Professional Documents
Culture Documents
net/publication/370036654
CITATIONS READS
0 506
4 authors, including:
Ravikant Singh
Swami Vivekanand University
55 PUBLICATIONS 48 CITATIONS
SEE PROFILE
All content following this page was uploaded by Ravikant Singh on 15 April 2023.
Chapter 4
GROWTH AND SURVIVAL OF PLANT PATHOGENS
Introduction
The initial step in an infection chain or illness cycle is the means of survival.
Primary infection and primary inoculum are terms used to describe the initial
infection that results from the sources of pathogen survival (infected host as a
reservoir of inoculum, saprophytic survival outside the host, dormant spores, and
other structures in or on the host or outside the host) in the crop. After the disease
has begun to affect the crop, the spores or other pathogen structures serve as sources
of secondary inoculum and secondary infection, which spreads the disease across
the field.
P. K. Singh (praveensingh4567@gmail.com)
Department of Botany, Swami Devanand Post Graduate College, Math-Lar, Deoria, U. P.,
India
R. C. Vishwakarma
Department of Botany, Buddha Post Graduate College, Kushinagar, U.P., India
J. Kaur
Department of Botany, Dr. Ambedkar Government P. G. College, Unchahar, Raebareli, U.
P., India
R. Singh
Department of Botany, Mahakaushal University, Jabalpur, Madhya Pradesh, India
The fungus that causes late blight on potatoes (Phytophthora infestans) can live in
seed tubers or soil (Anderson, 1951). Primary inoculum (PI), which is present in
the soil, produces primary infection of the crop from healthy seed whereas infected
tubers convey the infection to the field. The PI may also travel vast distances or
from nearby areas by wind. The fungus then spreads spores on leaves. When these
spores touch healthy plant surfaces, they spread by wind and water and start new
diseases. This infection is a secondary one. The disease is brought on by the main
infection, and it is disseminated by the secondary infection.
and multiply. In this way, the weed hosts facilitate the transition between two crop
seasons. For instance, Pyricularia grisea (Teleomorph: Magnaporthe grisea), the
fungus that causes rice blast disease, can persist during the off-season of the rice
crop by infecting grass weeds like Brachiaria mutica, Dinebra retroflexa, Leersia
hexandra, and Panicum repens. The conidia (inoculum) released by the weed host
and spread by wind as soon as a new rice crop is planted infect the new rice crop.
Alternative hosts, or wild hosts from different families: These hosts play a less
significant role than collateral hosts. However when a pathogen has a very broad
host range (like Sclerotium rolfsii, Rhizoctonia solani, Fusarium moniliforme, etc.)
and is adaptable to a wide range of environmental conditions, the disease's alternate
hosts become a crucial part of its survival (Blair, 1942; Abeygunawardena and
Wood, 1957). For the heteroecious rust pathogens to complete their life cycle, these
alternative hosts are crucial. For instance, the barberry bush (Berberis vulgaris), the
alternate host of Puccinia graminis tritici (the pathogen that causes black or stem
rust on wheat), coexists side by side with the cultivated host in temperate climates.
This wild host, which belongs to a separate family, is crucial to the fungus' survival
in such conditions.
Self-sown crops: Early-sown, self-sown, and voluntary crops are all sources of
various plant diseases. As an illustration, self-sown rice plants contain the pathogen
(Rice tungro virus) and the vector (Nephottetix virescens).
[2] Saprophytic survival outside the host: Several plant diseases may exist
without growing vulnerable plants thanks to their capacity for saprophytic life.
Typically, saprophytic life exists in or on the soil. According to Williams (1971),
there is a difference between soil invaders and soil dwellers. The former includes
the fundamental fungal flora of the soil, whereas the latter are transient exotics.
Fungi can exist without the cultivated host plant as saprophytes and fall into one of
three categories for study:
(i) Organisms that live permanently in the soil as saprophytes in the absence of the
host plant are referred to as soil residents. For instance, Pythium, Rhizoctonia, and
Sclerotium species.
(ii) Root occupants: These highly specialised parasites live in soils alongside their
hosts in close proximity. As long as the host tissue in which they are residing as
parasites has not entirely disintegrated, the active saprophytic phase continues.
Examples include cotton root rot, Fusarium species, and Verticillium (fungi that
cause vascular wilt) (Phymatotrichum omnivorum) (Anwar, 1949).
(iii) Rhizosphere colonisers: They are organisms that inhabit the dead substrates
near the roots and remain there for an extended period of time. They are more
resilient to soil antagonists. Example: Cladosporium fulvum, a tomato leaf mould.
[3] Survival as dormant spores or specialized resting structures:
Plant viruses do not have a resting phase and spread continuously along an infection
chain. The phytopathogenic bacteria do not also create dormant spores or other
similar structures. They continue to exist in either their active saprophytic stage on
the remains of dead plants or their active parasitic stage in the current host.
[4] In conjunction with-
(i) Nematodes: They can exist as active parasites that feed on a living host, as well
as dormant forms like eggs, cysts, and galls that grow inside the tissues of the host.
These structures could be found in seed lots or in the soil.
(ii) Phanerogamic parasites: They use seeds to stay latent for a long time. For
instance, Orobanchae seeds can last longer than 7 years in soil.
Fungi are the only organisms among plant pathogens that create spores, which are
similar to nematode eggs, and other resting structures for their inactive survival.
The following categories can be applied to these dormant structures of survival.
Soil borne fungi:
(a) Dormant spores, such as conidia (Taphrina deformans causes peach leaf curl),
chlamydospores (Fusarium species causes wilt) (Anwar, 1949), oospores (Downy
mildew fungi), and perithecia (Venturia inaequalis causes apple scab), etc.
(b) Other dormant components, such as thickened hypha, sclerotia (Cottony Rot
Fungus, Sclerotinia sclerotiorum), microsclerotia (Verticillium), rhizomorphs
(Armillaria mellea), etc.
yellow mosaic) and Spongospora subterranea (Potato mop top virus), which also
spread the viruses through the dormant spore (Baker, 1959).
DISPERSAL OF PLANT PATHOGENS
The spread of plant pathogens is the second link in the infection cycle. Dispersal,
dispersion, or transmission of plant pathogens refers to the movement of spores or
infectious materials functioning as inoculum from one host to another host at
varying distances that causes the disease to spread. The spread of the pathogen or
illness is crucial for the continuation of the pathogen's life cycle and its evolution,
as well as for the spread of plant diseases. As it may be possible to limit dispersal
and so break the chain of infection, understanding these processes of dispersal is
crucial for efficient treatment of plant diseases.
Asexual and sexual spore generation in fungi occurs in response to the fungus's
active vegetative development in or on the host tissues, and both types of spores
are mechanically distributed in both time and space via a variety of techniques. In
bacterial disorders, the bacterial cells manifest as oozing on the host's surface or
they may cause tissue breakdown, exposing the bacterial mass, which is then spread
by a variety of physical and biological agents. Insects, mites, phanerogamic
parasites, nematodes, and people can all transmit viral infections that lack these
organs.
Infectious plant diseases travel via space in one of two ways:
I. Direct, active, or autonomous dissemination.
II. Passive or indirect dissemination.
(I) Autonomous, direct, or active dispersal:
In this technique, plant diseases spread via soil, seed, and planting materials during
routine agronomic activities. With this method of dissemination, external forces
like insects, wind, water, etc. play a minor influence.
(i) Seed as the means of autonomous dispersal: As the majority of cultivated
crops are grown from seed, the movement of pathogens and the spread of illnesses
are significant. The pathogen's dormant structures, such as Cuscuta seeds, ergot
fungus sclerotia, smut, sori, etc., are discovered in seed lots and spread as seed
contaminants. Long distances are travelled by the bacterial cells or fungal spores
that are found on the seed coat, as in the smuts of barley, sorghum, etc. The dormant
mycelium of several fungi found in the seed is spread across great distances. There
are three types of seed dispersal: internal seed borne, external seed borne, and
contaminated seed borne.
(a) Seed contamination: Pathogens that are carried by seeds pass through seed lots
as separate contaminants, avoiding direct contact with crop seeds that are still
viable. During crop harvest, the seeds of the pathogen or parasite and the host are
combined. It can often be challenging to distinguish between the seeds of the host
and the pathogens. Ergot of rye and smut of pearl millet, for instance. During
harvest and threshing, smut sori and ergots are easily incorporated with the seed
lots.
(b) Externally seed borne: When the pathogen lodges itself on the seed coat as
latent spores or bacteria during the growth of the crop or at the time of harvest and
threshing, close contact between the structure of the pathogen and seeds is
established. For instance, loose sorghum, short sorghum, cotton with bacterial
blight, etc. Due to their innate ability for long life, outwardly seed borne entities
like smut spores found in many diseases can persist for many years. Ex: The spores
of Tilletia caries (the stinking smut of wheat) and Ustilago avenae (the smut of
oats) are still viable after 18 and 13 years, respectively (Williams et al., 1971).
(c) Internally seed borne: The pathogen may enter the ovary and infect the embryo
while it is still in the process of development. They develop an internal seed borne
state. Example: Loose smut of wheat.
Differentiate Seed infection and infestation-
Seed infection: A seed is only infected after the pathogen has grown in or on it for
a while and developed a connection with the tissues of the seed. For instance, in
loose wheat, the fungus develops in the tissues of the embryo and goes latent when
the seed goes into dormancy.
Seed infestation: When the fungus or pathogen is present on the seed coat and in
the seed lot, the seed is infected and serves merely as a vehicle for the infection.
(ii) The use of soil for autonomous dispersal: Facultative saprophytes or parasites
that live in the soil may do so. The pathogen may spread by moving through the
soil, growing there, or by moving soil that contains the pathogen. Whereas the latter
is referred to as dispersal by soil, the former is known as dispersal in soil.
The three steps of dispersal in soil are as follows:
Dispersal in soil:
(a) Soil contamination: The pathogen gradually spreads from an area that has been
infected to a new area, contaminating the soil.
(b) Growth and spread of a pathogen in soil: Depending on the pathogen's capacity
for multiplication and distribution, it is possible for it to expand and spread once it
has entered the soil. The pathogen's ability to adapt to the soil environment and its
capacity for saprophytic survival are its most crucial traits. High growth rates, quick
spore germination, greater enzymatic activity, the ability to manufacture
antibiotics, and tolerance to antibiotics produced by other soil-microorganisms are
all factors that affect a pathogen's ability to survive.
The three different types of pathogens in soil can be distinguished based on this
competitive saprophytic activity. In the absence of host plants, specialised
facultative parasites (Saprophytes) can survive in soil, but they are more dependent
on the host plant's leftovers (e. g., Armillariella mellea, Ophiobolous graminis etc.).
Facultative parasites without specialisation can live their entire lives in the soil
(Pythium sp., and Phytophthora sp.) (Barton, 1957). The presence of an active host
is necessary for soil-borne obligatory parasites like Syncytium endobioticum and
Plasmodiophora brassicae. The diseases remain in the soil as dormant organisms
such as oospores (Pythium, Phytophthora, Sclerospora, etc.), Chlamydospores
(Fusarium), smut spores (Ustilago) (Barton, 1957), and sclerotia (Rhizoctonia,
Sclerotium).
Dispersal by the soil:
During agricultural operations, the pathogen is distributed by the soil through
agricultural instruments, irrigation water, workers' feet, etc. As a result, fungus
propagules and plant detritus containing bacterial and fungal diseases spread over
the field (Beaumont, 1954). The most crucial way of disease dissemination involves
moving soil and propagation materials from one location to another. For instance,
transferring papaya seedlings from a nursery where Pythium ophiodermatid- the
pathogen responsible for papaya stem or foot rot, is present, can introduce the
infection into fresh pits where the seedlings would be transplanted. Similar to this,
diseases existing in the nursery might be transferred to the orchards by fruit tree
grafts that are transported with dirt surrounding their roots (Baker, 1959).
(iii) The plant and its organs as a method of self-sufficient dispersal:
The third mode of autonomous dispersal consists of plants, plant elements other
than seeds that are employed for vegetative growth, unprocessed field output, and
plant debris that builds up during cropping. Example: Seed tubers taken from the
native source of the potato in South America were used to introduce late blight in
North America and Europe. Asian immigrants brought citrus canker to California.
The climate in California was favourable for its pandemic.
(II) Passive or Indirect dispersal:
Plant pathogens can spread passively through both living and non-living entities.
(i) Insects: Insects either outwardly (epizoic) or inside (internally) transport plant
diseases (endozoic). They have the ability to spread bacterial, fungal, viral,
mycoplasmic, spiroplasmic, rickettsial, etc.
Fungal diseases:
Those fungi that produce conidia, oidia, and spermatia in honey secretions with
alluring odours are of particular importance in terms of external transmission. Ex:
Sorghum sugary illness. The ascomycetes' spermatial oozings around the mouth of
the spermagonia draw a variety of insects, including flies, pollinating bees, and
wasps that serve a dual purpose in plant pathogen transmission and pollination. Elm
bark beetles carry Dutch elm disease (Ceratostomella ulmi) internally.
Bacterial diseases:
The citrus canker bacteria (Xanthomonas axonopodis pv. citri) and the fire blight
organism (Erwinia amylovora) are spread by flies (bees), ants, and leaf miners,
respectively (Anderson, 1924). The spotted cucumber beetle (Acalymma vittata)
and the stripped cucumber beetle (Acalymma vittata) both transfer the bacteria
Erwinia tracheiphila that causes cucumber wilt (Diabrotica undecimipunctata).
When the beetles eat on the infected plant, the bacterium contaminates the
mouthparts, travels to the beetle's gut, and spends the winter inside the beetle. So,
the beetle aids the bacteria in two ways: in their spread and in their ability to
survive.
Viral illnesses:
Several types of insects disseminate more than 80% of phytoplasmal and viral
infections. Insect vectors are the particular kind of insects that spread diseases by
acting as specific carriers. In the order Hemiptera, the leaf hoppers (Cicadellidae or
Jassidae) and aphids (Aphididae) are both the most numerous and significant insect
vectors of plant viruses. Whiteflies (Aleurodidae), tree hoppers (Membracidae),
mealy bugs (Coccoidae), and scale insects (Coccoidae) are all Hemipteran species
that can spread virus infections. Few true bugs (Hemiptera), thrips (Thysanoptera),
beetles (Coleoptera), and grasshoppers are plant virus vector insects (Orthoptera).
Table 4.1. Arthropods as vector for viruses.
S.No. Vector Virus
1. Aphid transmitted viruses
Myzus persicae Beet mosaic, Lettuce mosaic,
Potato virus Y, Turnip mosaic,
Beet yellows
Acyrthosiphon pisum Bean common mosaic, Bean
yellow mosaic, Soybean
mosaic, Pea enation mosaic
Toxoptera citricidus Citrus tristeza
2. Leaf hopper transmitted viruses
Nephotettix impicticeps, N. Rice tungro virus
nigropictus, N. virescens
Nephotettix cincticeps, N. Rice dwarf virus
nigropictus
Circulifer tangelos Beet curly top
Agallia contricta Potato yellow dwarf
Graminella nigrifrons Maize chlorotic dwarf
3. Tree hopper transmitted viruses
Mycoplasma diseases:
Phloem is where plant MLOs reside, and insects that feed on plant phloem
distribute MLOs to other plants. The primary vector of mycoplasmal disease
that are transmitted in or on seed and propagation materials. For instance, fusarium
wilt of bananas, citrus canker, downy mildew of grapevines, late blight of potatoes,
etc.
Planting contaminated seed: Planting contaminated bulbs, bulbils, corms, tubers,
rhizomes, cuttings, etc. of vegetatively propagated plants like potato, sweet potato,
cassava, sugarcane, banana, many ornamentals and fruit trees, etc. aids in the spread
of pathogens from field to field, orchard to orchard, locality to locality, or from one
country to another.
When using standard farming techniques, humans who are involved in
preparational cultivation, planting, irrigation, weeding, pruning, etc., contribute to
the spread of plant infections. Workers' clothing, shoes, hands, and other items can
carry spores and other exterior structures of fungi from plant to plant and from field
to field (Berry and Davis, 1957). Via the use of contaminated tools: In the field,
tools used for different cultural operations (weeding, thinning, hoeing, etc.) can
spread pathogens from one region to another. For instance, soil-borne illnesses such
root rot and wilt. Knives used for cutting and pruning plants also aid in the transfer
of seeds from one plant to another. Example: Banana with a bunched up top. By
using grafting and budding material that is contaminated: The most efficient way
to spread infections among horticulture crops is by grafting and budding between
healthy and infected plants.
(iii) Phanerogamic parasite dispersal: Phanerogamic parasites spread viruses by
bridging the gap between infected and healthy plants as Dodder (Cuscuta
California, C. campesris, C. subinclusa etc.) Cucumber mosaic virus is called
Cuscuta subinclusa (California's Cuscuta); Tobacco mosaic virus, Viral tobacco
rattling virus causing tomato spotted wilt, virus that causes tomato bushy stunt,
Cuscuta campestris.
(iv) Birds are a key means of dispersal for the spread of some fungi and blooming
parasite seeds. Crows in the tropics scatter the seeds of gaint mistletoe
(Dendrophthoe sp.) on other trees with their excrement after eating its meaty,
sticky, and gelatinous berries. Birds spread Loranthus seeds by adhering them to
their beaks and through their excrement. Birds carry dodder stem fragments while
they prepare their nests, which allows them to travel to new locations. Moreover,
more than 18 species of birds spread the Endothea parasitica spores that cause
chestnut blight. Several powdery mildew fungi's cleistothecia are conveyed via bird
feathers.
(v) Domesticated and wild animals: Domesticated animals (cattle) consume live
fungal propagules (spores, oospores, or sclerotia) while consuming sick feed and
die as a result. When used as manure, this excrement acts as a source of inoculum
when placed on the ground. Moreover, soil-dwelling fungus, particularly sclerotia,
attach to the legs and hooves of animals and travel there.
Spread By some Non-living Entities:
(i) Wind: Anemochory is the scientific term for the movement of infections by
wind. The upward air currents, wind velocity, and the downward wind movements
all play a role in wind transmission. Fungi, bacterial, and viral propagules are all
effectively transported by wind.
Fungi: Fungal diseases are frequently lightweight and well-suited to wind
dissemination. Fungal infections have evolved to disperse via wind by producing a
large number of spores and conidia, releasing spores with enough force, and
producing spores that are extremely small and light to travel great distances. For
example, rusts, smuts, downy mildew, and powdery mildew.
Wind can be used for both short- and long-distance distribution.
Spores used for short-distance dispersal include rust fungus basidiospores,
powdery mildew fungus conidia, and downy mildew fungus sporangia. The
sole cause of the annual recurrence of grain rusts in the plains of northern India is
the wind, which carries uredospores from the source of survival in the hills in the
far north (Himalayas) and south (Nilgiris).
Long-distance dispersal-adapted spores, including conidia of Alternaria,
Helminthosporium, and Pyricularia, uredospores of rust fungi, and
chlamydospores of smut fungi (Anwar, 1949).
Air currents allow uredial rust fungi to spread over great distances, which causes
damaging epidemics to spread over a large area. For instance, Puccinia graminis
var. tritici uredospores have been found up to 14000 feet above diseased wheat
fields. Similar reports were made of Alternaria spores at 8000 feet, Puccinia
recondita spores at 12500 feet, and Cronartium ribicola spores at 14000 feet
(Atkinson, 1953).
Uredospores of this fungus are blown over a distance of more than 1000 miles in
just two days in the United States, from Mexico in the south to Dakota and
Minnesota in the north. If the uredospores are dispersed at a height of 5000 feet,
they can travel up to 1100 miles in a wind of 30 miles per hour without losing
viability.
Nematodes: Together with fungus, it also aids in the spread of nematode cysts and
the seeds of phanerogamic parasites. Ex: Dust storms from Rajasthan to Haryana
transport the nematode Heterodera major cysts that cause the molya disease of
wheat and barley.
Bacteria: Some pathogenic bacteria are transported over short distances by wind
with the infected material. Example: The bacterial exudates produced by Erwinia
amylovora, the cause of fire blight on apple and pears, are produced as fine strands
that can break off and be carried by the wind.
The insect and mite vectors that carry viruses and phytoplasmas move in different
directions and across greater distances depending on the direction and speed of the
air, but viruses and phytoplasmas are not directly transferred by wind.
(ii) Water: Hydrochory is the term for the transmission of plant pathogens through
water. Although water is less important than air in the long-distance transportation
of diseases, it is more effective because the viruses can germinate more quickly
once they settle on a wet surface. Surface running water and rain splash are the
principal channels via which water is disseminated.
The pathogens are transported over short distances by the surface flow of water
during irrigation from canals and wells or after heavy rains. For instance, rainwater
or irrigation water can spread the mycelial pieces, spores, or sclerotia of fungus
such as Colletotrichum falcatum (red rot of sugarcane), Fusarium, Ganoderma,
Macrophomina, Pythium, Phytophthora, Sclerotium, etc. Only when floods cover
a bigger region or when water flows farther from the sources of pathogen survival
is long distance dispersal by water also possible (ARK, 1932).
Splash dispersal is another name for rain splash dissemination. It is one of the most
effective ways for bacterial plant diseases to spread. Raindrops that hit sores,
pustules, cankers, or even soil surfaces with force may cause the propagules to
splash in small droplets and rest on nearby healthy surfaces that are susceptible, or
the water droplets may be blown over great distances by the wind. Examples
include green ear of bajra, bacterial leaf spot (Xanthomonas campestris pv. oryzae),
and bacterial leaf streak (Xanthomonas campestris pv. oryzicola) (Sclerospora
graminicola).
When rain splashes or irrigation drops fall from above, bacterial and fungal spores
that are already present in the air or on plant surfaces are carried downhill and
deposited on healthy, vulnerable plants. Water not only aids in the growth and spore
release of many fungi, but also plays a significant role in the spread of plant
infections. Moreover, it facilitates the infection and spore germination processes.
Conclusion:
Plant Pathogens adopt several mechanism for the survival and growth of spores.
Many biotic and abiotic factors can help them to survive and grow at max by
different methods. Plant pathogens use infection as a tool for the survival. Primary
infection is the base for the further development of plant pathogens. Primary
inoculum is the source of pathogen survival in the crops. Infected host act as a
reservoir of inoculum. After the disease has begun to affect the crop, the spores or
other pathogen structures serve as sources of secondary inoculum and secondary
infection, which spreads the disease across the field. So, we conclude that plant
pathogens are even smarter than our assumptions as they use different techniques
in different conditions for their survival. Nature nurture every creature on the earth.
References
Abeygunawardena, D. V. W. and Wood, R. K. S. 1957. Factors affecting the
germination of sclerotia and mycelial growth of Sclerotium rolfsii Sacc. Trans.
British Myc. Soc.40: 221–231.
Anderson, E. J. 1951. A simple method for detecting the presence of Phytophthora
cinnamomi Rands in soil. Phytopathology 41: 187–189.