UNIT 14 HOST PLANT RESISTANCE

Structure
Introduction
Objectives
Insect and Host Plant Relationships
The Insect Aspect-Host Selection
The Plant Aspect-Host Characteristics
Concept of Host Plant Resistance
Characterisation of Host Plant Resistance
Intensity of Resistance
Mechanisms of Resistance
Genetic Resistance
Ecological Resistance (Apparent ResistancelPseudo-Resistance)
Factors Affecting the Expression of Resistance
Physical Factors
Biological Factors
Breeding Insect-Resistant Varieties
Successful Uses of Insect Resistant Cultivars
Potential and Limitations of Host Plant Resistance
Summary
Terminal Questions
Answers

14.1 INTRODUCTION
You must have observed that some plants harbour fewer pests than the others. These Most plants are resistant
are said to show resistance to the pests. Conversely, other plants which are preferred to most pathogens and
herbivores.
by pests and are destroyed by them are called susceptible. Crop production in tropical
countries like India depends on many factors including biotic agencies. The biotic
agencies, viz., insect pests play a key role as far as full manifestation of yield potential
of a crop is concerned. Available estimates of crop yield losses vary from 40 percent
to 100 percent in different cereals, pulses, oilseeds, cotton and horticultural crops. To
achieve maximum productivity of crops, adequate crop protection methods have to be
adopted. Amongst various methods host plant resistance (HPR) to insects is an
effective, economical, and environment friendly method of pest management, further
it is compatible with other control strategies, such as biological, cultural and chemical
control. The most attractive feature of HPR is that farmers virtually do not need any
skill in application techniques, and there is no cash investment by the resource-poor
farmers.

Considerable progress has been made in identification and development of crop

cultivars (plant varieties in agriculture and horticulture) with resistance to the major
pests in different crops. There is a need to transfer resistance genes into high-yielding
cultivars with adaptatiori to different agro-ecosystems. Resistance to insects should
form one of the criteria to release varieties and hybrids for cultivation by the farmers.

HPR will not only cause a major reduction in pesticide use and slowdown the rate of
development of resistance to insecticides in insect populations, but also lead to
increased activity of beneficial organisms and reduction in pesticide residues in food
and food products.

Also, HPR is one of the preferred methods for minimizing the damage caused by
insect vectors of viruses, viz., whiteflies. It also protects the crop by making it less
suitable for the pest or by making the crop tolerant to the pest. The result is less crop
Methods of Pest damage. An example of this is the use of smooth-leaf (glabrous) cotton rather than
Management41 hairy-leaf cotton to reduce the impact of whiteflies on yield.

Objectives
After studying this unit you should be able to:
define HPR,
explain insect and host plant relationship,
list the major characteristics by which resistance can be assessed,
explain intensity of resistance,
discuss mechanisms of resistance,
define and differentiate between non-preference, antibiosis and tolerance and
appreciate the advantages of the use of insect resistant crop varieties.

14.2 INSECT AND HOST PLANT RELATIONSHIPS

For understanding the mechanisms of plant resistance to insects, it is important to
understand some of the basic relationships between these organisms. Such
relationships can be commonly referred to as insecdplant interactions, and lay
emphasis on insect attack and plants response to the attack.

14.2.1 The Insect Aspect-Host Selection

The insect aspect of the insecdplant interaction is often described as a series of steps,
in time and space, that lead to suitability of a plant for the insect. The major steps in
this direction include:
i) Location of host habitat;
ii) Location of the host plant;
iii) Selection of the host plant;
iv) Acceptance of the host plant; and
v) Adequacy of the host plant

Location of host habitat: Insects locate the general area of the host by means usually
unrelated to the plant. Various physical stimuli like light, wind, gravity etc. help orient
dispersing insects to the overall location of the host.

Location of the host plant: Once the insects are in close proximity of the host it must
find a proper host. Most insects rely on vision and/or smell to locate a host plant.
Other factors include colour, size and shape e.g., aphids and whiteflies are attracted to
yellow-green surfaces.

However, once the insect is in contact with the plant, short-range stimuli arrest further
movement. These stimuli are both physical, which excite the tactile receptors and
chemical, which excite the chemoreceptors on the antennae, tarsi, mouthpart, etc.

Host-plant selection: Host-plant selection by insects usually involves both primary i

and secondary metabolites. Usually host-plant odour or taste for insects comes from
nutrients and odd compounds that are present in the plant and these inputs are
interpreted by the insect's central nervous system to determine whether a given plant
is a host.

Acceptance of the host plant : Subsequent to host finding, insects may take test bites,
as with some caterpillars, to confirm host recognition.

Major physical factors involved in acceptance of a host may include such factors as
leaf and stem toughness, leaf surface waxes, and pubescence (density and type of
hairs). 'These factors may be important in relation to feeding and/or oviposition.
Adequacy of the host plant : Sufficiency of the plant as a host is finally determined Host Plant Resistance
during feeding. If nutrients are adequate and no toxicity occurs, the insect completes
its life cycle successfully.

14.2.2 The Plant Aspect-Host Characteristics

The plant provides various physical and chemical stimuli and thus becomes an
'important participant in the insect and host plant relationship. Both morphological
and physiological characteristics of a plant elicit insect responses.

Morphological characteristics: Plant morphological features may produce physical

stimuli or bar insect activity. Variations in foliage size, shape, color, surface texture
and presence or absence of glandular secretions, spines, hairs may determine the
degree of acceptance by insects. Pubescence and tissue toughness sometimes limit
insect responses.

Physiological characteristics: Physiological characteristics, which influence insects,

involve chemicals that are products of plant metabolism. Primary metabolic proceis in
plants produce substances to catalyze reactions, build tissues and supply energy. The
plant requires inorganic ions and produces enzymes, hormones, carbohydrates, lipids,
proteins, and phosphorus compounds for energy transfer. Together these primary
metabolites promote growth and reproduction of the plant. For insects, some of these
primary metabolites are feeding stimulants, nutrients and toxicants.

The chemicals produced during secondary metabolic processes in plants are known as
secondary metabolites. These may be stored in any convenient place in plant structure
and, often, are extruded from the outer layers of plant tissues. Here, the insects may
sense them as token stimuli. A token stimulus elicits a response initially but
afterwards has no effect.

The relationship between chemical stimuli and insect response is a form of chemical
communication between these organisms. Such chemicals are called semiochemicals
(semiosis: observation of signs). You have already studied about semiochemicals in
detail in Unit 11 of this course. Let us recollect the concept again. Among
semiochemicals are pheromones, which promote communication between members of
the same species, and allelochemicals, which promote communication between
members of different species. Allelochemicals can be subdivided further into
allomones and kairomones. Allomones are mostly defensive chemicals, producing
negative responses in insects and reducing chances of contact and utilization. They
include repellents, oviposition and feeding deterrents, and toxicants. Conversely,
kairomones are advantageous to an insect, promoting host'oviposition and feeding.
They include attractants, arrestants, excitants, and stimulants.

One of the most classical examples of exploitation of allelochemicals in an economic

crop is that of resistance in maize to first generation of the European corn borer,
Ostrinia nubilalis (Hubner). The chemical compound resistance factor, identified as
MBOA (6-methoxy-2-benzoxazolinone), appears to function as a repellent andlor
feeding deterrent. MBOA does not exist in uninjured corn tissues, but DIMBOA (2,4-
dihydroxy-7- methoxy-l,4-6 benzoxazine-3-one) in its glucoside form is present in
uninjured corn leaves. Injury to the plant tissue due to larval feeding results in the
enzymatic conversion of the glycoside to the glycone DIMBOA. The role of
allelochemicals in insect-plant interactions as stimulant and deterrent is given in Table
14.1.
Methods of Pest Table 14.1: Dual role of allellochemicals in insect-plant interactions
Management-I1
Allelochemical Insect Reaction

Stimulant for Deterent for

Cucurbitacin Diabrotica lindecimpunctata Epilachrza tredecimnotata
Cyanogenic glycoside Epilachna varivestris Many phytophagous insects
Furanocoumarins Papilio polyxenes Spodoptera exenzpta,
S. litura
Glucosinolate Pieris rapae Papilio polyxenes
Gossypol Anthonomus grandis Helicoverpa zea
Lupanin Macrosiphon albifons Acrythosiphon pisum
Tomatine Pieris brassicae Leptinotarsa decemlineata
Emuoasca fabae

SAQ 1
Fill in the blanks:
i) The major steps taken by an insect that lead to suitability of a plant for that insect
include:

c) ................................;
d) ................................; and
e) ................................ ;
ii) Both ....................... and .................................. characteristics of a
plant elicit insect responses.
iii) Plant morphological features like foliage .................... ., ........................,
........................., ..................and presence or absence of ...................,
................................, .............................may determine the degree of
acceptance by insects.
iv) Physiological characteristics, which influence insects, involve chemicals that are
products of ................................................
v) Some allellochemicals which play an important role in insect-plant interactions
are .........................................................................................,
..............................................................................................
,

14.3 CONCEPT OF HOST PLANT RESISTANCE

Although there are a number of agricultural pests throughout the world yet only a few
species of pests attack any plant species. This is because plants have some resistance
to most of the pests. Had it not been there, very few plants would have been around.
Since host plant resistance is the result of interactions between the plant and insect, it
is therefore assumed that optimum conditions under which a plant species is grown
are also favourable enough for growth and development of the insect so that the plant
species is accepted by the insect. The concept of host plant resistance should,
therefore, be viewed by comparing the performance of a variety under optimum
conditions for the growth and development of the plant in the absence and presence of
insect population capable of causing maximum loss to the host plant.
Snelling (1941) defined plant resistance as those characters that enable a plant to Host Plant Resistance
avoid, tolerate o r recover from attacks of insects under conditions that would
cause greater injury to other plants of the same species.

Painter (195 1) defined resistance as the relative amount of heritable qualities

possessed by the plant, which influence the ultimate degree of damage done by
the insect.

Beck (1965) defined resistance as collective heritable characteristics by which a

plant species, race, clone o r individual may reduce the probability of successful
utilization of that plant as a host by an insect species, race, biotype (new races o r
strains) or individual.

Maxwell and others (1972) extended the definition of Painter (1951) by considering
level of insect infestation and environmental conditions. According to them,
resistance is those heritable characteristics possessed by the plant, which
influence the ultimate degree of damage done by the insect.

According to Kogan (1994) resistance to insects is the heritable property that

enables a plant to restrain the growth of insect populations or to recover from
injury caused by populations that were not restrained.

So, in a nutshell we can say that Host Plant Resistance is amnheritable trait that
enables the plant to inhibit the growth of insect population or to recover from
injury caused by populations that were not inhibited to grow.

SAQ 2
Give a general definition of Host Plant Resistance.
......................................................................................................

14.4 CHARACTERISATION OF HOST PLANT

RESISTANCE
There are four major characteristics by which resistance can be assessed:
Resistance is heritable and controlled by one or more major genes.
Resistance is relative and can be measured only by comparing with a susceptible
cultivar of the same species.
Resistance is measurable and its magnitude can be determined quantitatively (by
insect establishment) and qualitatively (by analysis of the standard scoring
systems).
Resistance is variable and can be modified by abiotic and biotic components of
the environment.

Based on the above characteristics, resistance can be classified genetically and

epidemiologically. Genetically, there are three main types of resistance i.e.
monogenic, oligogenic and polygenic. Monogenic resistance is controlled by a single
major gene. Oligogenic resistance is the term used when the character is controlled by
several genes. Polygenic resistance is the result of many genes and is clearly more
difficult to incorporate into a plant breeding programme. Cytoplasmic resistance is
due to self-duplicating, mutable substances in the cytoplasm.

In epidemiological term, resistance is classified as either horizontal or durable

resistance, with a long lasting effect and effective against all genetic variations of a
I Methods of ~ & t
Management-I1
particular pest or vertical resistance or transient resistance that is effective for a short
period and against certain variants only. A few other terms are in use in plant breeding
for pest resistance. Qualitative resistance applies when the frequency distribution of
resistant and susceptible plants in the crop population is discontinuous and the plants
are easily categorized as either resistant or susceptible. Quantitative resistance is the
term used when a crop shows continuous gradation between resistant plants and
susceptible plants within the population.

Passive resistance is when the resistance mechanism is already present before the
pest attack e.g. thick cuticle or hairy foliage. Active resistance is a resistance reaction
of the host plant in response to attack by a parasite pathogens just like the human
production of antibodies in response to foreign matter in the blood.

SAQ 3
i) List the four major characteristics by which resistance can be assessed.

ii) How can the resistance be classified genetically?

14.5 INTENSITY OF RESISTANCE

Interactions between the host plants and insects are spread over a wide range of
intensity. From the point of view of the insect, the interaction varies from totally
unsuitable to completely suitable for its growth and development. On the other hand,
in terms of the host plant, lesser the population of the insect andfor lesser the damage
they cause to the plant, the more resistant the plant is likely to be. Therefore, intensity
of resistance is a relative term and should be discussed in relation to a susceptible
cultivar of the same species. The following scale may be used to classify degree of
resistance based on intensity:
i) Immunity: An immune variety is one, which a specific insect will never consume
or injure. There are thus, very few, if any, cultivars immune to the attack of
specific insects, which are otherwise known to attack cultivars of the same
species.
ii) High resistance: A variety with high resistance is one which possesses qualities
resulting in small damage by a specific insect under a given set of conditions.
iii) Low resistance: A low resistance level indicates the possession of qualities which
cause a variety' to show lesser damage or infestation by an insect than the average
foi the crop under consideration.
iv) Susceptibility: A susceptible variety is one, which shows average, or more than
average damage caused by an insect.
v) High susceptibility : A variety shows high susceptibility when much more than
average damage is done by the insect under consideration.

These terms are used for expressing resistance vis-A-vis screening of varieties under
field conditions and have nothing to do with the mechanism of resistance. An
intermediate level of resistance is , sometimes referred to as moderate resistance.
 Host Plant Resistancc
SAQ 4
Fill in the blanks:
i) The .................. of resistance is a relative term and should be discussed in
relation to a .................... cultivar of the same species.

ii) A rating scale for classifying the degree of resistance based on intensity may be
used, that includes:
........................,
........................
........................ ,
........................ ; and
........................ ,

14.6 MECHANISMS OF RESISTANCE

For normal growth and development to occur in insects, certain requisites, available in
proper amounts and at specific times, are necessary. Resistant cultivars, by one means
or the other, do not fulfill these and thereby interruptthe normal host selection
process. In some instances, the mechanisms of resistance involve new allomones or
increased amounts of the existing ones; in others they may be based on reduced levels
of kairomones. Also, physical factors may be involved.

Most authorities consider true plant resistance as being primarily under genetic
control. In other words, the mechanisms of resistance are derived from preadapted
inherited characters. Therefore, the expressions of these characters can also be
mediated by environmental conditions.

14.6.1 Genetic Resistance

The most widely accepted classification of genetic resistance modes or mechanisms in
plants is that of Painter (1951). These modes or mechanisms include:
i) Non-preference (Antixenosis)
ii) Antibiosis and
iii) Tolerance.

Non-Preference: It is the inherited feature of a host which discourages the

feeding, colonization or oviposition of an insect pest. Another term used for non-
perference is "antixenosis", (Xenosis is a Greek word for guest; therefore antixenosis
means against or expelling guests).

In other words, non- preference refers to plant characteristics that lead insects away
from a particular host; it includes activities of both the plant and the insect. With non-
preference, normal insect behaviour is impaired in such a way as to lessen chances of
the insects using a plant for oviposition, food, or shelter. Non- preference can be
expressed in a cultivar through either allelochemic or morphological characteristics.

Allelochemic non-preference:This form of non-preference is common among plants,

sometimes causing them to be totally rejected by insects. Allelochemic non-
preference occurs with insects like the spotted cucumber beetle, (Diabrotica sp.) on
cucurbits. In this insectlplant relationship, cucurbitacins (a class of tetracyclic
terpenes) produced by the cucurbit plants act as attractants and feeding stimulants for
the beetles. Cultivars that lack or display low levels of specific cucurbitacins attract
fewer beetles and therefore receive much less damage than those with the feeding
requisites.
Methods of Pest Morphological non-preference: This form of non-preference results from plant
Management-11 structural characteristics, which disrupt normal behaviour by physical means. For
example corn ear worm, Heliothis t e n prefers to oviposit on pubescent (hairy)
surfaces. Some common physical factors imparting resistance to crops plants are listed
below in Table 14.2.

Table 14 2: Physical factors imparting resistance to crop plants

- - - - - - - - - -

Plant factors Effect on insect

Thickening of cell walls
Proliferation of da~nagedand
wounded tissues
Solidness of stems and other
characteristics of the stems
Presence of pubescence ( Trichornes)
Anatomical adaptations
Si~rfacewaxes
Silica deposits
Husk tightness
Affects feeding and oviposition mechanisms
Insects do not survive after initial injury
Interference in feeding and oviposition
mechanisms, dehydration of eggs
Obstruction in feeding, digestion, ovi-
position. locomotion, attachment and
provision of shelter.
Affects various activities of insects
Colonization and oviposition affected
Feeding inhibition, abrasion in cuticle
Resists insect attack

Forms of morphological non- preference that impair feeding behaviour are very
important and may be the first line of defense against many pests. This is partly
because morphological non- preference provides long lasting effectiveness. compared
with most chemically based resistance; that is, insect populations have a difficult time
overcoming this form of resistance.

The term anlibiosis Antibiosis: It is the adverse effect the plant exerts on the growth and survival of the
encolnpasses all adverse insect. In other words plant resistance affects the biology of the insect, so pest
physiological effects of a
temporary or a permanent abundance and subsequent damage is reduced compared to that which would
nature resulting From the have occurred if the insect were on a susceptible crop variety. Antibiosis
ingestion of a plant by an resistance often results in increased mortality or reduced longevity and reproduction
insect. of the insect.

As with non-preference, both insect and plant factors are involved in antibiosis
mechanism. This mechanism usually impairs an insect's metabolic processes and
often involves consumption of plant metabolites.

Allelochemicals are generally involved with antibiosis. Some of the best-known

allelochemicals include the cyclic hydroxamic acids in corn (DIMBOA), gossypol and
related compounds in cotton, steroidal glycosides in potato, and saponins in alfalfa.

Quantity and quality of primary metabolites also may be important in conferring

antibiosis. Particularly significant are imbalances of sugars and amino acids that result
in nutritional deficiencies for insects feeding on the plant. For example: pea cultivars
with low amino acid levels and increased sugar content show resistance to the pea
aphid, and rice cultivars deficient in asparagines ( an amino acid ) cause reduced
fecundity in the brown planthopper.

Symptoms of insects affected by antibiosis include the following:

i) Death of' young lmmatures
ii) Reduced growth rate
iii) Illcreased mortality in pupal stage
iv) Small adults with reduced fecundity Host Plant Resistance
V ) Shortened adult life-span
vi) Morphological malformations
vii) Restlessness and other abnormal behaviour

Tolerance: It enahles the host plant to withstand or recover from damage caused
by insect pest abundance equal to that damaging a plant without resistance
characters (susceptible). Tolerance is a plant response to an insect pest.

Many factors are involved in plant tolerance, yet overall mechanisms are poorly
understood. Known components of this form of resistance include general vigor,
compensatory growth in individual plants andlor plant population; wound healing,
mechanical support in tissues and organs, etc. An example of tolerance is found in
several corn genotypes that have the ability to repair and replace roots fcd upon by the
western corn borer. Diahrotira sp., Such tolerance allows the plants adequate water
and nutrient uptake and anchorage despite heavy feeding. Surprisingly, tolerant
genotypes developed greater root volume with rootworm feeding than without it.

Thus, tolerance resistance differs from antibiosis and antixenosis resistance in how it
affects the insect-plant relationship. Antibiosis and antixenosis resistance cause an
insect response when the insect attempts to use the resistant plant for food,
oviposition, or shelter.

Tolerance is useful in pest management due to certain distinct advantages:

Tolerant varieties have a higher economic threshold level than the susceptible
varieties and hence require less insecticide application and promote bio-control.
Tolerant varieties do not depress insect populations nor do they provide any
selection pressure on the insect and thus are useful in preventing the development
of insect biotypes.
In varieties with a combination of three mechanisms of resistance, tolerance
increases yield stability by providing at least a moderate level of resistance, when
vertical genes providing a high level of resistance through antixenosis and
antibiosis succumb to the new biotype.

14.6.2 Ecological Resistance (Apparent Resistance/Pseudo-Resistance)

Ecological resistance sometimes also called as apparent resatarzce or pseudo-
re.si~tunce,is not considercd as true resistance by many authorities. This is because
ecological resistance relies more on environmental conditions than on genetics.

The characteristics of this resistance are temporary, and cultivarc involved ire
potentially susceptible. Apparent resistance is important in insect pest management,
but its use must be carefully synchronized with prevailing environmental conditions
for effectiveness. The three types of apparent resistance recognized by various
authorities include host evasion, induced resistance and host escape.

Host ewasion: Alterations in plant growth patterns that result in asynchronies of

insect-host phenologies constitute the modality of resistance called host evasion.
With host evasion the plant passes through a susceptible stage quickly or at a time
such that its exposure to potentially injurious insects is reduced. Often host evasion is
accomplished by planting early maturing varieties.

Induced resistance: The qualitative or quantitative enhancement of a plant's

defense mechanisms against pests in response to extrinsic physical or chemical
stimuli is called as induced resistance. Induced resistance is a form of temporary
resistance derived from plant condition or the environment. Here factors like
fertilization and changes in the soil moisture levels may make the plants more tolerant
Methods of Pest to insects than under other circumstances. For example: nitrogen and potassium levels
Management4 are known to affect aphid population on the plants. High nitrogen varieties usually
allow increases in the survival, but the opposite may occur for high levels of
potassium. Providing a proper balance of these nutrients in fertilizers has been
suggested as a means of inducing resistance to aphids.

The use of the chemical insecticide thiamethoxan 70 WS as seed treatment is a very

good example of induced resistance in soybean crop (Fig. 14.1). By using only one
application of this chemical, the highly susceptible soybean varieties like Pusa 16,
Pusa 20 behave like resistant varieties during the entire plant growth period against a
key insect pest, whitefly Bemisia tabaci. An example of resistant v&ety of soybean,
PK 1092 is shown in Fig. 14.2.

Fig. 14.1: Susceptible variety of soybean, Pusa-16 (right). Resistance to Yellow Mosaic Virus
transmitted by white fly has been induced by thiamethoxan @ 3gmlkg seed (left).

Fig 14.2: Resistant variety (PK 1092) of soybean released from Pant Nagar.
Host escape: This refers to the lack of infestation of susceptible plants in a Host Plant Resistance
population of otherwise infested plants. The principle of host escape recognizes that
the presence of an uninfested plant may not mean that it is resistant and emphasizes
that escapes occur in most plant populations, even with heavy insect infestations. The
reason for this escape is rarely understood.

SAQ 5
i) Fill in the blanks
a) The most widely accepted classification of genetic resistance modes or
mechanisms in plants by R. H. Painter ( 1 951) includes ...................... ...,
................................ .., and .....................................
b) The three types of ecological or apparent resistance recognized by various
authorities include ......................... ..., ................................ ., and

ii) List some physical factors imparting resistance to crop plants.

...................................................................................................

- -

14.7 FACTORS AFFECTING THE EXPRESSION OF

RESISTANCE
Resistance is primarily governed by genetics but physical and biotic factors of the
environment also often influence its expression. ,

14.7.1 Physical Factors

Weather, soil, plant architecture, and cultural practices are some of the most important
influences on the plant's physical environment. These factors can affect plant
resistance by influencing such elements like temperature, light intensity, and soil
fertility. Changes in these elements cause fundamental changes in plant physiological
processes and can alter levels of allellochemicals or cause imbalances in basic
nutrients.
i) Temperature: Abnormally high or low temperatures for a period of time may
cause loss of resistance e.g. loss of resistance to the Hessian fly has been found in
some wheat varieties at temperatures below 1 8 ' ~ .
ii) Light intensity: Shade induced loss of resistance has been found in several
instances e.g. wheat resistant to wheat stem fly, sugar beets resistant to green
peach aphids, potato resistant to Calarado potato beetle etc. With potatoes,
shading was found associated with reduced levels of steroidal glycosides in
leaves. These substances are known to retard feeding and development of the
beetle.
iii) Soil fertility:Changes in soil nutrient levels may also mediate the expressions of
resistance in some plants, but little is known about the mechanisms involved e.g.,
alfalfa resistant to spotted alfalfa aphid were found to have reduced resistance if
deficient levels of calcium or potassium or excess levels of magnesium or
nitrogen were present.

14.7.2 Biological factors

The most important biological factors are the selection of biotypes and changes in
resistance with plant ag
- F
 Methods of Pest
Management41
I Crop varieties that have
resistance to pests or have
i . desirable agronomic
characteristics can he
developed by cross
breeding crop varieties with
resistant or desirable plants
in thc same species.
~ Attention to terminology. In
plant breeding, resistant is
i a crop qualifier. A wheat
variety is resistant to the
rust pathogen. In
toxicology, resistant is a
pest qualifier. The Colorado
potato beetle is resistant to
parathion.
I
I control method
I CULTIVARS
16
i) Biotypes: When resistant cultivars are grown widely, selection pressure is
imposed by these hosts on the insect population. When capable, the insect
population responds with genotypes with virulence to overcome the resistance.
With time, these genotypes with superior fitness increase in number and displace
the earlier ones. This results in a situation of growing ineffectiveness of the
resistant cultivar e.g., biotypes of brown plant hopper in rice, whiteflies in cotton
etc.
ii) Plant age: Physiological responses in plants vary with age, and these can lead to
changes in the expression of cultivar resistance e.g., resistance in corn to the
European corn borer results from the presence of DIMBOA (2,4-dihydroxy-7-
methoxy-l,4-benzoxamine-3-one).DIMBOA levels are highest during the early
growing season, thus offering maximum resistance to the first generation corn
borers. DIMBOA levels decrease as the season progresses, and most cultivars
have very little resistance to second-generation corn borers.
SAQ 6
Enlist the physical and biotic factors of the environment which often influence the
expression of resistance.
14.8 BREEDING INSECT-RESISTANT VARIETIES
The basic approach underlying plant breeding for pest management is to develop a
crop variety that is resistant to the pests. Development and standardization of
screening techniques is an essential pre-requisite to any effective breeding
programme. For this, one may need information about the following:
Nature of crop
Insect pests involved
Initial information and data regarding
Possible sources of resistance from varieties and strains locally available
Possibility of acquiring new germplasm that may carry resistance
Determination of some of the basic properties of the plant responsible for
resistance
Hybridization to combine gene for resistance with desirable agronomical
characters
Study of resistance in advance generations
Study of genetics of plant resistance to the insect
Study of resistance of released varieties in plots to evaluate resistance on an insect
SAQ 7
What information is required for breeding insect resistant varieties?. ....................
14.9 SUCCESSFUL USES OF INSECT RESISTANT
Screening for insect resistance under natural conditions is a long-term process. It is
difficult to identify reliable and stable sources of resistance under natural field
conditions, as there is a lot of variation in insect population in space and time. In order
to overcome such problems, it is essential to develop and standardize suitable
 Host Plant Resistance
,screening techniques. Such techniques are available in India for leafhoppers, gall
midge, and borers in rice; shoot fly, stem borer, midge and ear head bugs in sorghum;
army worms in pearl millet and sorghum; leaf miner, aphids, jassids and Spodoptera
in groundnuts; pod-borer in chickpea and pigeonpea; stem borer in maize etc. (Table
14.3).

Table 14.3: A few selected Insect-resistant cultivars/sources of resistance

identifiedldeveloped for different crops in India

Common name Scientific name Cultivar(s)

(some examples)
1 2 3
CEREALS AND MILLETS
Maize (Zea mays)
Spotted stem borer Chilo partellus Ganga Safed, Kundan
I Rice (Oryza sativa)
1 Brown planthopper
1 I 1
Nilaparvata lugens Manasarowar*, Bhadra*,
Joyti*.

Yellow stem borer Scirpophaga incertulas Ratna*, ~aiyasree*,

Saket*, Amelbero*, IET

PULSES
Chickpea (Cicer arietinum)
Pod borer Helicoverpa annigera ICC 506, PDE 2, Dulia*,
Cowpea (Vigna unguiculata)
Aphid Aphis craccivora
I Jassid I Empoasca kerri I JG 10-72
I
Mung (Vigna radiata)
Galerucid beetle M. obscurella Jawahar 45*, PIMS 3,
Gujarat 1*
White fly Bemisia tabaci
Pea (Pisum sativum)
Pod Borer Etiella zinckenella EC 33860, Bonville*, T
6113*
Pigeonpea (Cajanas cajan)
Pod fly Melanagromyza obtusa ICP 10531-El, ICP 7941
El
Urd (Vigna mungo)
Galerucid beetle M. obscurella Pusa 1*, KG 3, Krishna*,
T9
Pod borer I H. a n n i ~ e r a I Kalai*, 338-3, Krishna*
OILSEEDS
Groundnut (Arachis hypogea)
Jassid E. kerri NCAC 2230, ICG 5049
Leaf miner Aproaerema modicella ICGV 8603 1, DRS 10,
7404 and 9883
/ Thrips I Thrips palmi I M 13*, Robut 33-1' I1
1 Tobacco leaf caterpillar I Spodoptera litura I ICGV 86031, FDRS lo*
Methods of Pest Rape and Mustard (Brassica
Management-I1
SPP.)
Mustard aphid Lipaphis erysinzi C 294, Laha 101*, Pusa
Kalyani*
1 Soybean (Glycine max) 1 1 1
Leaf miner ' A. rnodicella PI 227687, Nimsoy*, PL
507, J9 75-1, PK 1092
1
CASH CROPS
Cotton (Gossypiunt spp.)
Cotton jassid Amrasca biguttula Khandwa 2, Badnawar,
Krishna*, Mahalaxmi*,
Eknath*
Pink bolloworm Pectinophora gossypiella Lohit, Adadhita, MCU 7,
Sujata, Sanguineurn*
Spotted bollworm Earias vitella L 1245*, JK 119-25-54,
Supriya*, LPS 141*
Sugarcane (Saccharum
offlcinarum L.)
Stalk borer Chilo auricilius Co 7302*, CoS 767
Top borer Scirpophagu excerptalis Co 7224, Co 67, Co 1158
Tobacco (Nicotiana spp.) Jamaica*, Cuban*,
~ y z u persicue
s
Aphid . Sumatra*
Stem borer 1 Scrobipalpa heliopa 1 SBR 1*, SBR 2*
VEGETABLES
Lady's finger (Abelmoschus
esculentus)
Shoot and fruit borer E, virella AE 57, Parkins long
green*, Karnual special*
Potato (Solanum tuberosum L.)
Potato tuber moth Phthorirnuea operculella

Tomato (Lycopersicum
esculentum) H. arrnigera
Fruit borer BT 1, T 32*, T 27"
*Released for cultivation

SAQ 8
Name two resistant cultivars each against the following insect pests:
i) Brown plant hopper of rice
...................................................................................................

ii) Yellow stem borer of rice

..................................................................................................

iii) Pod borer of chickpea

...................................................................................................
! iv) Leaf miner of groundnut Host Plant Resistance
.................................................................................................

V) Mustard aphid
...................................................................................................

14.10 POTENTIAL AND 1,IMITATIONS OF HOST PLANT

RESISTANCE IN IPM
In contrast to natural ecosystems, most agricultural crop production systems are
ecologically unstable, non-sustainable, and energy dependent. Through plant
domestication and cultivation practices man has interfered with species diversity and
b natural defense inechanis~nsof plants in many ways. Cultivated crops originated from
genetically diverse plant types. However, crop plants are now grown in large,
genetically homogeneous stands, a practice that decreases genetic and species
I diversity and increases the likelihood of economically significant insect pest
infestations. Defense mechanisms of plants are re-created in resistant plants. Plant
defense mechanisms include escape in space and time, incompatible biological
associations, physically and chemically derived barriers, and accommodation by
replacement or repair of damaged plant parts. By re-creating plant defenses, genetic
resistance to insect pests plays a vital role in an environmentally compatible manner to
enhance ecological stability in agricultural crops.

An ZPM Direct Control Tactic: Plant resistance to insects is one of several cultural
control methods. Cultural control methods involve use of agronomic practices to
reduce insect pest abundance and damage below EIL, which would have occurred if
the practice had not been used. In IPM, plant resistance to insects refers to the use of
resistant crop varieties to suppress insect pest damage. Plant resistance is intended for
use in conjunction with other direct control tactics.

Advantages to the Use of Insect-Resistant Crop Varieties: Use of insect-resistant

crop varieties is economically, ecologically, and environmentally advantageous.
i) Economic benefits occur because crop yields are saved from loss to insect pests
and money is saved by not applying insecticides that would have been applied to
susceptible varieties. In most cases, seed of insect-resistant cultivars costs no
more, or little more, than for susceptible cultivars.
ii) Ecological and environmental benefits arise from increases in species diversity in
the agroecosystem, in part because of reduced use of insecticides. Increase in
species diversity enhances ecosystem stability, which promotes a more sustainable
system far less polluted and detrimental to natural resources.
The IPM concept stresses the need to use multiple tactics to maintain insect pest
abundance and damage below levels of economic significance. Thus, a major
advantage to the use of insect-resistant crop varieties as a component of IPM
arises from the ecological compatibility and compatibility with other direct
control tactics.
iii) Insect-resistant cultivars synergize the effects of natural, biological, and cultural
insect pest-suppression tactics. The "built-in" protection of resistant plants from
insect pests functions at a very basic level, disrupting the normal association of
the insect pest with its host plant. The compatible, complementary role host plant
resistance to insect pests plays with other direct control tactics is, in theory and
practice, in concert with the objectives of IPM -all crop cultivars should contaln
resistance to insect pests.
 Methods of Pest iv) Host plant resistance to insect pests has advantages over other direct control
Management-I1 tactics. For example, plant resistance to insects is compatible with insecticide use,
while biological control is not. Plant resistance to insects is not density dependent.
whereas biological control is.
v) Plant resistance is specific, only affecting the target pest.
vi) Often, the effects of use of insect-resistant cultivars are cumulative over time.
Usually the effectiveness of resistant cultivars is long lasting.
vii) HPR is usually the only tactic required to manage a pest when a highly resistant
variety is available.
viii)Protection is independent of weather.
ix) All parts of plant are protected including those that are difficult to treat using
conventional pesticides.
x) The pest is always influenced at the most sensitive stage.

The role of host plant resistance to insects in IPM has been well defined, at least in
theory. However, the specific role a resistant cultivar plays in a particular IPM
situation is crucial to successful deployment of the resistant cultivar. The impact of
the resistant cultivar on standard cultural, biological, and insecticidal control methods
should be well defined. Likewise, the impact of each of these control tactics on the
resistant cultivar also must be defined.
I

I
Several definitions have been used to convey the relative level of resistance in a plant.
However, the problem of quantifying resistance continues to be a problem influencing
farmer acceptance of insect-resistant cultivars. A better way to define resistance levels
in agronomically improved resistant cultivars is through quantified comparisons of
insect pest damage or plant yield loss of susceptible cultivars. Once insect pest
abundance or damage to yield-loss relationships have been determined, economic
threshold levels can be determined and combined with factors such as crop value and
I insect pest control costs to develop dynamic thresholds for use by producers. Dynamic
thresholds provide a description of resistance and can reduce crop loss risk because
limitations are known and remedial action can be taken when necessary. By using this
system to define relative differences in insect pest resistance between cultivars, it may
be possible to simply indicate that a resistant cultivar has a higher economic threshold
level than a traditional susceptible cultivar.

Although the advantages of plant resistance far outweigh the limitations, however the
drawbacks of plant resistance should also be recognized:
I

i) Time of development: This is a lengthy method hence is not adequate for solving
sudden or localized pest problems.
ii) Genetic limitations: Absence of resistance genes among available germplasm
makes the programme more complex.
iii) Biotypes: The occurrence of biotypes may limit in time and in space the use of
resistant varieties.
I iv) Conflicting resistance tracts: Some plant characteristics may act as resistance
factors for a species but include susceptibility to them.

SAQ 9
List a few advantages and limitations in the use of insect resistant crop varieties.

nn
 Host Plant Resistance
114.111 SUMMARY
After studying this Unit, you have learnt that:
Crop production in tropical countries like India depends on adequate crop
protection.
Host plant resistance (HPR) to insects is an effective, economical, and .
environment friendly method of pest management, further it is compatible with
other control strategies, such as biological, cultural and chemical control.
The most attractive feature of HPR is that farmers virtually do not need any skill
in application techniques, and there is no cash investment by the resource poor
farmers.
Considerable progress has been made in identification and development of crop
cultivars with resistance to the major pests in different crops.
HPR will not only cause a major reduction in pesticide use and slowdown the rate
of development of resistance to insecticides in insect populations, but also lead to
increased activity of beneficial organisms and reduction in pesticide residues in
food and food products.
Host Plant Resistance is an inheritable trait that enables the plant to inhibit the
growth of insect population or to recover from injury caused by populations that
were not inhibited to grow.
Interaction between host plants and insects are spread over a wide range of
intensity.
The mechanisms of resistance involves physical factors as well as levels of
allomones or kairomones.
'The most widely accepted classification of genetic resistance modes or
mechanisms in plants is that of Painter (195 1) and includes non-preference,
antibiosis and tolerance.
Ecological Resistance sometimes also called as apparent resistance or pseudo-
resistance, is not considered as true resistance by many authorities. This is because
ecological resistance, relies more on environmental conditions than on genetics
and includes host evasion, induced resistance and host escape.
Factors affecting the expressions of resistance are primarily governed by genetics
but physical and biotic elements of the environment also often influence its
expressions.
Physical factors like weather, soil, plant architecture, and cultural practices are
some of the most important influences on the plant's physical environment. These
factors can affect plant resistance by influencing such elements like temperature,
light intensity. and soil fertility.
Biological factors are the selection of biotypes and changes in resistance with
plant age.
Breeding insect-resistant varieties is an essential feature of host plant resistance
for which developnient and standardization of screening techniques is an essential
pre-requisite .

14.12 TERMINAL QUESTIONS

1

1. List the major steps involved in the insect / plant interactions.

Methods of Pest
Management-U

...................................................................................................
.What is apparent or pseudoresistance?

...................................................................................................
...................................................................................................
Discuss the factors affecting the expressions of resistance.
...................................................................................................
...................................................................................................

Discuss the essential pre-requisites for an effective breeding programme.

...................................................................................................
Discuss the potential of HPR in IPM.
...................................................................................................
...................................................................................................

Write short notes on the following:

i) Allellochemicals I

...................................................................................................
iii) Intensity of resistance

...................................................................................................
...................................................................................................
iv) Characteristics of HPR

...................................................................................................
...................................................................................................
Non - preference
....................................................................................................
...................................................................................................
...................................................................................................
 Host Plant Resistance
vi) Antibiosis

vii) Tolerance
...................................................................................................

14.13 ANSWERS
Self-assessment Questions
I. i) The major steps include
a) location of host habitat;
b) location of the host plant;
c) selection of the host plant;
d) acceptance of the host plant ; and
e) adequacy of the host plant
ii) Morphological and physiological
iii) Size, shape, color, surface texture and presence orabsence of glandular
secretions, spines, hairs
iv) Plant metabolism
v) Cucurbitacin , Furanocoumarins, Gossypol, Lupanin, Tomatine.

2. Resistance is the relative amount of heritable qualities possessed by the plant,

which influencesthe ultimate degree of damage done by the insect.

3. i) The four major characteristics by which resistance can be assessed are:

Resistance is heritable and controlled by one or more major genes.
Resistance is relative and can be measured only by comparing with a
susceptible cultivar of the same species.
~esistanceis measurable and its magnitude can be determined quantitatively
(by insect establishment) and qualitatively (by analysis of the standard scoring
systems).
Resistance is variable and can be modified by abiotic and biotic components
of the environment.
ii) Monogenic, Oligogenic and Polygenic resisstance

4. i) Intensity, susceptible
ii) a) Immunity
b) High resistance
c) Low resistance
I d) Susceptibility
e) High susceptibility

5. i)' a) Non-preference, Antibiosis and Tolerance.

b) Host evasion, induced resistance and host escape.
Methods of Pest ii) Thickening of cell walls, Proliferation of damaged and wounded tissues.
Management41 Solidness of stems and other characteristics of the stems, Presence of
pubescence (Trichomes), Anatomical adaptations, Surface waxes. Silica
deposits, Husk tightnessetc.

6. Physical Factors - Temperature, Light intensity, Soil fertility

Biological factors - Biotypes, Plant age

7. Refer to Section 14.8

8. Refer to Table 13.3

9. Refer to Section 14.10

Terminal questions
1. Refer to Section 14.2
2. Refer to Section. 14.6
3. Refer to Section 14.6.2
4. Refer to Section 14.7
5. Refer to Section 14.8
6. Refer to Section 14.16
7. i) Refer to Section 14..2.2
ii) Refer to Section 14.5
iii) Refer to Section 14.5
iv) Refer to Section 14.4
) Refer to Section 14.6.1
i ) Refer to Section 14.6.1
vii) Refer to Section 14.6.1