INTRODUCTION

Dependence of man on plants for food dates back to more than 12000 years. Other animals and plants
existed on this planet before the arrival of man and these later became the security against starvation of
man. The animals domes- ticated by man almost wholly depend on plants. In addition to food, plants
and animals are the primary sources of materials for other necessities of life such as clothing, house
building materials, furniture, drugs and fuel etc. Coal and petroleum products are also derived from the
remains of plants buried deep in soil. The congenial climate, the rains, etc are made and modified by the
plants.

The availability of food to human population in reasonable quantity is governed by the population
density needing food, cultivated land area avail- able for food production and production of food per
unit land area. Among these determinants availability of land is almost fixed. Production per unit land
area does not rise fast enough while the human populations is a fast rising phenomenon. If the increase
in food production can be matched with increase in the number of mouths to be fed, probably the
problem will not be acute. However, the rise in human population is much faster than the rise in food
production. Although total foodgrain production in the world has been rising and about 1400 million
hectares of land surface (12% of the total) is being cultivated for some food crop, the rise in production
has not matched with the rise in consumer numbers. For instance, the total food production in India has
shown an overall increase of about 65% since 1950 while the population during the same period showed
an increase of about 175%.

In the year 1940 the human population on this planet was around 2140 million which rose to 3280
million in 1965, to 3950 million in 1975 and to 4900 million in 1985. In the year 2000 it stood at around
6800 million. Overall rise in world population during the 60 years was about 218%. During the 25 years
since 1940 the average rise per year was around 2%. It has shown only slight increase in the average
percentage but the picture is different when these figures are considered on regional basis. Two third or
more of the total world population is in the countries of Asia, Africa and Latin America.

WHAT IS A PLANT DISEASE?

When the plant is under some sort of stress and is not developing and functioning in a manner expected
from it, we call it diseased. This does not define the term "disease". Often the symptoms appearing as a
result of a disease, the cause of the disease and the injuries caused to the plant have been considered
synonymous. However, they signify only the condition of the plant due to disease or the cause of the
disease.

In 1858, Julius Kuhn, in Germany, had defined plant disease as abnormal changes in the physiological
processes which disturb the normal activity of plant organs. A similar definition was given by H.M. Ward
in 1896 who defined disease as a condition in which the functions of the organism are improperly
discharged or, in other words, it is a state which is physiologically abnormal and threatens the life of the
being or the organ. In 1918, E.J. Butler had defined disease as variation from normal physiological
activity which is sufficiently permanent or extensive to check the performance of natural func- tions by
the plant or completion of its development. Similarly, according to the American Phytopathological
Society (Phytopathology 30: 361-368, 1940) disease is a deviation from normal functioning of
physiological processes of sufficient duration or intensity to cause disturbance or cessation of vital activi-
ties. The British Mycological Society (Trans. Brit. Mycol. Soc. 33: 154-160, 1950) also defined disease as a
harmful deviation from the normal function- ing of process. Wheeler (1975) broadly considered plant
disease as all the malfunctions which result in unsatisfactory performance of the plant or which reduce
ability of the plant to survive and maintain its ecological niche. Thus, the major thrust in defining a
disease since the time of Julius Kuhn was on the deviation from the "normal", a condition which may be
variable.

An analytical approach to definition of the term "disease" was made by Horsfall and Dimond (1959) who
clarified many misconceptions. According to them disease (i) is not a pathogen, it is caused by a
pathogen, (ii) is not the symptoms or effects seen on the plant, symptoms result from disease, (iii) is not
a condition as the condition results from the disease, (iv) is not any injury which results from disease as
well as from any traumatic cause and (v) cannot be catching or infectious, it is actually the pathogen
which is catching or trasmitted. They defined disease as a malfunctioning process in the plant body due
to cotinuous irritation which results in some suffering. It is a pathological process in plants and animals
(including man).

The pathogens bring about the irritating processes resulting in a diseased conditions of the plant
through different but interrelated pathways:

1) By utilizing the host cell contents.

2) By causing death of cells or by interfering with their metabolic activities through enzymes, toxins and
growth regulators.

3) By weakening of tissues due to continuous loss of nutrients.

4) By interfering with translocation of food, minerals and water.

Dispersal of Plant Pathogens

The dispersal, transport or transmission of the pathogen or the disease important not only for spread of
the disease in the population but also for continuity of the life-cycle and evolution of the pathogen. The
living host or other organic substrates cannot indefinitely provide space and nutrition for the growing
population of the animate pathogens. This compels them to move out to new, hitherto uninhabited,
sites otherwise the pathogen would die due to starvation resulting from exhaustion of the host and due
to overcrowding. Nature has provided the pathogens the necessary mechanisms for exit from the host
and move to new locations. In fungi, production of asexual and sexual spores follows the active
vegetative growth in or on the invaded host tissues. These are dispersed mechanically in time and space
by various means. In bacterial diseases, the bacterial cells come out on the host surface as ooze or the
tissue may be disintegrated to such an extent that the bacterial mass is exposed and then dispersed by
various physical and biological agencies. Viruses have no such organs. They are transmitted by insects
and man Nematodes are themselves motile and try to move out of the exhausted and overcrowded
sites and then may be moved with soil, water and other means

Pathogens of all infectious diseases are transmissible, i.e., they are carried by various means from the
source of survival or from a diseased individud in the population to the healthy individual. Some
diseases may even be contagious i.e., they can be transmitted by contact between diseased and healthy
plants. Many mechanisms described for survival of pathogens apply to dis persal of pathogens also. In
general the seed-borne pathogens or diseases are dispersed by the movement of seed, soil-borne
pathogens through movemen or displacement of soil and through root contacts, and those surviving
hosts (collateral or alternate hosts) in active stage through wind-borne spores other structures and
insects. However, in majority of diseases a combination on of mechanisms, aided by external physical
and biological forces, operates dispersal (as well as survival). The same pathogen may be carried by soil
as wild

Well as by seed and its transport may be aided by water, wind, insects, etc The two links in the infection
chain of an animate pathogen, viz., survivo through dormant structures and dispersal of the pathogen
are very closely bound with each other. Actually, the dormant structures of fungi provide means of
dispersal in time, i.e., the pathogen is retained in a viable condition over a period of time enabling it to
be transported through physical agencies without being harmed. In lower fungi, many pathogens are
transported only through their resting structures in soil. The resting structures of fungi on seed or in
plant debris are other examples. Among soil fungi, fungistasis provides a large, well distributed reserve
of inactive spores able to colonize new sub- strates quickly when available.

The dispersal of pathogens is accomplished in the following manner: 1. Direct (active or autonomous)
dispersal, such as dispersal in and by soil

and by seed and planting materials. 2. Indirect (passive) dispersal involving the role of man, insects,
nematodes and other animals, water and air.
The knowledge of these methods of dispersal is essential for effective man- agement of plant diseases
because possibilities of preventing dispersal and thereby breaking the infection chain exist.

AUTONOMOUS DISPERSAL

Autonomous dispersal of bacteria, fungi, viruses and nematodes is accom- plished through the agency of
soil, seed and plant organs during normal agronomic operations. There is no primary role of external
agencies like insects, wind, water etc. in this type of dispersal.

PASSIVE DISPERSAL

The passive dispersal of plant pathogens is accomplished through the agency of members of animal
kingdom (man, insects, nematodes, farm and wild animals, birds, etc.) and air and water (irrigation and
rains).

DISPERSAL BY MEMBERS OF ANIMAL KINGDOM

(i) Man: The most highly evolved member of the animal kingdom, man, can be considered as one of the
most important single factor affecting dispersal of plant pathogens in a limited area or over long areas
throughout the world. The day-to-day activities of man in normal farming practices and his trade or
commerce activities that are helping dispersal of pathogens and introduction of diseases in new areas
are:

(1) Vegetative propagation of ornamentals, fruit trees, tuber like crops potato and sweet potato,
and plantation crops like banana and sugarcane is a highly efficient means of introducing a
disease in new areas. In depending on vegetative propagation for such crops man invariably
helps dispersal by moving the diseased propagating material of plants from field to field,
orchard to orchard and from one geographic region to another. Late blight (Phytophthora
infestans), bacterial wilt (Ralstonia solanacearum) and viruses of potato, ratoon stunting disease
(Clavibacter xyli subsp. xyli) and mosaic of sugarcane, banana bunchy top virus and Shigatoka
(Cercospora musae), citrus canker (Xanthomonas axonopodis pv. citri) and many diseases of
grapevines are some of the examples. Citrus canker originated in South-east Asia and was
carried by propagation material to USA and South Africa. Its recurrence in USA and Australia
after eradication is also attributed to import of infected propagating stock. The downy mildew of
grapevines (Plasmopara viticola) originated in North America and was introduced into Europe
with the introduction of the planting material. Same is true for the powdery mildew of
grapevines (Uncinula necator). Role of commercial nurseries is very important. The nurserymen
who do not care to follow proper hygienic methods distribute infected or contaminated
seedlings, grafts, etc. to growers and thus supply the inoculum of pathogens also.

(2) The seed trade is another means of dispersal of pathogens in which man plays the most
crucial role. There is a close relationship between pattern of seed trade and that of dispersal of
pathogens (autonomous as well as passive). The import and export of contaminated seed
without proper precautions (cer- tification and quarantine) has caused entry of pathogens from
one country to another. Entry of soybean and sugarbeet seed from abroad had introduced
certain fungal pathogens that were not present in India. Karnal bunt (Neovossia indica) was
endemic to India. Movement of wheat seed for experimental purposes caused its detection in
Mexico and some European countries. Within India also this disease has spread to different
states all over the country through seed. The sale of seed from crops badly affected by a seed-
borne pathogen or disease is a common method of dispersal of such other destructive
pathogens as Ustilago segatum tritici (loose smut of wheat), Sphacelotheca sorghi (grain smut of
sorghum), Claviceps fusiformis (ergot of pearl millet) and Anguina tritici (ear cockle of wheat)
and yellow ear rot (Rathayibacter tritici).

(3) Spores and other structures of fungi, bacterial cells in ooze, and even viruses can be carried
by workers' clothings, shoes, hands, etc. from plant to plant and from field to field.

(4) The use of contaminated implements, cutting knives, etc. help in trans- mission of many soil-
borne and other pathogens and also the pathogens present in tubers, sugarcane cuttings, bulbs,
etc. Transmission of bacterial wilt (R. solanacearum), ring rot (Clavibacter michiganensis subsp.
sepedonicus), bacterial wilt of banana (R. solanacearum) by cutting knive or matchets during
planting and in banana wilt during harvest of fruit bunches is well documented. The ratoon
stunting bacterium has no other means of transmission except during harvesting by cutting
tools.

(5) Man is responsible for spread of almost all the graft transmissible diseases. Grafting and
budding between healthy and diseased plant is the most effective method of distribution of
virus pathogens of fruit trees. Careless selection of rootstock and scion of citrus is known to
transmit such diseases as Tristeza and other viruses.

(6) Cultural operations such as hoeing, weeding, pruning, harvesting and marketing also packing
of fruits in the orchards, all carried out by man, not only transmit diseases by contact between
the plant and the contaminated tools and by providing wounds on the plant for easy entry of
pathogens but also in transporting the pathogen to long distances as contaminants on the
packing cases or crates.

(iv) Dispersal by Other Animals: In addition to man, insects and nematode other larger animals
can also accidentally transmit plant pathogens. Although experimental evidence is lacking, the
possibility does exist that birds feeding on insect vectors can transport pathogens to distances
not ordinarily covered by the vector itself. Dispersal of spores by birds carrying them on their
feather is also possible. The dispersal of phanerogamic parasites by birds is an established fact.
Birds feeding on berries of Dendrophthoe deposite the seed with their excreta on other trees.
Stem fragments of dodder (Cuscuta) are carried by birds for preparing their nest and thus these
get transported to new locations. Rodents and fur-bearing animals can also be source of
transmission of pathogens. Among mammal cattle feeding on contaminated fodder often pass
out viable fungal propagules in the dung which can act as source of inoculum when used as
manure. Thus, conidia of Colletotrichum falcatum (red rot of sugarcane) and sclerotia of many
fungi have been detected in cattle dụng.

Dispersal by Fungi and Phanerogamic Plant Parasites

Fungi growing in soil are suspected to carry bacterial cells externally and put them in roots of
susceptible plants. However, the most interesting interac tion is between certain fungi and soil-
borne viruses. Many soil-borne viruses are transmitted by members of Chytridiales and
Plasmodiophorales of fungi. The established species number five. There are also reports of virus
transmis sion by Oomycetes, Erysiphaceae and Pucciniaceae. The chydrid fungi are characterized
by the size of posteriorly uniflagellate zoospores which have a characteristic “jerky” swimming
pattern, and by the morphology of their single- celled resting spores. The plasmodiophorales
have biflagellate, heterokont zoospores.

The fungus-transmitted viruses generally comprise of heterogeneous groups with respect to

their usual properties. Two types of virus-fungus relationships are recognized on the basis of
method of virus acquisition and by the location of the virus particles relative to the fungus
resting spore. The in vitro transmis sion method (Olpidium brassicae x tobacco necrosis virus)
involves in vitro acquisition, in which virions are not located within the resting spores. The in
vivo transmission method (Olpidium brassicae x lettuce big vein virus) involves in vivo
acquisition, in which virus enters the thallus as it grows in a virus infected host, and the virus is
located within the resting spores.

In vitro transmission is found with two species of Olpidium and with the polyhedral viruses. In
the transmission of tobacco necrosis virus by Olpidium brassicae acquisition begins when virus-
free zoospores released from resting spores or from vegetative sporangia encounter virions in
soil water. The virus particles are tightly and specifically adsorbed to the zoospore membrane.
This adsorption probably involves receptors in the zoospore membrane and the Particular coat
protein of the virions. In the transmission of lettuce big vein Virus by Olpidium brassicae the
virus is taken by the fungus thallus while It is growing in an infected host. The infectivity of air
dried soil is retained for 8 years. Similarly, soil-borne wheat mosaic virus and barley mosaic virus
are Internally zoospore borne in Polymyxa graminis and beet necrotic yellow vein Mosaic virus
in Polymyxa betae. Unconfirmed transmission of virus like par-Ticles from Chenopodium quinoa
by powdery mildew and rust fungi and the Transmission of virus to Phaseolus vulgaris by
Uromyces phaseoli and to barley By Erysiphe graminis are also reported.

Host specialization or host specificity of the fungus pathogen is an important characteristic of

fungal vectors of viruses. Host specialization has been known in Olpidium brassicae. The lettuce,
tomato and red clover isolates of the fungus are plurivorus and transmit the tobacco necrosis
virus to tobacco, tulip, bean and potato. More specific fungus isolates from oats, melon and
crucifers are less efficient or non-vectors. In O. bornovanus, there are at least three host specific
strains (cucumber, melon and squash) in the cucurbit group and they vary in their ability to
transmit different viruses. Spongospora subterranea f.sp. nasturtii is specific for watercress.
Some isolates of Polymyxa betae infect most chenopodiaceous hosts while others are highly
specific for Amaranthus spp. Or Portulaca spp.

The phanerogamic plant parasite dodder (Cuscuta spp.) is a total stem parasite of crops and fruit
and roadside trees. It establishes parasitic relation- ship with plants through haustoria sent into
the xylem and phloem of the host. Many viruses and the phloem-limited citrus greening
bacterium (Liberobacter asiaticum) have been shown to be transmitted by dodder (cf. Singh,
1998). Through the haustoria the dodder stems pick up the virus from the plant and when these
stem pieces are dispersed and lodged on new host plants the virus or other pathogens are
transmitted to the new plant during infection by dod- der. In plant species where vascular union
by grafting has failed to transmit the virus, dodder has been successfully used to act as a bridge
between diseased and healthy plant vascular bundles. Many viruses can multiply in dodder. If
this multiplication occurs and if the receiver (healthy) plant is kept in dark transmission by
dodder is more efficient.

Tobacco rattle virus (TRV) is transmitted by at least 6 species of Cuscuta and there is also
infection of the dodder. Cucumber mosaic virus (CMV) is trans- Imitted by at least 10 species.
The virus infects the vector and multiplies in it. White clover mosaic virus can infect dodder and
transmission can also occur. The alfalfa mosaic virus occurs in at least 5 species of Cuscuta. Beet
yellows and beet curly top viruses are also transmitted by dodder. Cuscuta species transmit the
tomato mosaic virus in winter but not in summer. Aphids can pick up a virus from dodder
established on a host.