defense or resistance a host plant employs against a pathogen or an abiotic
agent, it is ultimately controlled, directly or indirectly, by the genetic material (genes) of the host
plant and of the pathogen.
Plants are resistant because:
• They are immune to the pathogen (non-host resistance)
• They possess genes for resistance (R genes) directed against the avirulence genes of the
pathogen (true resistance).

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 • The plants, due to various reasons escape or tolerate infection by pathogen (apparent
resistance).
True resistance: located in the nucleus. It is genetically controlled by the presence of one, a few
or many genes for resistance in the host plant.

Reasons for true types of resistance:

a) Lack of recognition between the pathogen and the host plant.
b) The presence of resistance genes.
There are two kinds of true resistance: horizontal and vertical.

Horizontal resistance
All host plants possess some level of horizontal resistance. Such resistance also called non-
specific, general, quantitative adult plant, field, or durable resistance. Each of these genes
alone may be rather ineffective against the pathogen and may play a minor role in the total gene
resistance (minor gene resistance). Generally, horizontal resistance doesn’t effectively protect
plants against infection. However, it may slow down:
• the development of individual infection point on a plant
• the spread of the disease & development of epidemics in the field.

Vertical resistance
• Differentiates clearly between races of the pathogen since it is effective against specific
races of the pathogen.
• Sometimes called as strong, major, specific, qualitative, complete, stable or differential
resistance, but is most commonly referred to as vertical resistance.
• Controlled by one or a few genes (monogenic or oligogenic resistance).
• The host and the pathogen are incompatible.
• Inhibits epidemic development by limiting the reproduction of initial inoculum.

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Often, it is desirable to combine more than one resistance genes (R 1R2, R1R3, R1R2R3) in the same
plant, which is then resistant to all races of the pathogen. A plant species may have as many as
20 to 40 genes for resistance against a particular pathogen (e.g., wheat against the leaf rust
pathogen - Puccinia recondita). In some plants, genetic material for resistance against the
disease is present in the cytoplasm instead of nucleus, of cell. The two best known examples of
such cytoplasm resistance in corn are the southern corn leaf blight and yellow leaf blight caused
by Bipolaris maidis and Phyllostica maidis, respectively.

Apparent Resistance
This type of resistance occurs as a result of disease escape or tolerance to disease.

Disease escape: occurs when genetically susceptible plants don’t become infected because, three
factors necessary for disease, i.e., susceptible host, virulent pathogen, and favorable environment
don’t coincide and interact at the proper time. Plants may escape disease because:
• Their seeds germinate faster
• ,, seedlings harden before the pathogen attack
• Some plants susceptible to the pathogen only at a particular growth stage (young leaves,
stems or fruits). E.g., young tissues and plants are infected much more severely by
pythium, powdery mildews, and most bacteria and viruses, than are older ones.

Disease Tolerance: the ability of plants to produce a good crop even when they are infected with
a pathogen.
Tolerance results from:
• Lack of receptor sites for the pathogen
• Inactivating the irritant excretion of pathogen
• Productions of efficient branch, tiller, or root to compensate the affected part of the plant.

Sources of Disease Resistance

• Adapted cultivars: particularly useful to improve levels of quantitative resistance
• Breeders’ stock: lines that are relatively well adapted, but haven’t been released as
cultivars.

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 • Cultivated Germplasm: maintained at plant introduction stations & other collections of
easy accessible genetic diversity.
• Wild or related species: much more difficult to incorporate resistance from wild species
than from Germplasm of the same species.
• Artificially induced variation within a plant species: this serves when genetic variation
for resistance doesn’t exist within a species.

Varieties with monogenic or oligogenic resistance generally show complete/stable resistance to a

specific pathogen under most environments. However, varieties with general (polygenic)
resistance are less stable under different environments. As a rule, a combination of major and
minor genes through cross breeding for resistance against a pathogen is the most desirable
makeup for any plant variety. Multiline cultivars or cultivar mixtures also have been promoted as
a means to achieve durable resistance to highly variable pathogens such as the cereal rusts.

Breakdown of resistance
• When genetically uniform plants are grown over large areas, a new pathogen race will
appear that can attack the previously resistance plants (e.g., southern corn leaf blight by
Helminthosporium spp.)
• Occasionally by mutation.

9. EPIDEMIOLOGY OF PLANT DISEASES

The Elements of an Epidemic: timely combination of susceptible host plants, a virulent

pathogen, and favorable environmental conditions over a relatively long period of time.
Epidemiology: The study of epidemics and of the factors that influence them. Plant disease
epidemics sometimes also called epiphytotics; occur on most crops in many parts of the world.

9.1. Different Factors Affecting Epiphytotics

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9.1.1. Host factors
• Levels of genetic resistance or susceptibility: host plants with high levels of (vertical)
resistance do not allow a pathogen to become established, unless a new race of pathogen
develops. Plants with lower levels of (horizontal) resistance become infected that depends
on the level of resistance and the environmental conditions.
• Degree of genetic uniformity of plants: when genetically uniform host plants are grown
over large area, there is much possibility of the appearance of a new race of pathogen to
attack them leading to an epidemic.
• Type of crop and age of plants: in annual crops like corn, wheat, vegetables, cotton,
tobacco, etc. epidemics is generally develop much more rapidly (in few weeks) than in
the perennial woody fruit and forest trees.

9.1.2. Pathogen factors

Virulence level: virulent pathogens infect host rapidly and produce large amounts of inoculum at
a faster rate.
Inoculum density: the greater the number of pathogen propagules (bacterial cells, fungal spores
& sclerotia, nematode eggs, virus-infected plants, etc.) within or near the fields of host plants, the
more inoculum reaches the fields and greatly increasing the chances of an epidemic.
Reproduction pattern of pathogen: pathogens, such as most fungi, bacteria and viruses, have
short reproduction cycles and thus can produce many generations in a single growing season.
Ecology of pathogen: in most fungi and parasitic phanerogams, inoculum is produced on the
surfaces of aerial parts of plant. This can be dispersed easily to longer distances and can cause
widespread epidemics.
Mode of spread of pathogen: inocula of rusts, mildews and leaf spot fungi are spores that can
be easily dispersed by air and strong winds over long distance. They cause most frequent and
most widespread epidemics. Seed and soil borne pathogens suffer from several limitations and
hardly cause local, slow-spreading diseases.

9.1.3. Environmental factors

The concurrent presence of susceptible plants and virulent pathogens doesn’t always guarantee
numerous infections and the development of an epidemic in the absence of favorable

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environmental factors. The environment may affect each component of the epidemic, i.e. host as
well as pathogen. In addition, the environment may affect:
• Growth stage, succulence, and genetic susceptibility of the host
• Survival, vigor, multiplication rate, sporulation, direction and distance of dispersal of the
pathogen
• The rate of spore germination and penetration
• The number and activity of the vectors of the pathogen
The most important environmental factors that affect disease epidemics are moisture and
temperature.

9.1.4. Human factors

Many activities of humans, chiefly the cultural practices and the control measures used, have
direct or indirect effect on plant disease epidemics. These may favor or reduce the rate and
frequency of epidemics.
• Site selection: low-lying and poorly drained fields, especially nearest to other infected
fields, tend to favor the appearance and development of epidemics.
• Selection of propagative material: use of infected seed, nursery stock and other
propagative material. On the contrary, use of pathogen-free propagative material reduces
the chance of epidemic.
• Cultural practices: continuous monocropping (large areas planted with the same variety
of crop), high nitrogen fertilizer levels, dense planting (increasing foliage density),
overhead irrigation, injury by herbicide application, and poor sanitation etc. all increase
the possibility and severity of epidemics.
• Disease control measures: sometimes the control measures like use of certain chemicals
or planting of certain variety may lead to selection of virulent strains of the pathogen.
This results into serious outbreak of the disease either due to resistant of the pathogen to
chemical or the pathogen can attack the resistance of the variety.
• Introduction of new pathogen: the movement of seeds, tubers, nursery stock and other
propagative parts worldwide increased the chances of introduction of pathogens to new
virgin areas. Once to new areas, such pathogens lead to severe epidemics. For example,
Citrus canker (Xanthomonas citri).

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9.2. Measurement of Plant Diseases (pathometry)
Pathometry: the quantitative study of suffering plant. It involves disease measurement, yield
loss estimation, forecasting of plant disease and predicting their impact.
The objective of disease measurement is to take decisions on:
• Development of epidemics in time and space
• Analyzing the factors that affect disease development
• Evaluation of control measures (fungicides & host resistance)
• Prioritization of the allocation of time and money for various problems
• Growers judicious decisions for crop protection by using information on diseases, yield
loss and threshold levels.

Assessment of Plant Disease and Yield Loss

While measuring disease, one is interested in measuring:
• Disease Incidence: the number of plants or proportion of plant units (leaves,
stems, and fruit that show any symptoms) in relation to the total number of units
examined.

Disease Incidence (%) = No of diseased plants x 100

Total No of plants examined

The term disease prevalence also tells about occurrence of disease on large geographical area.
E.g., out of 10 fields assessed in a given area, 6 are observed to show disease symptom.
Therefore, the disease prevalence = 6/10 x 100 = 60%
• Disease Severity: the proportion of area or amount of plant tissue that is
diseased (expressed in %).

Disease Severity (DS) = Area of plant tissue infected x 100

Total area examined

Assessment keys
Methods for assessing severity can be conveniently separated into two types.

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(I) Descriptive keys: prepared by describing the characteristic symptoms of disease or
amount of disease along with description. Disease scoring scales could be 0-5, 0-9 or 1-9.
Field Keys for the Assessment of Potato Late Blight caused by Phytophthora infestans (1-9 scale
by British Mycology Society, 1945)).

Disease scale % infection Description of P. infestans

1 0 No disease on plant
2 0.1 Only few plants affected (1 or 2 spots) in 12 yards’ radius.
3 1 Up to 10 spots per plant, or general light spotting.
4 5 About 50 spots per plant or up to 1 leaflet in 10 attacked.
5 25 Nearly every leaflet with lesions, plants still remaining
normal; but looks green, although every plant is affected.
6 50 Every plant affected & about ½ of leaf area destroyed by
blight; field looks green flecked with brown.
7 75 About 3/4th of leaf area destroyed by blight; field looks
neither predominantly brown nor green.
8 95 Only a few leaves left green but stems green.
9 100 All leaves dead, stems dead or dying.

(II) Standard Area Diagrams: a set of pictures (diagrams, sketches, or photographs) having
different amount of disease that can help in measurement of disease under field conditions.
(c) Yield Loss: the proportion of the yield that the grower will not be able to harvest due to
disease (expressed in kg, qt, t).
Yield loss almost always results in economic loss from disease. Economic loss occurs whenever
economic returns from the crop decrease because of reduced yields.
The yield loss is the difference between attainable yield and actual yield.

Yield Loss = Attainable – Actual yield x 100

Attainable yield

Concepts of Crop Loss Assessment

• To obtain quantitative data on occurrence and development of disease

66
 • To assess the relative importance of different diseases
• To determine the relationship b/n disease severity and yield (quality) loss
• To calculate economic loss
• To distinguish b/n treatment; e.g., fungicides
• To detect differences in disease resistance among varieties

Yield data could be obtained from:

• Statement of authority
• Questionnaires
• Survey
• Arial photograph
• Field experiment – real data

In managing plant diseases, the grower can justify applying disease control measures only when
the incremental costs of control are generally smaller than the increase in crop returns. The level
of disease, i.e., the amount of plant damage, at which control costs just equal incremental crop
returns is called the economic threshold of the disease. The economic threshold of a crop–
pathogen system varies with the tolerance level of the crop, which depends on the growth stage
of the crop when attacked, crop management practices, environment, shifts in pathogen
virulence, & new control practices.

9.3. Simple and Compound Interest Plant Diseases

Simple Interest (monocyclic disease): when there is only one generation of disease in the crop
season. e.g.; root rot of peas, barley smut, etc.
Compound Interest (polycyclic disease): when there are several generations of the pathogen
within a life cycle of a crop. E.g., Late blight of potato and tomato, Chocolate spot, etc.

9.4. Monitoring and Forecasting of Plant Disease Epidemics

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Forecasting: saying in advance what is likely to happen. Therefore, Disease forecasting allows
the prediction of probable outbreaks or increases in intensity of disease. It is extremely useful to
farmers in the practical management of crop disease.

Monitoring: observing the field every time, frequent inspection of certain disease or pathogen.
Forecasting involves all the activities in ascertaining and notifying the farmers in a community.

Practical advantage of forecasting

• To determine when and where a particular management practice should be applied.
• Farmers always weigh the risks, costs and benefits of numerous decisions (e.g., relevant
cultural practices).
• Farmers can reduce the expenditure on chemicals, etc.

Basis of forecasting
• The amount of initial inoculum (e. g., greenhouse tests and direct assessment of target
fields (soil borne fungi structures such as sclerotia and nematode cysts).

• Weather conditions favoring development of secondary inoculum. This involves

assessing the rate of occurrence of the infection cycles. For example, late blight of potato
and tomato can be predicted when constant cool temperatures b/n 10 and 24 0C and
relative humidity over 75% for at least 48 hours coincide with several rainy days. This
result in infection that can lead to a late blight outbreak within 2-3 weeks later.
10. GENERAL CONTROL METHODS OF PLANT DISEASES

Except trees, damage or loss of individual plants is usually taken insignificant and therefore
control measures are generally aimed at saving the populations rather than a few individual
plants and measures used are preventive rather than curative.

10.1. Regulatory Methods (Quarantine, Sanitation & Exclusion of Plant Pathogens)

Quarantine: Control of import and export of plants, plant parts and their products to prevent
spread of diseases and pests. This measure is taken by state through legislation with the objective

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of preventing the entry of disease or pathogen into the country or into a region in the country.
Some of the worst plant disease epidemics that have occurred throughout the world include:
• Downy mildew of grape in Europe.
• The bacterial citrus canker, the chestnut blight, the Dutch elm disease and the soybean
cyst nematode in the USA were introduced from other countries.
Pathogens are likely to be introduced in or on live plants, seeds, tubers, and other planting
material, in imported grain, fruit, vegetables and other foodstuffs. The plants or their products
must be imported only from disease-free areas. There should be effective examination of
incoming plant material by adequately trained and experienced inspectors at ports, railway
stations and frontier ports.

Sanitation: Destruction of diseased plant material whether in the form of plant parts in the field
or crop residues after harvest. It is one of the effective recommendations in the management of
viral diseases of field crops, loose smut of wheat and production of disease-free seed. Rouging of
infected plants is recommended in several other diseases such as smut of sugarcane, red rot of
sugarcane, downy mildews of sorghum, and maize, wilts of pigeon pea and several viral
diseases. The destruction of crop debris especially the roots to reduce source of survival and
multiplication of inoculum help in control of many diseases. Pruning infected dead branches,
and removing infected fruit and other plant debris, help reduce the inoculum. Deep ploughing,
especially during summer in the tropics, buries the debris to such depths where the pathogens are
automatically destroyed. Fallows also help in the reduction of inoculum through sanitary effects
of decomposition of crop debris in absence of a host.
Exclusion of pathogen (legislation): This includes the measures designed to keep the pathogen
away from entering the area in which the host is growing, or to reduce the pathogen to its
minimum extent. This can be achieved by regulatory methods as: Inspection, crop rotation, etc.
Crop rotation is an old practice where sources of primary inoculum are eradicated. Soil borne
pathogens with narrow host range can be reduced in the soil by planting, for 3 to 4 years, crops
belonging to species or family not attacked by that particular pathogen.

10.2. Cultural Methods/Eradication of Plant Pathogens, etc.

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These methods include all man aimed activities in controlling diseases through the cultural
manipulation of plants such as:
• Sanitation - reduce inoculum potential
• Clean cultivation - use of disease-free seed and propagating material
• Crop rotation - control soil borne diseases
• Adjustment of planting time/sowing date
• Adjustment of planting density/spacing
• Optimum watering/fertilizer management
• Tillage practices

Cultural practices often offer the opportunity to alter the environment, the condition of the host,
and/or the behavior of the causal agent, to achieve economic management of disease. Most
cultural practices used to control plant diseases are preventive in nature. Cultural practices may
be employed, before or after planting. Deep ploughing and flooding are used before planting
while irrigation and fertilization can be applied several times during the crop season for disease
management.

Host eradication: Destruction of infected plant residues or removal of sources of infestation. All
the host plants infected by or suspected to harbor the pathogen are to be removed and burnt. This
results in elimination of the pathogen and preventing greater loss from the spread of the pathogen
to more plants. Host eradication is used in eradication of over-wintering/over-summering hosts,
eradication of alternate hosts and eradication of diseased plants. Host eradication is also done
routinely in nurseries, greenhouses, and fields to prevent the spread of many diseases by
eliminating infected plants, a ready source of inoculum. Removal and destruction of plants as
they become diseased is called rouging. Some pathogens of annual crops over-winter or over-
summer mainly in other perennial, usually wild plants or alternate hosts (wild or economically
less important). For example, Puccinia graminis tritici can be controlled by eradicating barberry
(wild alternate host).

10.3. Use of Resistance Genes and Plant Breeding for Resistance

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Resistance in plants, pre-existing or induced, is the natural trait developed during course of co-
evolution over millions of years. Thus, disease management through host resistance genes should
be considered most natural way to counter the threat posed to plants. Undoubtedly the most
attractive form of defense is provided by disease resistance, for as long as it remains effective it
provides protection at no cost to the farmer or the community.
The role of resistant genes in plant disease management should consider:-
• Easy availability of resistant genes: use of diverse type of genes.
• Easy transfer of genes in crop cultivars: production and insertion of resistance genes in
any plant genome using novel biotechnology techniques.
• Deployment of resistance genes: running out of cultivars due to ‘break down’ of
resistance.
Breeding for disease resistance
Breeding for disease resistance varieties isn’t only cost effective but also grower and consumer
friendly. For a successful and efficient resistance-breeding programme search for sources of
resistance is the pre-requisite step. The sources of resistance can be located in any of the
following forms:
• Land races/cultivated plants
• Wild/alien species
• Cytoplasm as source of resistance
• Induced/created variability through mutations

10.4. Biological Control of Plant Diseases

The practice in which the undesirable effects of an organism are reduced through the agency of
another organism. These methods aim at either direct protection of plants from pathogens or at
eradication or reduction of inoculum by using antagonistic organisms (mostly microorganisms).
Antagonism is a phenomenon, in which the activity of an organism (antagonist) inhibits or
limits or harms the activity of other organism. Microorganisms that have been identified and
exploited as biological agents include: Trycoderma, Gliocladium, Aspergillus, fluorescent
Pseudomonas and Bacillus species of bacteria have been widely recommended for the
management of several soil borne plant pathogens.

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The mechanism of biological control:
• Reduction of inoculum density.
• Suppression of germination and growth of pathogen.
• Displacing the pathogen from host residues (crop debris).
• Protection of an infection court (encouraging the soil microflora to prevent infection by
the particular pathogen).
• Stimulation of resistance response of the host
• Fungistasis – inhibition of the spore germination in soils.
• Suppressive soils (the pathogens can’t establish themselves or they are established but
fail to cause disease).

10.5. Chemical Protection of Plants

Chemicals used in plant protection are termed as pesticides. Depending on the kind of pathogens
they affect, the chemicals are called fungicides, bactericides, vermicides, nematicides,
insecticides, acaricides, rodenticides, herbicides, and so on. Application of fungicides is a
common and effective technique to manage plant diseases. Fungicides are applied to plants in
different forms and to various parts of plants.

Foliar Sprays and Dusts:

Chemicals applied as sprays or dusts on the foliage of plants are usually aimed at control of
fungus diseases and, to a lesser extent, control of bacterial diseases.

Contact Protective Fungicides: Most fungicides and bactericides are protectants and must be
present on the surface of the plant in advance of the pathogen in order to prevent infection. They
include Thiram, Zineb, Mancozeb, Carbamate, Chlorothalonil (available as Bravo, Daconil),
Captan, etc.

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Systemic Fungicides: are absorbed through the foliage or roots and are translocated within the
plant through the xylem. Some of the most important systemic fungicides are Metalaxyl,
Benomyl, Carboxin, Oxycarboxin, etc.

Antibiotics: are substances produced by one microorganism and toxic to another microorganism.
Most antibiotics known to date are products of branching bacteria (Streptomyces) and some fungi
(Penicillium) and are toxic mostly to bacteria and also certain fungi.

Eradicants: kill the fungus inside the host or may suppress the sporulation of the fungus without
killing it.
Disinfectant: A physical or chemical agent that frees a plant, organ, or tissue from infection.
Disinfestant: An agent that kills or inactivates pathogens in the environment or on the surface of
a plant or plant organ before infection takes place.
Fumigation: The application of a fumigant for disinfestations of an area or soil.
Fungistatic A compound that prevents fungus growth without killing the fungus.
Seed Treatment: Seeds, tubers, bulbs and roots are often treated with chemicals to prevent their
decay after planting or the damping-off of young seedlings.
Soil Treatment: Volatile chemicals (fumigants) are often used to fumigate the soil before
planting for reducing the inoculum of nematodes, fungi, and bacteria.
Surfactants: are often added to fungicides so that they spread better, thereby increasing the
contact area between fungicide and the sprayed surface.
Stickers: compounds with good sticking ability are added to increase the adherence of the
fungicide to the plant surface.

10.6. Integrated Control of Plant Diseases

Integrated control:
* An approach that attempts to use all available methods of control of a disease of a crop plant
for best control results but with the least cost and the least damage to the environment.
* The use of any two or more control measures that is compatible to the farming systems and
each other.

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In an integrated control program several control methods are used including:
• Regulatory inspections for healthy planting material,
• Cultural practices (crop rotation, sanitation, pruning, etc.),
• Biological control (resistant varieties),
• Physical control (storage temperature) and
• Chemical controls (soil fumigations, planting material treatment, sprays, disinfestations
of cuttings, tools, crates, warehouses, etc.).

The main goals of an integrated plant disease control program are to (1) eliminate or reduce the
initial inoculum, (2) reduce the effectiveness of initial inoculum, (3) increase the resistance of the
host, (4) delay the onset of disease, and (5) slow the secondary cycles

11. BIOTECHOLOGY AND PLANT PATHOLOGY

Bio – means life or living, so biotechnology is the application of living processes to technology.
One of the more popular definitions is the use of microorganisms, animal cells, plant cells, or
components of cells to produce products with living organisms.
Genetic engineering: the movement of genetic information in the form of genes from one cell to
another.
Recombinant DNA technology: the process of removing and inserting genes into DNA.
DNA (Deoxyribonucleic Acid): the coded material in a cell.

11.1. Application of Biotechnology

Farmers and plant breeders have relied for centuries on crossbreeding, hybridization and other
genetic modification techniques to improve crops with built-in protection against insect pests,
disease-causing organisms and harsh environmental conditions. Microscopic organisms lend
themselves well to genetic engineering. Microbes reproduce quickly and can be genetically
engineered to produce products needed by other plants, animals and humans. One of the first
commercial products made by genetic engineering was insulin, previously, available only from
animal pancreas tissue. However, a bacterium called E. coli was genetically engineered to
produce insulin much like cows produce milk and bees produce honey.

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Biotechnology:-
• Far more precise and selective than traditional breeding in which thousands of genes of
unknown function are moved into our crops.
• Allow plant breeders to select single genes that produce desired traits and move them
from one plant to another.
• Can help meet the ever-increasing need by increasing yields & decreasing crop inputs.
• Open up new avenues for working with nature by providing new biopesticides.

11.2. Tissue Culture Techniques of Significance to Pathology

Advances in plant tissue culture include meristem tip propagation, single cell culture, haploid
plant production, and protoplast isolation and regeneration into whole plants. Genetic
engineering techniques allow the detection, isolation, modification, transfer, and expression of
single genes, or groups of related genes, from one organism to another. Tissue culture of disease-
resistant plants is particularly useful with clonally propagated plants such as strawberries, apples,
bananas, sugar cane, cassava and potatoes. When plants were regenerated from leaf protoplasts
of a potato variety susceptible to both Phytophthora infestans and Alternaria solani, were
resistant to these pathogens. Similarly, plants exhibiting increased resistance to disease caused by
Cochliobolus and Ustilago were obtained from tissue cultures of sugar cane. Small or large
pieces of DNA are introduced into plant cells or protoplasts, and the DNA may be integrated in
the plant chromosomal DNA. Genetic material (DNA) can be introduced into plant cells or
protoplasts by several methods of which use of the natural gene vector system of A. tumefaciens
is one. The best documented cases involve plants engineered for resistance to viruses: papaya
engineered for resistance to papaya ring-spot virus, potato engineered for resistance to potato
leaf roll and potato Y viruses and wheat engineered for resistance to wheat streak mosaic virus.

11.3. Prospects of Biotechnology in Pathology

The “Green Revolution” had its origins in the science of genetics and technology, so the
application of new biotechnological methods could lead to a new revolution – the “Gene
Revolution”. Because genetic engineering is a relatively new field and the applications are so
numerous, the opportunities will expand as the field develops. It is now evident that genetic

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engineering and other forms of biotechnology hold great promise in controlling diseases, insects,
weeds and other pests. Further, the environment will be enhanced by less use of chemical
pesticides and greater use of biological controls.
12. INTRODUCTION TO MAJOR PLANT DISEASES

12.1. Root Rots and Damping-off

12.1.1. Damping-off: Diseases of seedlings occur in tropical and temperate climates, and in
every greenhouse. The fungi live on dead plant and animal materials as saprophytes or as
parasites of fibrous roots of plants. The disease affects seeds, seedlings, and roots of plants of
almost all kinds of vegetables, flowers, cereals and many fruit and forest trees. In all cases,
however, maximum damage is done to the seed and seedling roots during germination either
before or after germination. Older plants are seldom killed, but they develop root and stem
lesions and root rots, their growth may be retarded considerably and yields drastically reduced.
The most important and common damping-off fungus is pythium. The pathogen needs free water
for its zoospores to swim and infect. When a wet soil is infested heavily with Pythium, any seeds
or young seedlings in such a soil may be attacked by the pathogen. Certain other fungi such as
Phytophthora, Rhizoctonia, Fusarium and even some bacteria, when carried in or on the seed,
also cause damping-off and kill seedlings.

12.1.2. Root rots: Disintegration or decay of part or all of the root system of a plant. Several
Ascomycetes fungi such as Cochliobolus, Gibberella, and Gaeumannomyces attack primarily the
roots and lower stems of plants. Soil borne fungi, such as Fusarium solani and other species
cause root and lower stem rots of many vegetables, ornamentals, field crops, and even trees. As
a general rule, the root and stem rot diseases caused by these fungi appear on the affected plant
organs at first as water-soaked areas that later turn brown to black. In some diseases, lesions are
frequently covered by white fungal mycelium. The roots and stems are killed more or less
rapidly, and the entire plant grows poorly or is killed. These fungi are favored by high soil
moisture and high relative humidity in the air.

12.1.3. Galls and root-knots

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Galls: Enlarged portions of plant organs (stems, leaves, blossoms, roots). Gall is formed
primarily by bacteria of the genus Agrobacterium and by certain species of Pseudomonas,
Rhizobacter, and Rhodococcus. It affects woody and herbaceous plants belonging to 140 genera
of more than 60 families. Tumors or galls of varying size and shape form on the lower stem and
main roots of the plant. Infected nursery plants are unsalable. Plants with tumors at their crowns
or on their main roots grow poorly and their yields are reduced. Severely infected plants or vines
may die. Crown gall first appears as small, round, whitish, soft overgrowths on the stem and
roots, particularly near the soil line. As the tumors enlarge, their surfaces become convoluted,
and the outer tissues become dark brown due to the death and decay of the peripheral cells.

Root-knots: This nematode disease of root system of several hundred vegetable occurs
throughout the world, especially in areas with warm or hot climates and in greenhouses
everywhere. Though vegetables are attacked by a number of plant parasitic nematodes, root-knot
nematodes (Meloidogyne spp. Such as M. incognita and M. javanica) rank first among the
nematodes that cause damage to vegetable crops. Nematodes attack more than 2,000 species of
plants, including almost all cultivated plants. All common vegetables, such as cucumber, carrot,
cabbage, potato, onion, sweet potato, tomato, eggplant, beans, pumpkin, lettuce, celery, chilly,
radish and squash are attacked. Estimated crop losses due to this disease range from 5 to 60% in
different vegetables. Root-knot nematodes damage plants by causing the formation of swellings
of the roots. These effects not only deprive plants of nutrients, but also disfigure and reduce the
market value of many root crops. When susceptible plants are infected at the seedling stage,
losses are heavy and may result in complete destruction of the crop. Characteristic symptoms of
the disease appear on the underground parts of the plants. As the entire root system is affected
and becomes deformed, its efficiency decreases, causing reduction in top growth, stunting,
chlorosis, yellowing, vigor loss, premature defoliation and wilting. Heavily infected roots
become shorter with fewer branched roots and root hairs. The typical root galls or knots are two
to several times as large in diameter as the healthy root.

12.2. Stem and Branch Diseases

12.2.1. Stem rusts: Stem rust is a well recognized disease of cereal crops, occurring almost
everywhere these crops are grown. The rust usually appears late in the season when the

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temperature starts rising. The rust appears in the form of elongated, reddish-brown pustules,
primarily on the stem, followed leaf-sheaths and leaves. Thus, stem is most severely attacked
(stem rust). Black or stem rust is caused by Puccinia graminis tritici which often infects wheat,
barley and triticale. Three spore forms are produced on the cereal hosts (Urediospores,
Teliospores and Basidiospores). The Urediospores are disseminated over long distances by wind
and germinate on healthy parts of plant causing secondary infection. Thus Urediospores
propagate the disease in the entire field. As the plants approach maturity, Urediospores
production ceases and teliospores develop. The teliospores are dark brown and have thick walls.
If conditions are suitable each cell of Teliospore germinates to form Basidiospores. The
Basidiospores can’t infect wheat but infect barberry spp. which serves as alternate hosts.

12.2.2. Cankers: Cankers are localized wounds or dead areas in the bark of the stem or twigs of
woody and other plants that are often sunken beneath the surface of the bark. Innumerable kinds
of pathogens cause cankers on trees. The most common causes of tree cankers are Ascomycetes
fungi, although some other fungi, some bacteria, and some viruses also cause cankers.
Particularly common are cankers caused by several species of the Oomycetes Phytophthora spp.
The basic characteristic of cankers is that they are visible dead areas that develop in the bark and,
sometimes, in the wood of the tree. Cankers generally begin at a wound and much more serious
on fruit trees such as apple and peach.

12.2.3. Diebacks: Extensive necrosis of twigs beginning at their tips and advancing toward their
bases. In India, where the citrus root nematode (Tylenchulus spp.) was first reported (1961), up
to 80% orchards were found infested in certain areas. The pathogen causes slow decline of citrus
trees with die-back of twigs and debilitation of the trees. Affected trees show reduced terminal
growth, chlorosis, shedding of terminal leaves, die-back of branches and reduction in number
and size of fruits. Infected roots show brownish discoloration and ultimately decay. Die-back
usually appears after the rains when there is prolonged deposition of dew on plants.

12.3. Foliar Diseases

Many species of Ascomycetes and Imperfect fungi cause primarily foliage diseases but some
may also affect blossoms, young stems and fruit and even root. Most of the foliar Ascomycetes

78
reproduce by means of conidia formed on free hyphae or in pycnidia. The primary inoculum of
these fungi may be either ascospores or conidia and usually originates from infected fallen or
hanging leaves of the previous year.

12.3.1. Downy mildews: Downy mildews are primarily foliage blights. They attack and spread
rapidly in young, tender green leaf, twig, and fruit tissues. Downy mildews can cause severe
losses in short periods of time. The severity of loss depends on the prolonged presence of wet,
cool weather during which the downy mildews sporulate profusely, cause numerous new
infections, and spread into and rapidly kill young succulent tissues. All species of the family
Peronosporaceae are obligate parasites of higher plants and cause downy mildew diseases on
most cultivated grain crops, vegetables, field crops, ornamentals, shrubs, and vines. Downy
mildew Oomycetes produce sporangia on sporangiophores; sporangia germinate by producing
zoospores or, at higher temperatures, by producing germ tubes.

Some of the most serious downy mildew Oomycetes and the diseases they cause include:
• Peronospora: causing downy mildew of onion (P. destructor), of soybeans (P.
manchurica) & of tobacco (P. tabacina);
• Plasmopara: causing downy mildew of grape (P. viticola), and
• Sclerophthora: causing downy mildew of cereals (corn, rice, sorghum, wheat).

The pathogen Plasmopara viticola affects the leaves, fruit, and shoots of grapevines. When the
weather is favorable and no protection against the disease is provided, downy mildew can easily
destroy 50 to 75% of the crop in one season. At first, small, pale yellow, irregular spots appear
on the upper surface of the leaves, and a white downy growth of the sporangiophores of the
Oomycetes appears on the underside of the spots. Infected grapes are quickly covered with the
downy growth may become distorted or thickened, and may die.

12.3.2. Powdery mildews: Powdery mildews are probably the most common, widespread, and
easily recognizable plant diseases. They affect all kinds of plants except gymnosperms. Powdery
mildews appear as spots or patches of a white to grayish, powdery, mildew growth on young
plant tissues or as entire leaves and other organs being completely covered by the white powdery

79
mildew. Powdery mildew is most common on the upper side of leaves, but it also affects the
underside of leaves, young shoots and stems, buds, flowers, and young fruit.
Fungi causing powdery mildews are obligate parasites: they cannot be cultured on artificial
nutrient media. The powdery mildew diseases of the various crops are caused by many species of
fungi of the family Erysiphaceae.
The most important diseases they cause are:
• Erysiphe graminis: causes powdery mildew of cereals and grasses
• Erysiphe polygoni: ,, ,, ,, ,, legumes, cabbage and crucifers
• Sphaerotheca pannosa: ,, ,, ,, ,, peach and rose
• Erysiphe cruciferarum: ,, ,, ,, ,, rape seed and mustard

12.3.3. Leaf rusts: The plant rusts, caused by Basidiomycetes of the order Uredinales, are
among the most destructive plant diseases. The name rust fungi come from the fact that often
their pustules and spores have a rusty yellow color. Leaf rust is a serious disease of wheat, barley
and many grasses. Pustules first develop on the lower leaves and leaf sheaths and progressing up
to the flag leaf. Under favorable conditions, symptoms may also occur on stems, awns and
glumes of the head. Pustules are small, round to oval, raised and orange yellow in color. Leaf
rust of cereals is caused by species of fungi of the genera Puccinia. Leaf rust also attacks
vegetables, ornamentals and trees such as apple, coffee and others. The most commonly known
diseases are:
• Leaf rust on wheat (Puccinia recondita)
• Leaf rust on barley (Puccinia hordei)
• Faba bean rust (Uromyces viciae fabae)
• Garlic rust (puccinia allii)
• Coffee leaf rust (Hemileia vastatrix)

12.3.4. Anthracnose, blights and leaf spots

Anthracnose: Necrotic and sunken ulcer-like lesion on the stem, leaf, fruit, or flower of the host
plant caused mainly by a certain group of fungi. Anthracnose diseases are caused by
Ascomycetes and Deuteromycetes. Anthracnose, meaning blackening, is disease of the foliage,
stems, or fruits that typically appear as dark-colored spots or sunken lesions with a slightly raised

80
rim. Anthracnose diseases of fruit often result in fruit drop and fruit rot. Although severe
everywhere, anthracnose diseases cause their most significant losses in the tropics and
subtropics. Some causal agents of anthracnose diseases:
• Sorghum anthracnose (Colletotricum graminicola)
• Bean anthracnose (Colletotricum lindemuthianum)
• Mango anthracnose (Glomerella cingulata)

Leaf spots and blights: Almost all bacterial spots and blights of leaves, stems, and fruits are
caused by bacteria in the genera Pseudomonas and Xanthomonas.
• Pseudomonas syringae: causing wildfire of tobacco, angular leaf spot of cucumber, halo
blight of beans and bean leaf spot.
• Xanthomonas compestris: causing common blight of beans (X. campestris pv. phaseoli),
angular leaf spot of cotton (X. campestris pv. malvacearum), bacterial spots of stone
fruits (X. arboricola pv. pruni) and of tomato and pepper (X. campestris pv. vesicatoria).

Blight: General and extremely rapid browning and death of leaves, branches, twigs, and floral
organs. Leaf spots and blights can also be caused by the following fungal species:
• Early blight of potato (Alternaria solani)
• Late blight of potato and tomato (Phytophthora infestans)
• Chocolate spot of faba bean (Botrytis fabae)
• Banana leaf spot or Sigatoka disease (Mycosphaerella musicola)

12.3.5. Streaks and stripes: The bacterial stripe (Pseudomonas andropogoni) and streak
(Xanthomonas holcicola) diseases are common on sorghum. Symptoms are more pronounced
and defoliation is more severe than with the similar diseases on sugarcane. Stripe rust (Puccinia
striiformis) is also a serious disease of wheat and barley that occur at lower temperatures than are
optimal for leaf and stem rust.

12.3.6. Viral mosaics: Mosaics are characterized by light-green, yellow, or white areas
intermingled with the normal green of the leaves or fruit or flowers. Depending on the intensity
or pattern of discolorations, mosaic-type symptoms may be described as mottling, streak, ring

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pattern, line pattern, vein-clearing, vein-banding, or chlorotic spotting. There are two closely
related viruses of economic importance in the genus Tobamovirus: tobacco mosaic virus (TMV),
which infects tobacco and many other, mostly solanaceous (tomato mosaic virus), which infects
tomato.

12.4. Floral and Seed Diseases

12.4.1. Smuts and ergot

Smuts: Plant smuts, caused by Basidiomycetes of the order Ustilaginales, occur throughout the
world. There are approximately 1,200 species of smut fungi. Most smut fungi attack the ovaries
of small grains and many grasses, and produce black spore masses (teliospores) that partially or
completely replace heads, spikelets or kernels. Most smut fungi produce only two kinds of
spores: Teliospores (the sexual, thick-walled resting spores of rust and smut fungi) and
Basidiospores (sexually produced spores borne on a basidium).smut diseases caused by
Ustilago species are usually initiated via floral infection.

Smuts generally overwinter as teliospores on contaminated seed, in plant debris, or in the soil.
The Teliospores are not infectious but produce Basidiospores, which on germination infect or
penetrate the host tissue. Smut fungi have only one generation per year, each infection resulting
in one crop of Teliospores per growing season. The control of smuts is primarily by use of
resistant varieties and seed treatment. The most common smut fungi and the diseases they cause
are the following:
• Ustilago: causing corn smut (U. maydis), loose smut of cereals (U. avenae, U. nuda, and
U. tritici), and sugarcane smut (U. scitaminea).
• Tilletia: causing covered smut of wheat (T. caries).
• Sphacelotheca: causing the sorghum smuts (S. sorghi).
• Urocystis: causing onion smut (U. cepulae)

Ergot: Ergot occurs worldwide, most commonly on rye but it also occurs on wheat, triticale,
barley, oats, millet and many grasses. The fungus (Claviceps purpurea) attacks only the seed-
producing organs. The disease destroys 5 to 10% of the grains in infected heads, but its main
importance is due to the fact that it replaces the grains with fungal sclerotia, which are poisonous

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to humans and animals. Grain containing more than 0.3% by weight of the ergot sclerotia can
cause ergotism, and therefore such grain may not legally be sold and milled for flour and human
consumption. Moreover, feeding livestock with cleanings from contaminated grain or grazing in
pastures that have infected grass heads can lead to reproductive failure or “gangrene” of the
peripheral parts of the animals.

Ascospores are carried by wind or by insects to young open flowers, where they germinate and
infect the ovaries directly or by way of the stigma. During infection, the fungus secretes the
enzyme catalase, which apparently plays a role in development of the ergot disease by
suppressing the defenses of the host. Within about a week, the fungus in the ovary forms conidia.
The conidia exude from the young florets as creamy droplets known as the “honeydew” stage.
The honeydew attracts insects, which become smeared with the conidia of the fungus and carry
them to healthy flowers, which the conidia infect. Conidia are also spread to flowers by
splashing rain. Gradually, infected ovaries, instead of producing normal seed, become replaced
by a hard mass of fungal mycelium, which eventually forms the characteristic ergot sclerotium.
The sclerotia have developed into conspicuous dark, hornlike structures, two or three times as
large as normal kernels. The sclerotia mature about the same time as the healthy seeds and either
fall to the ground, where they overwinter, or are harvested with the grain and may be returned to
the land with the seed. The fungus over-winters on the soil or as sclerotia in plant debris. Ergot is
more prevalent in cool, temperate climates.

The control of ergot depends entirely on using clean seed or seed that has been freed from ergot.
Sclerotia may be removed from seed by machinery or by soaking contaminated seed for three
hours in water and then floating off the sclerotia in a solution of about 18 kilograms of salt in
100 liters of water. Ergot sclerotia do not survive for more than a year and do not germinate if
buried deep in the ground. Therefore, deep plowing or crop rotation with a non-cereal for at least
a year helps eliminate the pathogen from a particular field.
12.4.2. Soft Post-harvest Decays:
Some of the most common Ascomycetes that cause post-harvest diseases are listed here.

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Alternaria: The various species of Alternaria cause decay on most fresh fruits and vegetables
either before or after harvest. Symptoms appear as brown or black, flat or sunken spots with
definite margins.
Fusarium: causes post-harvest pink or yellow molds on vegetables especially on root crops,
tubers, and bulbs. Contamination with Fusarium usually takes place in the field before or during
harvest, but infection may develop in the field or in storage. Losses are particularly heavy with
crops such as potatoes that are stored for long periods of time.
Geotrichum: causes the sour rots of citrus fruits, tomatoes, carrots, and other fruits and
vegetables. The fungus penetrates fruits, usually after harvest, at wounds of various sorts.
Infected areas appear water soaked and soft and are punctured easily. Fruit flies, which are
attracted to tissues affected with sour rot, spread the pathogen further.
Penicillium: The various species of Penicillium cause the blue mold rots and the green mold
rots. They are the most common and usually the most destructive of all post-harvest diseases,
affecting most kinds of fruits and vegetables. Penicillium enters tissues through wounds.

12.5. Fruit Diseases and Soft Rots

12.5.1. Brown rots: Brown rot occurs wherever stone fruits are grown and there is sufficient
rainfall during the blossoming and fruit ripening periods. It affects peaches, cherries, apricots,
and almonds with about equal severity. The brown rot of stone fruits is caused by the fungus
Monilinia fructicola.

12.5.2. Soft rots of vegetables and fruits: Bacteria are invariably present whenever fleshy plant
tissues are rotting in the field or in storage, and the foul smell given off by such rotting tissues is
due to volatile substances released during the disintegration of plant tissues by such bacteria.
Rotting tissues become soft and watery, and slimy masses of bacteria and cellular debris
frequently ooze out from cracks in the tissues. Some bacteria that cause soft rots in the field or in
storage are listed below.
Erwinia: the “carotovora” group causing soft rots of numerous fleshy fruits, vegetables, and
ornamentals (E. carotovora).
Pseudomonas: causing soft rots of fleshy fruits and fleshy vegetables (P. fluorescens), such as
the pink eye disease of potato, the slippery skin disease of onion and the sour skin of onion.
Bacillus: causing rotting of potatoes and tobacco leaves in storage.

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Clostridium: causing rotting of potatoes and tobacco leaves in storage.
The control of bacterial soft rots is difficult and depends on proper sanitation, avoiding injuries,
keeping storage tissues dry and cool, assuring good insect control and practicing crop rotation.

12.6. Vascular Wilts

Vascular wilts are widespread, very destructive, spectacular, and frightening plant diseases. They
appear as more or less rapid wilting, browning, and dying of leaves and succulent shoots of
plants followed by death of the whole plant. Wilts occur as a result of the presence and activities
of the pathogen in the xylem vessels of the plant. Entire plants may die within a matter of weeks,
although in perennials, death may not occur until several months or years after infection. As
long as the infected plant is alive, wilt-causing fungi remain in the vascular (xylem) tissues and a
few surrounding cells. Only when the infected plant is killed by the disease do these fungi move
into other tissues and sporulate at or near the surface of the dead plant. The most common two
genera of fungi that cause vascular wilts are Fusarium and Verticillium. Each of them causes
disease on several important crop, forest and ornamental plants. All vascular wilts have certain
characteristics in common. The leaves of infected plants or of parts of infected plants lose
turgidity, become stunted and soon wilt and finally die. Occasionally, entire fields of tomatoes
are killed or damaged severely before a crop can be harvested.

12.6.1. Fusarium wilts: Fusarium causes vascular wilts of vegetables and flowers, herbaceous
perennial ornamentals and plantation crops. Most of the wilt causing Fusarium fungi belongs to
the species Fusarium oxysporum. The pathogen is a soil inhabitant. Between crops it survives in
infected plant debris in the soil as mycelium and in all its spore forms but, most commonly as
Chlamydospores (cooler temperate regions). It spreads over short distances by means of water
and contaminated farm equipment and over long distances primarily in infected transplants or in
the soil carried with them.

12.6.2. Verticillium wilts: Verticillium causes vascular wilts of vegetables, flowers, field crops,
perennial ornamentals, and fruit and forest trees. Two species, Verticillium albo-atrum and V.
dahliae, attack hundreds of kinds of plants, causing wilts and losses of varying severity.
Verticillium attacks more than 200 species of plants; most of them are vegetables (tomato,

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pepper, watermelon, flowers, etc.); fruit trees (cherry, peach, strawberries, roses, etc.) and field
crops (cotton, potato, peanuts, etc.). Verticillium infection results in defoliation, gradual wilting
and death of successive branches or an abrupt collapse and death of the entire plant. The fungus
(Verticillium) spread by contaminated seed, by vegetative cuttings, and tubers, by scions, and
buds, by wind and surface ground water and by the soil itself.

12.6.3. Bacterial wilts: Vascular wilts caused by bacteria affect mostly herbaceous plants such
as several vegetables, field crops, ornamentals and tropical plants. The bacteria and the most
important vascular wilts they cause are listed.
Corynebacterium: causing bacterial canker and wilt of tomato, and wilt of bean. Erwinia:
causing bacterial wilt of cucurbits (E. tracheiphila), and fire blight of pome fruits (E.
amylovora). Bacterial vascular wilts are similar to the fungal vascular wilts caused by Fusarium
and Verticillium. However, whereas in fungal wilts the fungi remain almost exclusively in the
vascular tissues until the death of the plant, in bacterial wilts the bacteria often destroy (dissolve)
parts of cell walls of xylem vessels or cause them to rupture quite early in disease development.
The bacteria spread from plant to plant through the soil, through handling and tools, through
direct contact of plants, or through insect vectors
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