Dr. Muthu Venkateshwaran, Ph.D. N.R.

McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
Name of Disease: Apple scab

Hosts: In addition to infecting its namesake domestic apple, flowering crabapple and other

Malus species, apple scab also attacks other closely-related genera within the tribe Maleae,

including hawthorn (Crataegus ssp.), mountain ash (Sorbus ssp.), firethorn (Pyracantha ssp.),

and loquat (Eriobyotrya japonica) (Sutton et al.). Given the hosts are all incident to one tribe

within the Rosaceae, this pathogen has a relatively narrow host range.

Causal Organism: Apple scab is caused by Venturia inaequalis (Cooke) G. Winter (anamorph:

Spilocaea pomi). An ascomycete with septate hyphae and haploid nuclei in the subclass

Pleosporomycetidae, order Pleosporales, family Venturiaceae, the fungus is bipolar heterothallic,

producing ascocarps (in this case pseudothecia) in a stroma in overwintering leaves or fruit on

the orchard floor (Fig. 10.). Pseudothecia are negatively geotropic, solitary, dark brown to black,

spherical in shape, and approximately 90 to 150 μm in diameter (Fig. 6.). Each pseudothecium

has a distinct ostiole with single-celled apical bristles atop a short beak (Fig. 7.) The centrum is

pseudoparenchymatous with pseudoparaphyses (MacHardy).

Asci – of which approximately 50 to 100 are contained per pseudothecium – are six to twelve μm

wide by 55 to 75 μm long, fasciculate, cylindrical, and eight-spored with a short stipe; the ascus

wall is thin and bitunicate (Fig. 8.). Contained within an ascus are eight ascospores, each of

which are yellowish-green to tan and unequally two-celled, meaning the upper cell is shorter and

wider than the lower cell. The unequal size of the two cells in the ascospore is indicative of V.

inaequalis, it being the only member of the genus Venturia with such unique ascospore

morphology, and thence gives the species its binomial name (Fig. 9.) (Gadoury and MacHardy).

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
As it relates to asexual reproduction, conidiophores arise from shortened, erect, closely septate

brown mycelia. Likewise, the conidiophores themselves are brown, wavy to nodulose, being

either septate or non-septate. Conidia, each six to nine μm long by twelve to 22 μm wide are

yellowish olive, one- or two-celled, single, terminal, and ovate to lanceolate, with irregular

shapes found on occasion (Fig. 11.). Conidia are produced sequentially, abscising from the

conidiophore and leaving behind abscission ridges indicative of V. inaequalis. Conidia are

produced from lesions on the substrate, typically on the surface of leaves, flowers or fruit (Fig.

12.). However, lesions yielding conidia may also be located on shoots or bud scales (MacHardy).

Diagnosis: Physiological responses on the part of the host to being infected by the pathogen are

succinctly known as symptoms, whereas organismal structures illustrating the presence of the

pathogen on a host are known as signs. Symptoms diagnostic for apple scab occur on the aerial

parts of the host, including leaves, petioles, flowers, sepals, fruit, pedicels, young shoots, and bud

scales (Sutton et al.). Signs indicative for V. inaequalis largely entail the presence of

pseudothecia, mycelia within lesions, and conidia. Microscopy is required to accurately locate

and characterize signs of V. inaequalis (Keitt and Jones).

Nonetheless, the symptoms incident to apple scab are generally most noticeable and serious on

leaves and fruit. [Symptoms on blossoms usually occur as small, dark green lesions at the base of

the flower, on the sepals, and on the stem pedicel before and during bloom. When pedicels

become infected the developing fruit may drop, resulting in lower fruit yield (MacHardy).] The

first lesions seen in the spring are usually on the underside of expanding leaves. Once the leaves

open, the upper surfaces also become vulnerable to infection. A lesion first appears as an area

which is a lighter shade of green than the surrounding leaf. The lesion is usually circular and as it

increases in size it becomes olive-colored and velvety due to production of asexual spores known

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
as conidia (Fig. 2.) (Sutton et al.). Lesions that form on young leaves may be quite large, some

more than one centimeter in diameter. Lesions that form on expanded leaves are usually smaller

because older leaves are more resistant to infection. Affected tissues eventually may become

distorted and puckered, and the leaf lesions often become cracked and torn. Lesions on the leaves

and fruit are generally blistered and scabby in appearance, with a distinct margin (Fig. 3-4.). The

earliest noticeable symptom on fruit is water-soaked areas which develop into velvety, green to

olive-brown lesions. Infections of young fruit will cause fruit distortion (Fig. 5.) (MacHardy).

Severely infected leaves or fruit will often drop from the tree. Infection which causes significant

defoliation for two or three years in a row can result in weakened trees that are more susceptible

to winter injury and long-term decline (Sutton et al.).

Disease Cycle and Epidemiology:

Inoculation: Ascospores initiate primary infections on new growth, therefore acting as

primary inoculum. In most years and locations, the first ascospores are mature and

capable of causing infection at about the time of bud break. Ascospores continue to

mature and are discharged over a period of five to nine weeks (Stensvand et al.). The

peak period of ascospore discharge is usually between the pink and full-bloom stages of

blossom development. However, if rain starts during the night and continues throughout

the following morning, few ascospores are released until after sunrise. In the same breath,

low temperatures (i.e., less than ten degrees Celsius) slows the pace – or may even inhibit

– ascospore release. Secondary inoculum consists of conidia carried on the wind or

otherwise splashed onto uninfected leaves, flowers, fruit, etcetera, throughout the summer

(Keitt and Jones). All forms of inocula are partial to infection courts – or, sites on a host

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
where infection can occur – coinciding with younger leaves, flower buds, developing

fruit, twigs and bud scales (James and Sutton).

Penetration: Free moisture is required for the initiation of germination in ascospores

deposited onto a viable substrate. Following initiation however, germination proceeds as

long as the relative humidity is or exceeds 95%. The timetable necessary for infection to

occur is proportional to the number of hours of wetness and ambient temperature. The

duration of the wet period required for infection increases with the maturity of the fruit

(Gadoury and MacHardy).

Once an appressorium and penetration peg has formed on a germinating ascospore,

hyphae directly penetrate the surface of the substrate, becoming established between the

epidermis and the cuticle. Infection may occur in a variety of temperature extremes, from

as little as one degree Celsius to as many as 26°C. The duration of the wetting period for

successful ascospore infection varies with ambient temperature, ranging from as much as

18 to 24 hours at six degrees Celsius to as little as six to nine hours at 16 to 24°C

(Brook). Infection rarely occurs under conditions where air temperature exceeds 26°C

(Keitt and Jones). The physiological phenomenon contributing to this metabolism is not

entirely known, however. (Stensvand et al.).

Infection: After penetrating the cuticle, the infecting mycelium forms a subcuticular

stroma. Said stroma produces conidiophores and conidia in a localized, visible lesion on

primarily leaves and fruit, but also produced on flower pedicels, twigs, and bud scales

(MacHardy). Lesions with conidia at the center are visible nine to seventeen days

following infection. This incubation period, however, is dependent upon both

temperature and relative humidity. Nonetheless, the minimum relative humidity required

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
for sporulation is between 60 and 70%, and periods of subminimal humidity do not

reduce the virulence or survival of the fungus (Brook).

Dissemination: Following leaf drop in autumn, mycelia of compatible mating types

conjugate, undergoing plasmogamy, karyogamy and eventually meiosis, thusly forming

pseudothecia on the leaf tissue. Most pseudothecia form within four weeks of leaf drop,

the ascogonium developing until the lumen in the pseudothecium is filled with

pseudoparaphyses. Immediately following a distinct period of dormancy, the

pseudothecium proceeds with development, maturing in late winter or early spring and

culminating in the synthesis of asci and ascospores. Moisture is necessary for the

development of pseudothecia, and persistent drought dramatically diminishes the rate of

maturation (Brook).

Likewise, the optimum range in temperature for ascogonial development is eight to

twelve degrees Celsius, whereas the optimal temperature range for ascospore maturation

is between sixteen and eighteen degrees Celsius (Stensvand et al.). Factors such as type

of cultivar or the date of leaf fall have no known effect on the date of ascospore

maturation. Nevertheless, mature asci expand through the ostiole and forcibly discharge

ascospores throughout the subsequent spring and early summer. These asci, stimulated to

sporulate from frequent wetting wrought by rain or heavy dew, may be found on

overwintered leaves strewn about the orchard floor (Brook). Upon discharge, the

ascospores are disseminated by the wind – a form of passive dispersal.

Dormancy and Survival: V. inaequalis overwinters as pseudothecia on infected leaves

and fruit on the orchard floor. The fungus can also overwinter as mycelium in twig

lesions and bud scales, but only in cool maritime climates such as Northern Europe.

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
Survival of overwintering mycelia is otherwise uncommon outside of said regions due to

desiccation (MacHardy). Given V. inaequalis is a facultative pathogen, pseudothecia may

remain viable in the absence of a living host as a saprophyte on dead host tissues – such

as leaf and fruit lesions – for the next growing season, in which said pathogen initiates

primary inoculum in the form of ascospores (Keitt and Jones). Furthermore, since V.

inaequalis is a necrotroph capable of surviving on either already dead substrates or host

tissue killed directly by the pathogen via mycotoxins, it may continue to produce

secondary inoculum (i.e., conidia) through the course of the current growing season

(Brook). Apple scab is therefore a polycyclic disease (Fig. 13.).

Management: Apple scab is effectively managed with multisite protectant fungicides, such as

captan, dithianon, and ethylenebisdithiocarbamate (EBDC); single-site protectants, such as

quinone outside inhibitors (QoIs); or, single-site curative and protectant fungicides, such as

dodine, benzimidazoles, demethylation inhibitors (DMIs), and anilinopyrimidines (MacHardy).

Fungicides with curative properties are primarily used in postinfection applications, whereas

protectants are applied prior to infection as preventatives. Single-site protectants are susceptible

to becoming useless when repetitively used to manage apple scab epidemics, as V. inaequalis is

capable of rapidly evolving resistance to any environmental stressors – including fungicides –

that inhibit or promote a single gene or group of genes within the fungal metabolism.

Nevertheless, chemically-organic fungicide applications are the norm for any integrative pest

management program against apple scab (Mills).

Traditionally, scab sprays have been applied in accordance with key host phenological stages. A

typical spraying regime would have the first application at or immediately after bud break, with

subsequent applications at 1-cm green, tight cluster, full pink, bloom, and petal fall stages

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
(MacHardy). Such control programs require absolute precision, reflecting the potential risk of

infection posed to the fruit crop. The scope of these computerized predictions typically includes

infection periods, models for ascospore maturation, tree phenology, cultivar susceptibility, and

leaf litter degradation as either independent or dependent variables (Mills).

In organic apple production, variations on copper, sulfur, and lime sulfur are the most widely

employed inorganic fungicides to control apple scab. These compounds are all multisite

protectants, being very effective in killing any spores (including those of beneficial epiphytes) on

leaf surfaces, although lime sulfur appears to have some curative activity. However, the efficacy

of said protectants is subject to the weathering effects of rain. Likewise, rapid expansion of the

tree canopy at springtime (i.e., between the mouse ear and tight cluster stages) increases the risk

by which infection occurs on unprotected host tissue. As a result, repeat applications may be

necessary to control apple scab inocula, especially when rainy conditions persist over a multiday

period (Mills). In spite of the industry’s reliance upon organic and inorganic fungicides, there

have been several attempts at the development of biocontrol measures to combat the pathogen.

Unfortunately, none of said measures have been widely adopted into commercial agricultural

production systems (Sutton et al.).

In terms of other control and preventative measures, orchard sanitation practices in the autumn

can significantly reduce the number of ascospores (i.e., primary inoculum) the following spring,

thereby augmenting the efficacy by which fungicide applications control the fungus (Keitt and

Jones). For example, applications of urea prior to leaf fall mitigates the formation of

pseudothecia, increases the breakdown of leaf litter, and may potentially decrease the amount of

ascosporic inoculum by an order of 50% to 90%. Urea may also be directly applied to the leaf

litter in either late autumn, early spring prior to bud break, or both. These urea treatments can be

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
combined with the shredding of leaves with a flair mower to promote leaf decomposition and

interspecific competition betwixt V. inaequalis and soil inhabitants on leaf residues (Mill). Or, as

a substitute for urea, commercial powdered mixes containing various epiphytic microorganisms

may be applied to shredded leaf residue for a similar effect (MacHardy). However, even a 90%

reduction in pseudothecia and ascospores as a source of primary inoculum does not sufficiently

reduce scab infection under favorable conditions without a diverse mix of other control measures

(Mills).

Chemical practices are not the only means by which to prevent or control apple scab epidemics.

On the contrary, cultural practices can go a long way towards keeping disease abated. In general,

trees should be pruned regularly so that their canopies are relatively open. This management

technique promotes circulation around and through the canopy, thereby facilitating the drying of

foliage between periods of wetting, and furthermore preventing the development of

environmental conditions ideal for apple scab infection. In addition, this pruning regime

improves spray coverage during fungicide applications (Keitt and Jones).

Another means of preventing apple scab – without needing to rely on chemical inputs – is the

selection of genetically superior plant material for the home and commercial orchard. Even

amongst the commercially-available apple cultivars, the most frequented host of apple scab,

there is considerable variation in susceptibility to the pathogen. ‘Gala,’ ‘Jonagold,’ ‘McIntosh,’

and ‘Cox’s Orange Pippin’ are by far the most susceptible cultivars to V. inaequalis, whereas

‘Discovery,’ ‘Elstar,’ ‘Honeycrisp,’ and ‘Ingrid Marie’ are all examples of cultivars with

moderate to low susceptibility (Sutton et al.). For successful organically-produced operations as

well as home orchards, choosing apple cultivars with low susceptibility to scab is essential.

Altogether, more than 60 cultivars have been released worldwide in the last thirty years with

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
resistance to races of apple scab. The gene most commonly conferring resistance in these

resistant cultivars is known as the Vf gene complex (Sutton et al.). The resistance conferred by

this gene complex, originating from Malus floribunda ‘Siebold’ x ‘Van Houtte,’ has

unfortunately been overcome in several European states by V. inaequalis. In response to the

failure of cultivars with specific resistance, apple breeding programs worldwide are now

addressing apple scab with cultivars exhibiting broad resistance to the pathogen, with some

success (MacHardy). ‘Liberty,’ and ‘Florina’ and ‘GoldRush’ in particular, have been hits with

the apple growing industry; ‘Florina’ is now the second most widely grown cultivar in France

after ‘Golden Delicious,’ meanwhile ‘GoldRush’ has quickly become the sixth most widely

grown cultivar in North America, as of 2006, with a large following amongst consumers in New

York and New England (MacHardy). These three cultivars are a testament to the ongoing – and

likely never ending – work of developing apple germplasm capable of tolerating or resisting

infection on the part of Venturia inaequalis, the notorious apple scab.

Importance of Disease: Apple is one of the most important fruit crops grown and sold both

domestically and abroad, wine grape being first. Apple scab is the most economically important

disease of apples worldwide (Sutton et al.). The disease, first described in Sweden by E.M. Fries

in 1819 but known from paintings dating back to the fifteenth and sixteenth centuries, is present

wherever apples and other host plants are cultivated (Keitt and Jones). However, severity is

reduced, or the pathogen is altogether rare in semiarid regions of production, such as western

North America and western Australia (James and Sutton). Crop losses can be pronounced in

regions dominated by humid, cool weather conditions, as is the case across Northern Europe, the

northeastern United States, and Canada. Economic loss from apple scab is directly attributable to

fruit or pedicel infections (reducing shelf appeal to the consumer at market alongside poor

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
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postharvest viability) and indirectly due to repeated premature defoliation, which reduces host

growth and yields for several years (MacHardy). Likewise, repeated defoliation places

physiological stress on the host, thereby increasing susceptibility to winter injury or long-term

decline (Keitt and Jones).

References:

Blaise, P.H. et al. 1987. Apple Scab: A teaching aid on microcomputers. Plant Disease 71 (7):

574-578.

Brook, P. J. 1969. Effects of light, temperature and moisture on release of ascospores by

Venturia inaequalis (Cke.) Wint. New Zealand Journal of Agricultural Research 12:214-

227.

Gadoury, D.M., and W.E. MacHardy. 1982. A model to estimate the maturity of ascospores of

Venturia inaequalis. Phytopathology 72: 901-904.

James, J.R., and T.B. Sutton. 1982. Environmental factors influencing pseudothecial

development and ascospore maturation of Venturia inaequalis. Phytopathology 72: 1073-

1080.

Keitt, G.W., and L.K. Jones. 1926. Studies of the epidemiology and control of apple scab.

Wisconsin Agriculture Experimental Station Research Bulletin 73.

Merwin, I.A., et al. 1994. Scab-resistant apples for the Northeastern United States: New

prospects and old problems. Plant Disease 78 (1): 4-10.

MacHardy, W.E. 1996. Apple Scab: Biology, Epidemiology, and Management. APS Press: St.

Paul, Minnesota.

Mills, W.D. 1944. Efficient use of sulfur dusts and sprays during rain to control apple scab. New

York Agriculture Experimental Station, Ithaca Bulletin 630.

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation
Stensvand, A., D.M. Gadoury, T. Amundsen, and R.C. Seem. 1997. Ascospore release and

infection of apple leaves by conidia and ascospores of Venturia inaequalis at low

temperatures. Phytopathology 87: 1046-1053.

Sutton, T.B., H.S. Aldwinckle, A.M. Agnello, and J.F. Walgenbach, eds. 2014. Compendium of

Apple and Pear Diseases and Pests, Second Edition. APS Press: St. Paul, Minnesota. 8-

11.

Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The Plant Health Instructor. DOI:

10.1094/PHI-I-2000-1005-01. Updated 2005.

Definitions:

1. Ascocarp, or ascoma (pl. ascomata): The fruiting body of a fungus in the phylum

Ascomycota, consisting of very tightly interwoven hyphae and may contain millions of

asci, each of which typically contains four to eight ascospores.

2. Ascus (pl. asci): A sac, typically cylindrical in shape, in which the spores of ascomycete

fungi develop.

3. Bipolar heterothallic (from bipolar heterothallism): A fungus which requires the

conjugation of mycelia from two different mating types in order to sexually reproduce,

and yet is capable of self-fertilization on occasion.

4. Centrum (pl. centra): Collectively, all the structures enclosed within the ascocarp of a

fungus.

5. Fasciculate: Formed of, or growing in, bundles or clusters akin to fasces, a weapon used

by the ancient Romans.

6. Negatively geotropic (from negative geotropism): The act of growing against gravity.

7. Nodulose: To have minute nodules; finely knobby.

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
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8. Pseudoparaphysis (pl. pseudoparaphyses): A hypha growing betwixt asci derived from

within the pseudothecium, but originating from above (i.e., the tectum, the structure that

covers the ostiole while the pseudothecia develops), and not from the base of the

ascocarp.

9. Pseudoparenchymatous: Having a compact mass of tissue, made up of interwoven hyphae

or like filaments, that superficially resembles plant tissue.

10. Pseudothecium (pl. pseudothecia): An ascocarp resembling a perithecium, but whose asci

are not regularly organized into a hymenium and are instead bitunicate, having a double

wall which expands when it takes up water and shoots the enclosed spores out suddenly

to disperse them.

11. Stroma: A cushion-like mass of fungal tissue, having spore-bearing structures either

embedded in it or on its surface.

Register of Figures:

Fig. 1. ‘Jonathan’ fruit with apple scab lesions. (Courtesy J. Hartman and The American
Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The Plant
Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation

Fig. 2. Apple scab lesions on young ‘Golden Delicious’ leaves. (Courtesy W.E. MacHardy and
The American Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab.
The Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

Fig. 3. Scab lesions coalescing into blisters at the cuticle-epidermal interface of the leaf.
(Courtesy W.E. MacHardy and The American Phytopathological Society, Vaillancourt, L.J. and
J.R. Hartman. 2000. Apple scab. The Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-
01. Updated 2005.)

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation

Fig. 4. Blistered scab lesion on fruit. (Courtesy W.E. MacHardy and The American
Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The Plant
Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

Fig. 5. Distortion of apple fruits due to apple scab infection. (Courtesy W.E. MacHardy and The
American Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The
Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation

Fig. 6. Pseudothecium of Venturia inaequalis embedded in leaf tissue. The arrows point to
hyphae which may represent two different mating types. (Courtesy W.E. MacHardy and The
American Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The
Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

Fig. 7. Cross section of a pseudothecium of Venturia inaequalis. The cross section has been
stained so that the asci and ascospores are pink. (Courtesy W.E. MacHardy and The American
Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The Plant
Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation

Fig. 8. Asci of Venturia inaequalis, each containing eight, two-celled ascospores. (Courtesy
W.E. MacHardy and The American Phytopathological Society, Vaillancourt, L.J. and J.R.
Hartman. 2000. Apple scab. The Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01.
Updated 2005.)

Fig. 9. Two-celled ascospores of Venturia inaequalis. (Courtesy W.E. MacHardy and The
American Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The
Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation

Fig. 10. Depiction of a fallen apple leaf, showing the mature pseudothecia of Venturia inaequalis
protruding above the substrate, ready to release ascospores. (Courtesy W.E. MacHardy and The
American Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The
Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

Fig. 11. Conidia of Spilocaea pomi (teleomorph: Venturia inaequalis). (Courtesy J. Hartman and
The American Phytopathological Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab.
The Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

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Dr. Muthu Venkateshwaran, Ph.D. N.R. McVay
SCSCI 4530 – Plant Pathology Plant Disease Investigation

Fig. 12. Cross section through a leaf infected with apple scab. Arrow indicates the leaf cuticle
which has been ruptured and pushed back by the mass of erupting conidia and conidiophores.
(Courtesy W. E. MacHardy and The American Phytopathological Society, Vaillancourt, L.J. and
J.R. Hartman. 2000. Apple scab. The Plant Health Instructor. DOI: 10.1094/PHI-I-2000-1005-
01. Updated 2005.)

Fig. 13. Illustrated disease cycle of Venturia inaequalis on domestic apple. The disease cycle is
similar for other host plants of the pathogen. (Courtesy of The American Phytopathological
Society, Vaillancourt, L.J. and J.R. Hartman. 2000. Apple scab. The Plant Health Instructor.
DOI: 10.1094/PHI-I-2000-1005-01. Updated 2005.)

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