Survival and Dissemination of Plant Pathogens

Plant Pathogens & Principles of Plant Pathology
Organisms in Seeds
Organisms can easily live in seed and are often spread from garden to garden in this way.
For this reason, unless a unique garden variety is being preserved, gardeners should not save
seeds from their garden, but should purchase seeds that were produced in parts of the country
where diseases do not occur. Seedborne diseases can also be greatly reduced by using a chemical
seed treatment.
Survival of Plant Pathogen
Any pathogen can cause disease under favourable conditions. The only requisite factor is
that the pathogen must come in contact with the host for the development of the disease.
Pathogen itself or its parts that are capable of causing disease when brought near a host is called
inoculum. Fungal pathogens are diversified, where the vegetative body (hyphae), dormant
mycelium, (embedded in the embryo of seeds or other plant parts), special reproductive
structures (rhizomorphs, sclerotia, chlamydospores), various types of asexual spores (sporangia,
sporangiospores, zoospores, conidia) and sexual spores (oospores, zygospores, resting spores,
ascospores, basidiospores, etc.), serve as inocula. In the case of viruses and plant pathogenic
bacteria, the individuals are acting as inocula, since they do not produce any special type of
infective units like resting spores or endospores, etc. But in the case of Streptomyces sp.
(Actinomycetes), fragments of filaments and spore-like cells serve as inocula. In phanerogamic
parasites, seeds are the potential inocula.
Seeds in the soil survive for longer period. Orobanche seeds survive for about 13 years.
Seeds are abundantly produced for their multiplication, which could attack the host plants. But
dodder is an exception because broken bits of shoot can attack host plant. In any locality a time
lag exists between harvest of a crop and subsequent sowing. Year after year, diseases appear in
the newly sown crops. There should be some link between the previous crop and the subsequent
new crop to revive or continue the life cycle. The existence of the pathogen between the two crop
seasons is the vulnerable period in its life cycle. Hence knowledge of the survival ofthe pathogen
in the off-season is useful for the plant pathologists to device effective control measures.
The establishment of a plant pathogen in a geographical location presupposes its ability
to survive, not only during its parasitic relations with its hosts, but also during off-seasons in
which the hosts are not growing. In temperate zones, plant pathogens must be adapted for
survival overwinters or oversummers, like the powdery mildew pathogen that attacks fall-seeded
176 www.AgriMoon.Com
wheat. In the Torrid Zone, plant pathogens must be able to survive the dry seasons, during which
susceptible plants are not growing.
These sources of survival of pathogens or the sources for renewal of infection chain can be
grouped as follows:
1. Survival by means of specialized resting structures
2. Survival as saprophytes
3. Survival in vital association with living plants
4. Survival in association with nematodes and fungi
5. Survival in association with insects
6. Survival on agricultural materials
7. Survival on surface water
1. Survival by means of specialized resting structures
Enduring structures of plant pathogens may be as simple as conidia or as complex as
perithecia. Apparently, ascospores or conidia derived from them, serve to carry the pathogen
causing peach-leaf curl (Taphrina deformans) over the winter. Conidia of Alternaria solani, the
pathogen of early blight of potato and tomato, survive for eighteen months in dried diseased
leaves. Specialized thick-walled chlamydospores of Fusarium and other Imperfect fungi, spores
of many smut fungi and the amphiospores, uredospores and teliospores of certain rust fungi also
are important enduring structures. The resting spores of Plasmodiophora brassicae may survive
for ten years in soils infested upon the disintegration of clubbed roots. The oospores of downy-
mildew fungi survive in the soil between growing seasons. In fact, oospcres of the fungus that
causes onion mildew do not germinate until several years after their formation.
Some fungi survive unfavourable seasons in the form of sclerotia. Those produced by the
omnivorous cottony-rot fungus, Sclerotinia sclerotiorum,can survive for years in a dry
atmosphere. They decay rapidly, however, in warm moist soil. Cold induced dormancy probably
accounts for their ability to survive winters in temperate zones. Some powdery mildew fungi and
other ascomycetes survive with plant refuse. Parasitic phanerogams survive in the form of seeds
and as eggs, cysts and larvae of parasitic nematodes serve as overseasoning structures.
2. Survival as saprophytes
The ability to live saprophytically enables many plant pathogens to survive in the absence
of growing susceptible plants. Saprophytic survival usually occurs in the soil. Waksman (1971)
distinguished between soil inhabitants soil invaders; the former comprise the basic fungal flora
of the soil, whereas the latter are short-lived exotics. As applied to the root infecting fungi soil
inhabitants are unspecialized parasites with a wide host range that are able to survive-indefinitely
in the soil as saprophytes; soil invaders (root inhabiting fungi) are more specialized parasites that
survive in soils inclose association with their hosts. Most plant pathogenic fungi and bacteria are
soil invaders, but some pathogens, notably Rhizoctonia solani and Pythium debaryanum that
cause seedling blights and root rots, live saprophytically in the soils.
The microbiological balance in the soil markedly affects the survival of saprophytic plant
pathogens there. Apparently, Sanford (1926) was the first to suggests that control of potato scab
by green manuring with grass might be due antagonistic action of saprophytic organisms
flourishing on the green manure. Not only do soil saprophytes antagonize other microorganisms
by toxic but also some such as Trichoderma Iignorum actually parasitize Rhizoctonia solani and
other soil-borne pathogens. Despite antagonism and parasitisim by other organisms, many plant
pathogens survive in the soil as inhabitants or invaders. The special conditions that favour
biological control of plant pathogens in sterilized soil or in culture are nonexistent in field soil, in
which there is a complex microflora and a low concentration of nutrients.
Certain plant pathogens survive between growing seasons as saprophytes in the dead
tissues of susceptible plants. Such organisms are only incidentally associated with the soil, and
live only as long as tissues of susceptible plants are available to supply nutrients. Most plant-
pathogenic bacteria and many specialized parasitic fungi survive in this manner. The apple scab
pathogen (Venturia inaequalis) lives parasitically in leaves and fruits during the growing season,
but becomes saprophytic in fallen leaves. Perithecia form in these leaves during the winter, but
ascospores do not form until spring. Ascospores of certain other ascomycetes mature during the
winter, but are protected from adverse conditions by perithecial walls. Soil inhabitants include
obligate saprophytes and facultative parasites and they are exo - pathogens. Whereas soil
invaders (root inhabitants) include facultative saprophytes and obligate parasites and they are
endopathogens (root infecting fungi).
Plant pathogenic bacteria can saprophytically survive or actively multiply in the
rhizosphere or rhizoplane of healthy host and non-host plants. Erwinia carotovora subsp.
carotovora has been considered to survive in soil. However, some recent studies have shown that
soft rot Envinia cannot persist for a long time in fallow soil. E. carotovora subsp. carotovora
multiplies in the rhizosphere of many cruciferous plant species, where the population can readily
increase from 102 cells/g in fallow soil to 104 to 106 cells/g in soil subjected to the rhizosphere
effect of chinese cabbage. Pseudomonas glumae, the causal agent of bacterial grain rot of rice,
remains on rhizosphere and / or rhizoplane of the rice plant from germination to tillering stage.
Burkholderia solanacearum and species of Agrobacterium are best known with a prolonged soil
phase, which can be regarded as the true soil-borne pathogens.
3. Survival in vital association with living plants
Survival of the plant pathogens in vital association with living plants is grouped into
a. Seed
The pathogen of loose smut of wheat, Ustilago nuda tritici, enters the stigma and style
and infects the young seed, in which it survives as mycelium. The seed-infecting pathogens that
cause loose smut of wheat and loose smut of barley are strikingly different from other smut fungi
that attack cereal crops. Most of the others survive from season to season either in non-
pathogenic association with seed or as spores in the soil. Colletotrichum lindemuthianum, the
causative organism of bean anthracnose, can also infect the seed; unless the seed is killed, the
fungus in newly sprouted bean seedlings initiates new infections. The bacteria that cause bean
blights and bacterial blight of cotton survive the winter in infected seed. In Mexico, the fungus of
late blight of potatoes (Phytophthora infestans) produces oospores but in colder regions of the
world, the fungus overwinters as mycelium in diseased tubers.
Lenticels of potato tubers may carry soft rot Erwinia at the maximum level of 100
cells/lenticel, although the infested tubers do not necessarily develop soft rot in the field.
Examples of the plant pathogenic bacteria that survive in seed/planting materials are given in the
following Table 1.
Table 1. Plant pathogenic bacteria surviving in seeds and planting materials.
SI. Disease Bacteria
No
a. Seed
1. Bacterial canker of tomato Clavibacter michiganensis subsp
michiganensis.
2. Bacterial brown stripe of rice Pseudomonas avenae
3. Bacterial blight of cotton Xanthomonas axonopodis pv.
malvacearum
b. Planting material
3. Ring rot of potato Clavibacter michiganensis subsp.
sepedonicus
4. Bacterial wilt of carnation Pseudomonas caryophylli
5. Bacterial wilt of potato and Burkholderia solanacearum
ginger
b. Collateral hosts
Collateral hosts are those, which are susceptible to the plant pathogens of crop plants and
provide adequate facilities for their growth and reproduction of these pathogens. Weeds, which
survive and live during non-cropping season provide for the continuous growth and
multiplication of the pathogen. For example, the fungal pathogen for blast disease of rice,
Pyricularia oryzae can infect the grass weeds like Bracguarua mutica, Dinebra retroflexa,
Leersia hexandra, Panicum repens, etc., and survive during off-season of rice-crop. As soon as a
fresh rice crop is raised, the conidia (inoculum) liberated from the weed host disseminated by
wind infect the fresh rice crop. Thus the weed hosts help to bridge the gap between two rice
crops. Hence the pathogen can be able to line continuously in the vicinity on these hosts inspite
of the non-cropping period intervening between two cropping periods. Intensive cultivation of
particular crop repeatedly and constantly also provides perpetual inoculum. Powdery mildew and
viral diseases of cucurbits are also best examples, where, number of cultivated crops serves as
collateral hosts.
The survival of the plant pathogens on collateral hosts/ alternative hosts which include
the weed hosts also. The collateral/weed hosts which are present in the field and in the bunds
harbour the plant pathogens during cropping season. But the collateral/weed hosts present in the
bunds harbour the plant pathogens during off-season. The pathogens can survive in active
sporulating stage on wild collateral hosts and from their wind or insect to primary inoculum.
Plant pathogenic bacteria may be able to disseminate in the parasitic form on annual and
perennial weeds. For example, the long-term survival of Pseudomonas avenae in Florida is
attributed to the association with a perennial grass, vasey grass (Paspalum urvillei), through
repeated infection of its vegetative growth as well as seed transmission. Bacterial leaf blight of
maize may have its origin in the infected vasey grass distributed in the field. This list of
collateral weed hosts for the plant pathogens is given in the Table below:
Table: Collateral hosts (alternative hosts) of plant pathogens
Collateral hosts
Pathogen Disease Principal host
/ Alternative hosts
1. Fungal diseases
Sclerospora Downy mildew Pearlmillet, Echinochloa sp.,
graminicola Fox-tail, Millet Euchlaena sp.,
Panicum sp.
Peronosclerospora Downy mildew Com Euchlaena sp.,
heterogoni Heteropogon sp.
P. sorghi Sorghum downy Corn, Sorghum Andropogon spp.,
mildew Euchlaena sp.,
Heteropogon contortus,
Panicum trypheron
Erysiphe Cucurbit powdery Cucurbits weeds Many cucurbitaceous
cichoracearum mildew
Rhizoctonia solani Web blight Cowpea Amaranthus spinosus, Aspilia
africana
Bacterial diseases
Xanthomonas Bacterial leaf blight Rice Cyperus spp
oryzae ov oryzae Leersia hexandra
Pseudomonad Red stripe anad top Sugar cane Sorghum halepense
rubrilineans rot S. sudanense
X. axonopodis pv. Bacterial blight Cotton Eriodendron anfructuosum,
malvacearum Jatropha curcas
Lochnera curcas
Thurbaria thespesoides
Virus and Phytoplasma diseases
Cucumber mosaic Mosaic Saffflower Amaranthus blitum, datura
virus metel
Physalis minima
Solanum nigrum
Rice tungro virus Rice tungro Rice Oryza spp,
Echinochloa colona,
E. crusgalli,
Leersia hexandra
Bhendi yellow vein Bhendi yellow vein Bhendi Hibiscus tetraphyllus
mosaic virus mosaic virus
Tomato spotted Ring mosaic Groundnut Bidens pilosa
wilt virus Tagetes sp.
Phytoplasma Datura sp.,
Little leaf BrinjaI
Catharanthus sp.
c. Alternate hosts
The role of alternate hosts is not important as of collateral hosts. However, when a
pathogen has very wide host-range and is tolerant to wide range of weather conditions the
alternate hosts become very, important source of survival of the pathogen. These alternate hosts
are very important for the completion of the Iifecycle of heteroecious rust pathogens. e.g. in
temperate regions the alternate host Berberis vulgaris of Puccinia graminis tritici (black/stem.
rust pathogen on wheat), the barberry bush, grows side by side with the cultivated host, wheat. In
such areas this wild host belonging to a different family is important for survival of the fungus. It
helps in completion of heterogeneous infection chain of the rust fungus. The list of alternate
hosts for important plant pathogenic survival is given in the Table 4.
Table 4. Alternate hosts of plant pathogens
Fungal pathogen Disease Primary host Alternate host
Puccinia graminis Stem rust / Black Wheat Barberis vulgaris
tritici rust
Puccinia recondita Leaf rust / Thalictrum flavum
Brown rust/
Orange rust
Puccinia coronata Crown rust Oat Rhamnus

Puccinia anomala Brown rust Barley Lily
Puccinia dispersa Brown rust Rye Anchusa sp
Puccinia purpurea Rust Sorghum. Oxalis corniculata
Puccinia substriata Rust Pearl- millet Brinjal
var. penicillariae
Puccinia sorghi Leaf rust Maize Oxalis corniculata
Cronartium ribicola Blister rust Currant Pine
Gymnosporangium Cedar rust Cedar Apple
juniperi-virginianae
d. Self sown crops

Self-sown plants, voluntary crops and early sown crops are reservoirs of many plant
pathogens e.g., groundnut rust pathogen, Puccinia arachidis and ring mosaic of groundnut
caused by tomato spotted wilt virus. Self-sown rice plants harbour the pathogen as well as
vector. e,g., rice tungro virus and its vector, Nephotettix virescens.
Perennial crops also playa major role in the survival of the plant pathogens. Pathogens,
which infect perennial plants, can survive in them during low winter temperature and/or during
the hot, dry weather of the summer. They survive in the lesions on perennial host plants, which
may be actively growing or are donnant. Disease like bunchy top of banana survives
continuously in the suckers produced by the mother plants. In citrus canker the bacterium,
Xanthomonas axonopodis pv. citri survives in the cankers formed on the leaves and twigs of the
standing tree. They mostly survive in mild or vigorous active form on hosts, Citrus sp. Other
examples are Erwinia amylovora and Xanthomonas campestris pv. pruni.
e. Ratoon Crops
Sometimes ratoon crop also harbour the plant pathogens e.g., sugarcane mosaic.
f. Survival by latent infection
Latent infection refers to the conditions in which the plant pathogens may survive for a
long time in plant tissue without development of visible symptoms. Eg. Pseudomonas syringae
pv. syringae and X axonopodis pv. citri can survive in apparently healthy bark tissues of their
tree hosts. Xylella fastidiosa, the causal agent of Pierce's disease of grapevine and leaf scorch
disease of various fruits and leaf ornamental trees infect diverse kinds of weeds without
developing visible symptoms. Because these weeds are usually favourable habitats for the vector
insects, latently infected weeds become an important source of the carrier insects.
g. Survival as residents
S.No Disease Bacteria

1. Fire blight of apple and pear Erwinia amylovora
2. Soft rot of chinese cabbage Erwinia carotovora subsp.carotovora
3. Bacterial grain rot of rice Pseudomonas glumae
4. Bacterial blight of soybean Pseudomonas· syringae pv. glycinea
5. Angular leaf spot of cucurbits Pseudomonas syringae pv. lachrymans
6. Bacterial canker of stone fruits Pseudomonas syringae pv.
morsprunorum
7. Bacterial brown spot of bean Pseudomonas. syringae pv. syringae
8. Bacterial speck of tomato Pseudomonas syringae pv. tomato
9. Bacterial blight of cotton Xanthomonas axonopodis pv.
malvaceqrum
10. Bacterial blight of cassava Xanthomonas axonapodis pv.manihotis
11. Common blight of bean Xanthomonas campestris pv. phaseoli
Plant pathogenic bacteria have the capacity to grow on the surface of host and non-host plants
utilizing the small amount of nutrients that are secreted on the plant surface. Survival as residents
in the phyllosphere by bacteria is given below:
4. Survival in association with nematodes and fungi
Plant viruses like wheat mosaic, wheat spindle streak virus, lettuce big vein, tobacco
necrosis, tobacco rattle and tobacco ring spot viruses survive with nematodes or fungi found in
the soil between crop seasons. Tobacco ring spot virus is associated with the nematode,
Xiphinema americanum. The fungus, Polymyxa graminis (Barley yellow mosaic, oat yellow
mosaic, wheat soil-borne mosaic, wheat spindle-streak mosaic) and Spongospora subterranea
(potato mop top) carry the viruses internally and transmit.them through their resting spore.
Viruses are retained by nematode vectors for long times (stable). Xiphinema sp. retained viruses
for a considerable Iength of time, while Longidorus spp and Trichodorus spp. retained them for
a much less period of one or two months only.

5. Survival in association with insects
Many insects are carriers of inocula during the growing season and several important
plant pathogens survive between growing seasons within insects. Some bacterial plant pathogens
may survive within the insect body and over winter therein. The com flea beetle, Chaetocnema
pulicaria Melsh carries inside its body, the com wilt pathogen, Xanthomonas stewartii and thus
helps in its overwintering. The cucumber beetles, Diabrotica vitata Fabr. and D.
duodecimpunctata Oliv., which chew the plant parts affected by Erwiniatra cheiphila carry the
pathogen inside their body, where it over winters. In the following seasons the insect passes on
the bacterial pathogen to the host plant.
These insect vectors effectively transmit the bacterial pathogen causing wilt of cucurbits.
It is reported that the bacterial pathogen and the -insect vectors live in a symbiotic relationship,
the insects helping the bacterium with protection from adverse weather conditions and the
bacterium helping the insects with a supply of some digestive enzymes while it is inside the
insect's body. Plant viruses and phytoplasmas multiply within the vectors and can overwinter in
those insects. Semi-persistent viruses are retained in the vectors for periods ranging from hours
to days. Example, citrus tristeza virus is retained in the aphid Toxoptera Citricida. Persistent
viruses retain the viruses from days to week. Most of the hopper borne viruses multiply in their
vectors. Viruses are retained through the moult and the vectors frequently remain viruliferous for
life important. Vectors, which retain the viruses, are given below:
Leaf hopper transmitted viruses
Vector Virus
Leaf hopper Circulifer tenellus Beet curly top virus
Plant hopper Cicadulina mbila Maize streak virus
Green leaf hopper Nephotettix cincticeps Rice dwarf virus
Brown plant hopper Nilaparvata lugens Rice grass stunt virus
Hopper Agallia constricta Wound tumour virus
Transovarial transmission of the virus to the eggs of the vectors occurs and the virus can
multiply within a viruliferous hopper even if the insect is feeding on an inimune host plant. Eggs
carrying viruses may overwinter and provide a source of virus to infect spring crops, even in the
absence of diseased plants. Phytoplasmas attacking plants also multiply in the insects and remain
infective throughout their life period. e.g. Rice dwarf virus (RDV) is transmitted through the
eggs to about 60% of the progeny of the infective female leafhopper, Nephotettix cincticeps.
RDV passes through the eggs to six succeeding generations. Clover club leaf is transmitted
through 21 generations of the leafhopper vector, Agalliopsis novel/a over a span of five years.
6. Survival on agricultural materials
Clavibacter michiganensis subsp. michiganensis has been shown to survive in air-dried
conditions for 7 to 8 months on the surface of wooden stakes and boxes or wires or for 15
months in air-dried tissues of diseased· tomato plants.
7. Survival on surface water
Erwinia carotovora subsp. carotovora is detected from water from drains, ditches,
streams, rivers and lakes in mountainous upland and arable areas of Scotland and Colorado
throughout the year.
Dispersal of Plant Pathogens
Transport of spores or infectious bodies, acting as inoculum, from one host to another
host at various distances resulting in the spread of disease, is called dissemination, dispersal or
transmission of plant pathogens. It is very important for spread of plant diseases, for continuity
of the life cycle and evolution of the pathogen. The spores of some fungi are expelled forcibly
from the sporophore or sporocarp by a squirting or puffing action that results in successive or
simultaneous discharge of spores up to a centimetre or so above the sporophore.
The seeds of some parasitic plants are also expelled forcibly and may arch over distances
of several metres. These are dispersed mechanically by various means. In bacterial diseases, the
bacterial cells come out on the host surface as ooze or the tissues may be disintegrated so that the
bacterial mass is exposed and then dispersed by various physical and biological agencies.
Insects, mites, phanerogamic parasites nematodes and human beings transmit viral diseases,
which have no such organs.
The knowledge of these methods of dispersal is essential for effective control of plant
diseases because possibilities of preventing dispersal and thereby breaking the infection chain
exist.
The dispersal of infectious plant pathogens occurs through two ways,
I. Autonomous or direct or active dispersal
II. Indirect or passive dispersal

I. Autonomous dispersal
It is also known as active or direct dispersal. In this method the dispersal of plant
pathogens (fungi, bacteria, and viruses) takes place through soil and seed or planting materials
during normal agronomic operations.
1. Soil as means of autonomous dispersal
Soil-borne facultative saprophytes or facultative parasites may survive through soil. The
dispersal may be by movement of the pathogen in the soil or by its growth in soil or by
movement of the soil containing the pathogen. The former is known as dispersal in soil while the
latter is called dispersal by soil.
a. Dispersal in soil
The following are the three stages of dispersal in soil.
i. . Contamination of soil
ii. Growth and spread of the pathogen in soil
iii. Persistence of the pathogen
i. Contamination of soil
Contamination of the soil takes place by gradual spread of the pathogen from an infested
area to a new area or by introduction of contaminated soil, plant debris to a new area or by
introduction of infected seed or planting materials.
ii. Growth and spread of the, pathogen in soil
Once the pathogen has reached the soil it can grow and spread "depending on the
multiplication and spread. Multiplication and spread depends on the characters of the pathogen,
presence of susceptible host and cultural practices. The adaptability of the pathogen to the soil
environment includes saprophytic survival ability. The survival ability of the pathogen is
governed by high growth rate, rapid spore germination, better enzymatic activity, capability to
produce antibiotics and tolerance to antibiotics produced by other soil microorganisms. The
active saprophytic survival of facultative saprophytes and facultative parasites in soil is affected
by soil structure, moisture, organic matter, pH; antagonism etc., Specialized facultative parasites
(or saprophytes) can pass their life in soil in the absence of the hOst plants, but they depend more
on the residues of their host plant. e.g., Armillariel/a mel/ea, Ophiobolus graminis,
Phymatotrichum omnivorumand Fusarium. The non-specialized facultative parasites can pass
their entire life in the soil. e:g., Pythium sp., Phytophthora sp., The soil-borne obligate parasites.
such as Plasmodiophora brassicae, Synchytrium endohioticum requires the presence of active
host.
Ill. Persistence of the pathogen
The pathogens persist in the soil as dormant structures like oospores (Pythium;
Phytophthora, Sclerospora, etc.) chlamydospores (Fusarium) or smut spores (Ustilago) or
sclerotia (Rhizoctonia, Sclerotium, etc.)
b. Dispersal by the soil
The pathogen enters the soil, grow and spread in the soil. During the cultural operations
in. the field, soil is moved from one place to the nearby place through the agricultural
implements and irrigation, worker's feet. Propagules of fungi or the dormant structures of fungi
and the plant debris containing the fungal and bacterial pathogens thus spread throughout the
field.
2. Seed and seed materials as the source of autonomous dispersal
The seeds serve as medium for autonomous dispersal of pathogens. Since most of the
cultivated crops are raised from seed the transmission of diseases and transport of pathogens by
seeds has much importance. The dormant structures of the pathogen (e.g., seeds of Cuscuta,
sclerotia of ergot fungus, smut sori, etc.,) are found mixed with seed lots and they are dispersed
as seed contaminant. The bacterial cells or spores of fungi present on the seed coat (such as in
smuts of barley, sorghum, etc.,) are transported to long distances. Dormant mycelium of many
fungi present in the seed is transmitted to long distances.
This type of dispersal is highly erratic. The most important methods of dispersal of
pathogen by the soil are transfer of soil from one place to another along with plant parts or
propagating materials. e.g., transfer of papaya seedlings from a nursery infested with Pythium
aphanidermatum (the cause of stem or foot rot of papaya) C311 introduce the pathogen in new
pits for transplanting the seedlings. Similarly grafts of fruit trees transported with soil around
their roots can transmit pathogens present in the nursery to the orchards. By this method,
pathogens are not only spread from field to the field but also from district to district, State to
State and often from country to country. There are three types of dispersal by seed viz.,
a. Contamination of the seed
b. Extemally seed - bome, and
c. Intemally seed - bome

Contamination of the seed
Seed -borne pathogens move in seed lot as separate contaminants without being in
intimate contact with the viable crop seeds. The seeds of the pathogen or parasite and the host are
getting mixed during harvest of the crop. In many cases, the identity ofthe seeds of the two
entities (host and the parasite) is difficult to separate. e.g., smut of pearl millet (Tolyposporium
penicillariae), ergot of rye and pearlmillet (Claviceps purpurea and C. jusiformis respectively).
Smut soil and ergots mix easily with the seed lots during harvest or threshing. In many smuts
such as Kamal bunt of wheat (Neovossia indica) and bunt of rice (Neovossia horrida) the
infected kernels containing smut sori are mixed with the seed. In leaf smut of rice (En tyloma
oryzae) leaf pieces containing smut sori are mixed with the seed.
Externally seed-borne
Close contact between structure of the pathogen and seeds is established in
diseases like covered smut and loose smut VI Daney, snoll smut ot sorghum, stinking smut of
wheat and bacterial blight of cotton where the pathogen gets lodged in the fomi of donnant
spores or bacteria on the seed coat during growth of the crop or at the time of harvest and
threshing. In many pathogens the externally seed-borne Structures such as smut spores can
persist for many years due to their inherent capaCity for long survival. The spores of Tilletia
caries (stinking smut of wheat) remain viable even after 18 years and those of Ustilago avenae
(oat smut) for 13 years.
Internally seed-borne
The pathogen may penetrate into the ovary and cause infection of the embryo while it is
developing. They become internally seed-borne. Internally seed borne pathogens like Usti/ago
nuda tritici are viable for more than 15 years. Other examples include Helminthosporium oryzae,
Sclerospora graminico/a, etc. The bacterial pathogens include Xanthomonas oryzae pv. oryzae
on rice, Pseudomonas syringae pv. syringae in cucurbits, Xanthomonas campestris pv.
campestris on crucifers, etc.
Mainly man distributes seeds of cultivated crops. Sometimes animals and birds also help
in distribution of crop seeds. Man and animals are the main agencies of dispersal of pathogen
through seed. The pathogens thus mixed with the seed or on the seed are transmitted.
Passive dispersal
Passive dispersal of plant pathogens happens through
I. Animate agents
a. Insects
b. Mites
c. Fungi
d. Nematodes
e. Human beings
f. Farm and wild animals
g. Birds
h. Phanerogamic parasites
II. Inanimate agents
a. Wind
b. Water
1. Animate agents
a. lnsects
Insects carry plant pathogens either externally or internally. Gaiiman (1950) used the
terms epizoic and endozoic respectively for these two types of transmission. The external
transmission of plant pathogens is of special interest in those fungi, which produce their conidia,
oidia and spermatia in sweet or honey secretions having attractive' odours. Some Qfthe well
known diseases of this type are the ergot, the Sc1erotinia brown rot of pear and apple, the honey
dew stage in the 'sugary disease' of sorghum and pearlmillet in parts of India and the pycnial
nectar in the cluster cup stage of rusts. The spermatial oozing at the mouth of spermagonia in the
Ascomycetes attract various types of insects, flies, pollinating bees and wasps which playa dual
role viz., pollination and transmission' of pathogens. The fire blight organism (Erwinia
amylovora) pathogens and citrus canker bacterium, (Xanthomonas axonopodis pv. cirri) are also
carried in this manner, the former by flies (bees) and ants and the latter by the leaf miner. The
black leg of potato caused by Erwinia carotovora is disseminated by maggots, wilt of com
caused by X. stewartii, gummosis of sugarcane caused by X vasculorum are the other examples
for bacterial diseases transmitted by insects.
Ingenious transmission of pathogens, of an internal nature (endozoic) is provided by the
Dutch elm disease (Ceratostomella ulmi) and the olive canker (Bacillus savastano i). The former
is transmitted by the elm bark beetles and the latter by the olive fly (Olea . europaea). These
insects, unlike the epizoic group, appear to have a close biologic relationship with the pathogens,
as they have not been reared without the contaminating pathogens.
Insects spread few important plant pathogenic bacteria. The cucumber wilt bacterium,
Erwinia tracheiphila is spread by the striped cucumber beetles (Acalymma vitata) and the
spotted cucumber beetle (Diabrotica undecimpunctata). When the beetles are feeding on the
diseased plant, the bacterium contaminates the mouthparts and passes into the gut of the insect.
During the winter season, the bacterium overwinters inside the beetle. Thus the beetle helps the
bacteria in two ways, i.e. in their transmission and survival.
Different types of insects spread more than 80 per cent of the viral and phytoplasmal
diseases. The insect, which act as specific carriers in disseminating the diseases, are called insect
vector.
Both aphids (Aphidae) and leafhoppers (Cicadellidae or Jassidae) in the order Homoptera
contain largest number and the most important insect vectors of plant viruses. Certain species of
mealy bugs and scale insects (Coccoidae), whiteflies (Aleurodidae) and treehoppers
(Membracidae) in the same order (Homoptera) also transmit virus diseases. Insect vectors of
plant viruses are few in true bugs (Hemiptera), thrips (Thysanoptera),beetles (Coleoptera) and
grasshoppers (Orthoptera). Aphids, leafhoppers and other groups of Homoptera and true bugs
have piercing and sucking mouthparts. Thrips have rasping and sucking mouthparts. All other
groups of insect vectors have chewing mouthparts and they transmit only very few viruses.
Aphids
Aphids are the most important insect vectors of plant viruses and transmit the great
majority of all stylet - borne viruses. As a rule several aphid species can transmit the same stylet
- borne virus and the same aphid species can transmit several viruses, but in many cases the
vector-virus relationship is quite specific. Aphids generally acquire the stylet-borne virus after
feeding on a diseased plant for only a few seconds (30 seconds or less) and can transmit the virus
after transfer to and feeding on a healthy plant for a similarly short time of a few seconds. The
length of time aphids remain viruliferous after acquisition of a stylet-borne virus varies from a
few minutes to several hours, after which they can no longer transmit the virus. In few cases of
aphid transmission of circulative viruses, aphids cannot transmit the virus immediately but must
wait several hours after the acquisition feeding, but once they start to transmit the virus, they
continue to do so for many days following the removal of the insects from the virus source. In
aphid transmitting stylet-borne viruses, the virus seems to be borne on the tips of the stylets, it is
easily lost through the scouring that occurs during probing of host cells, and it does not persist
through the moult or egg. The examples of aphid transmitted plant viruses are given in the
following Table.
S.No Virus Vector Type of transmission
1. Bean common Mosaic Acyrthosiphon pisum Non persistent
2. Bean yellow mosaic A. pisum Non - persistent
3. Citrus tristeza Toxoptera citricida Non - persistent
4. Pea enation mosaic A. pisum Persistent
5. Beet yellows M. Persicae Semi persistent
Leaf hoppers
Leaf hoppers are phloem feeders and acquire the virus from the phloem region. All leaf
hoppers, transmitted viruses are circulatory. Several of these viruses multiply in the vector
(propagative) and some persists through the moult and are transmitted through the egg stage of
the vector. Most leaf hopper vectors require a feeding period of one to several days before they
become viruliferous, but once they have acquired the virus they may remain viruliferous for the
rest of their lives. Usually there is an incubation period of 1 to 2 weeks between the time a leaf
hopper requires a virus and the time it can transmit it for the first time.
b. Mites
Mites belonging to the families eriophyidae (eriophid mite) and tetranychidae (spider
mite) transmit plant viruses.
c. Fungi
C. Fungi
Some soil - borne fungal plant pathogens transmit plant viruses. Olpidium brassicae,
Ploymyxa graminis, P. betae and Spongospora subterranea are the fungi involved in
transmission of virus disease. The viruses are apparently borne in or on the resting spores and the
zoospores, which upon infection of new host plants introduce the virus and cause symptoms
characteristic of the virus they transmit. All these fungi are pathogens of the host, which carry of
viruses. The zoospores of the fungi are released from the host and the zoospores carry the virus
and transmit it to the susceptible hosts during their infection process. In some cases plant viruses
are carried on the outside of the fungi. Examples are tobacco necrosis virus and cucumber
mosaic virus.
The viruses like lettuce big vein virus are found inside the zoospores. They persist for
years in viable resting sporangia. The types of transmission by fungi can be considered as non-
persistent and persistent transmission. The list of fungi and the virus diseases transmitted by
them are given in the following table.
S.No Fungal vector Disease

1. Olpidium brassicae Lettuce big vein, Lettuce necrosis,
Tobacco stunt, Tobacco necrosis
satellite
2. Synchytrium endobioticum Potato virus
3. Spongospora subterranea Potato mop top
d. Nematodes
Nematodes are soil borne organisms. Some of the nematodes act as agents for
dissemination of pathogenic fungi, bacteria and viruses. For example, the bacterium
Corynebacterium tritici that causes yellow ear rot of wheat is disseminated by ear cockle
nematode. Similarly, some pathogenic fungi such as, Phytophthora, Fusarium, Rhizoctonia, etc.,
are carried on the body of nematodes. Nematodes help these pathogenic fungi to enter into the
host through punctures for their own entry and enter into hosts along with the nematodes. Plant
nematodes play a vital role as vector in transmitting certain virus diseases. Nematode vectors
transmit viruses by feeding on roots of infected plants and then moving on roots of healthy
plants. Larvae as well as adult nematodes can acquire and transmit viruses, but the virus is not
carried through the larval molts or through the eggs. After moulting, the larvae or the resulting
adults must feed on a virus source before they can transmit again. Xiphinema, Longidorus and
Trichodorus transmit both the polyhedral and tubular type of viruses. The important viral
diseases transmitted by nematodes are given below:
Virus group Virus Vector

Tobra virus (Tobacco rattle Pea early browning, Paratrichodorous sp.
group virus) Tobacco rattle Trichodorous spp
Nepo virus (Nematode Grapevine chrome virus Xiphinema index
transmitted polyhedral Tobacco Ringspot X. americanum
virus) Tomato ringspot X. americanum
e. Human being
Man is the most important factor responsible for.' short distance and 'long distance
dispersal of plant pathogens. He helps in dissemination unknowingly by his usual agricultural
practices. Human being's role in dissemination of plant pathogens is more direc of plant
pathogens by human beings is known anthropochory. The ways and means by which human
beings help in dispersal are as follows.
i. Transportation of seeds (Seed trade)
Seed trade is one of the different means of dispersal of plant pathogens in which man
plays an important role. The import and export of contaminated seeds without proper precautions
lead to movement of pathogens from one country to another or from one continent to another.
Through this way pathogens of soybean and sugarbeet hither to not prevalent in India got
introduced. Human agencies of individual, official and unofficial have transported new plants
and plant products, the seed, the tubers, the propagating stock and fruits, which carried the plant
pathogens, many times in a latent condition and which ultimately lead to the outbreaks of new
diseases in places, hither to free from them. The diseases which are amenable to such
transmission are mainly those that are carried in or on the propagative parts and seed such as late
blight of potato, the downy mildew of grapevine, citrus canker, chestnut blight, Dutch elm
disease, Fusarium wilt of banana, Katte disease of cardamom and buncilY top of banana. A few
such diseases together with their places of origin and years and introduction are given in Table
below:
Disease Original home Introduced Year of
country introduction
Citrus canker Asia USA 1907
Fireblight of apple USA New Zealand 1919
Powdery mildew of USA Europe 1845

grape vine
Downy mildew of USA France 1878
grapevine
Late blight of potato South America USA 1830
Panama disease of Panama Islands Bonbay 1920
banana
Bunchy top of Sri Lanka South India -
banana
Many of these diseases, not very destructive in their homelands, have brought in ruin and
devastation. The sale of seeds for crops badly affected by a seed-borne pathogen is a common
method of dispersal of destructive pathogens e.g. loose smut of wheat (Ustilago nuda tritici),
grain smut of sorghum (Sporisorium sorghi), ergot of pearl millet (Claviceps fusiformis) and
Kamal bunt of wheat (Neovossia indica).
ii. Planting diseased seed materials (vegetatively propagated materials)
Planting diseased bulbs, bulbils, corns, tubers, rhizomes, cutting etc. of vegetatively
propagated plants such as potato, sweet potato, cassava, sugarcane, banana, many ornamentals
and fruit trees etc. help in dispersal of pathogens from field to field, orchard to orchard, locality
to locality or from one country to another.
iii. By adopting farming practices
Human beings (men and women) engaged in preparatory cultivation, planting, irrigation,
weeding, pruning etc. help in dispersal of plant pathogens. The fungal spores (oospores,
chlamydospores), dormant structures like sclerotia are carried by worker's clothing, shoes, hand
etc. from field to field. Men or women engaged in intercultivation in tobacco field spread the
dreaded tobacco.
iv. Through clothing
Palm workers engaged in cleaning coconuttrees spread bud rot disease.
v. By use of contaminated implements
Pathogens are transferred from one area to another through implements used in various
cultural operations (weeding, hoeing thinning etc.) in the field. e.g: root rot of pulses and cotton
(Macrophomina phaseo/ina, bacterial angular leaf spot of cucumber (Pseudomonas /achrymans)

and bacterial canker of tomato (Corynebacterium michiganensis). Cutting knives and pruning
knives help in dissemination from one plant to another e.g., Bunchy top of banana.
vi. By use of diseased grafting and budding materials
Grafting and budding between healthy and diseased plants is the most effective method
of distribution of pathogens of horticultural crops (fruit trees, ornamentals etc.) e.g., Careless
selection of stocks and scions in propagation of citrus trees.
f. Farm and wild animals
Farm animals (cattles) while feeding on diseased fodder ingest the viable fungal
propagules (spores or oospores or sclerotia) into their digestive system. Animals which feed on
downy mildew affected pearlmillet or sorghum take the oospores along with the fodder.
Oospores pass out as such in the dung. This dung when used as manure spread in the field and
act as source of inoculum. Smut fungi like grain smut of sorghum, loose smut and head smut of
sorghum are carried from field to field through the alimentary canals of farm animals. Soil
inhabiting fungi especially sclerotia adhere to the hoofs and legs of animals and get transported
to other places. Animals passing through the tobacco fields help in transmission of TMV.
g. Birds
In general, transmission by birds is of minor importance. But this method is important in
dissemination of seeds of flowering parasites and certain. fungi. Many migratory birds, such as
mistle thrush (Turdus viscivorus) in the temperate region and the crows (Crovos
brachyrhynchos) in the tropics, take active part in the transmission of giant mistletoe
(Dendrophthoe spp.) either through external contamination of their beaks and feathers or
internally through the alimentary canals. These birds feeding on the fleshy, sticky and gelatinous
berries of giant mistletoe deposit the seeds on the other trees with the excreta.
Stem segments of dodder (Cuscuta spp.) are carried by birds for buildingtheir' nests. Thus
the phanerogamic parasites are getting transported to new locations. Spores of chestnut blight
fungus, Endothea parasitica are disseminated by not less than 18 species of birds. Internal
transmission of this pathogen is carried out by the birds, which visit such diseased plants and get
contaminated by the spores. Birds are also known to carry the spores of fungi on their, body.
h. Phanerogamic parasites
Plant viruses are transmitted from one plant to another through the bridge formed
between the two plants by the twining stems of the parasitic plant dodder (Cuscuta spp). Dodder
is yellow vine without green leaves. In this way viruses are transmitted between plants belonging
to families widely separated taxonomically. The virus is transmitted in the food stream of the
dodder plant, being acquired from the vascular bundles of the infected plant by the haustoria of
dodder. After translocation through the dodder phloem the virus is introduced in the next plant
by the new dodder haustoria produced in contact with the vascular bundles of the inoculated
plant. Cuscuta californica, C. campestris, C. subinclusa are usually employed for dodder
transmission of viruses and phytoplasmas. C. europaea, C. epilinum and C. lupuliformis are also
employed in transmission of viruses.
2. Inanimate agents
a. Wind
The wind dispersal of plant pathogens is known as anemochory. It is one of the most
common methods of the dispersal of plant pathogens. It is the most dangerous and potent mode
of travel for plant pathogenic fungi. It acts as potent carrier of propagules of fungi, bacteria and
viruses. Usually the fungal pathogens are light in weight and are well adapted to wind dispersal.
Some pathogenic bacteria are carried along with the infected material to short distances by wind.
Damping-off pathogen (Pythium spp.), wart disease pathogen of potato (Synchytrium
endobioticum); root rot pathogens (Sclerotium and Rhizoctonia) and seeds of phanerogamic
parasites witchweed (Striga) are efficiently carried by wind. Viruses and phytoplasmas are not
directly transmitted by wind, but the insect and mite vectors that carry the'viruses move to
different directions and distances depending upon the direction and speed of air.
The adaptations for wind dispersal in fungal pathogens include, production of numerous
spores and conidia, discharge of spores with sufficient force, production of very small and light
spores so that they can move to long distances. The duration and periodicity of sporulation and
discharge are also important factors for wind dispersal. Some fungal pathogens causing powdery
mildews, downy mildews, rusts, smuts, sooty moulds, leaf spots, blast, apple scab etc., produce
large number of very light spores and conidia on the surface of the host. Uredial stages of the
rust fungi travel long distances through air currents and are thus responsible for destructive
epidemics over wide areas.
Wind transmission involves the upward air currents, velocity and the downward
movements of wind. All are equally responsible for the spread of infection and ultimate outbreak
of diseases and have been of special significance in the rust, smut and t>last fungi. Uredospores
of rust fungi have been carried to long distances, both cross-wise and upwards. Christensen
(1942) and Stakman (1946) determined by exposure of Vaseline slides in the upper air through
aeroplane flights, that uredospores and" aeciospores of Puccinia gramminis tritici could be
gathered in a viable condition up to a distance of 4,200 m, above infected fields, Alternaria sp. at
2,400 m. and those of Puccinia tritieina at 3,750 m. The transmission of aecial spores of
Puceinia graminis trWei from several groups of barberry bushes to the wheat crop showed that
these spores traveled successfully over a radius of 3 kms round about these bushes. The blister
rust fungus, Cronartium ribicola, is known to travel to a distance of 500 metres or 3,750 m.
inside a plantation that the range is probably more in the open. Similar observations have been
made in respect of dissemination of chlamydospores ofthe smut fungi.
In long distance dissemination with intervening stages of infection, the retention of
viability of spores is an important factor that determines the extent and severity of epidemics,
over wide areas. The outbreaks of cereal rusts and blast of rice are examples of such
dissemination. Spores differ widely in their ability to survive long distance travel through air.
Uredospores of rusts, chlamydospores or smut fungi and conidia of Alternaria,
Helminthosporium, Pyricularia and others are well adapted for long distance travel in a viable
condition and are known to play a vital role in epidemiology. The conidia of downy mildews,
powdery mildews and the aeciospores and basidiospores of the rust fungi are unable to withstand
such long distance dissemination when they are exposed to desiccation and direct sunshine and
thus are only capable of producing local epiphytotics of limited magnitude.
The bacteria causing fire blight of apple and pear (Erwinia amylovora) produce fine
strands of dried bacterial exudates containing bacteria and these strands may be broken off and
they are transmitted by wind. Bacteria and nematodes present in the soil-may be shown away
along with soil particles. Wind also helps in the dissemination of bacteria, fungal spores and
nematodes by blowing away rain splash droplets containing these pathogens. Wind carries
insects and mites that may contain _ or are smeared with viruses, bacteria or fungal spores to
short or long distances. Wind also causes adjacent plants or plant parts to rub against each other.
The wound created in this manner help the spread by contact of bacteria (citrus canker), fungi,
some viruses (Tobacco mosaic virus) and viroids and possibly of some nematodes.
b. Water
Transmission of plant pathogens by water (hydrochory as called by Gaiimann, 1950) is
not as significant as wind transmission. Although water is less important than air in long-distance
transport of pathogens, water dissemination of pathogens is more efficient, in that the pathogens
land on an already wet surface and can move or germinate immediately. In case of some diseases
the surface flow of water after heavy showers of rains or irrigation water from canals and wells
carries the pathogens to short distances. Soil inhabiting fungi like, Fusarium, Ganoderma,
Macrophomina Phytophthora, Plasmodiophora, Pythium, Rhizoctonia, Sclerotium, Sclerotinia,
Sporisorium, Ustilago, Verticillium etc., in the form of mycelial fragments, spores or sclerotia,
soil-bone bacteria and nematodes carrying viruses are transmitted through the above process.
They are transmitted through rain or irrigation water that moves on the surface or through the
soil.
All bacteria and the spores of many fungi are exuded in a sticky liquid and depend for
their dissemination on rain or (overhead) irrigation water, which either washes them downward
or splashes them in all directions.
Raindrops or drops from overhead irrigation pickup the fungal spores (uredospores of
Hemileia, Puccinia and Uromyces and bacteria (bacterial blight pathogen of rice, Xanthomonas
Olyzae pv. oryzae; bacterial leaf streak pathogen, X oryzae pv. translucens; citrus canker
pathogen, X axonopodis pv. citri; tomato bacterial blight pathogen, Ciavibacter michiganensis
and cotton bacterial blight pathogen, X.axonopodis pv. malvacearum present in the air and wash
them downward where some of them may land on susceptible plants.

Survival and Dissemination of Plant Pathogens

Uploaded by

Copyright:

Available Formats

Survival and Dissemination of Plant Pathogens

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Survival and Dissemination of Plant Pathogens

Uploaded by

Copyright:

Available Formats

Plant Pathogens & Principles of Plant Pathology

Puccinia coronata Crown rust Oat Rhamnus

d. Self sown crops

S.No Disease Bacteria

a much less period of one or two months only.

II. Indirect or passive dispersal

c. Intemally seed - bome

S.No Fungal vector Disease

Virus group Virus Vector

Powdery mildew of USA Europe 1845

(Macrophomina phaseo/ina, bacterial angular leaf spot of cucumber (Pseudomonas /achrymans)

You might also like

Pfad - The Proxy pFad of © 2024 Garber Painting. All rights reserved.