New Zealand Journal of Agricultural Research

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Pre-harvest lenticel spot of apples

P. J. Brook

To cite this article: P. J. Brook (1968) Pre-harvest lenticel spot of apples, New Zealand Journal of
Agricultural Research, 11:2, 237-244, DOI: 10.1080/00288233.1968.10431423

To link to this article: https://doi.org/10.1080/00288233.1968.10431423

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 237

PRE·HARVEST LENTICEL SPOT OF APPLES

By P. J. BROOK*

(Received 22 November 1967)

ABSTRACT
An investigation was made of lenticel spot which develops prior to
harvest on Sturmer and Golden Delicious apples. Fungi pathogenic
to apples were isolated from only a small proportion of spots, and few
spots developed into rots when fruit was held in store. Saprophytic
species, e.g., Stemphylillm botryosum and Ulocladium consortiale.
were isolated from the majority of spots. Fungicides applied through
summer reduced the amount of pre-harvest lentical spot, indicating that
the spots were caused by fungi. It is concluded that the death of apple
tissue under lenticels, constituting pre-harvest len tical spot, is caused
by saprophytic fungi which make limited growth in the cavities on the
f roit surface.

INTRODUCTION

"Lenticel spot" is the name given to disorders of apple fruits

whose symptom is a small brown or black spot centred on a lenticel.
Carne (1948) and Padfield (1954) describe lenticel spots as developing
in store or on fruit on the tree. By definition of these authors. lenticel
spot is not caused by pathogenic organisms, although the spots are
sometimes invaded subsequently by secondary rot fungi. "Lenticel
spot" is sometimes used to denote small lesions caused by Gloeosporium
spp. To avoid the confusion which arises from this different usage of
the name, the present author follows Carne (loc. cit.) and Padfield
(loc. cit.) in excluding spots known to be of pathogenic origin from
being termed "lenticel spot."

The present paper discusses lenticel spots which develop on fruit

while it is still on the tree, that is, "pre-harvest lenticel spot". Varieties
examined were Sturmer and Golden Delicious. Several years' observa-
tions on Sturmer in Nelson have established that pre-harvest lenticel
spot appears on occasional fruits in February and March, but that the
main onset of spotting occurs in April, coinciding with the period of
harvesting this variety for !>torage. Pre-harvest lenticel spot on
Golden Delicious has been observed for one year, and it seems that
spotting of this variety, too, develops as the fruit nears the maturity at
which it is harvested for storage.
• Plant Diseases Division, Department of Scientific and Industrial Research.
Private Bag, Auckland.
'N,Z. II agric, Res, (1968). 11: 237-44
238 Pre-harvest lenticel spot of apples

Pre-harvest lenticel spots occur predominantly on the apple face

exposed directly to the sun and are rarely found on the shade face
(Figs. I, 2.). They are readily distinguished from normal lenticels,
being pale brown, dark brown, or black, of pinpoint size up to about
1 mm diameter, or occasionally larger. The spots are often surrounded
by a halo of epidermal cells which contain red pigment.

1. W. Endt
Fig. I. (top). Fig. 2 (bottom) .-Unsprayed Stunner apples on 26 June. Fig. I
shows lenticel spots on the face exposed to the sun. Fig. 2 shows the shaded
face of the same apples.

The brown tissue of the spot consists of dead cells of epidermis,

hypodermis, and outer cortex, in which there is usually brown granular
matter. A cambium is often found beneath the dead cells (Fig. 3).
 P. J. BROOK 239

H o c

A . Underhill
Fig. 3 (top) .- Section through epidermis and lenticel spot on exposed face of
Sturmer apple, harvested 26 JUne. H, hypha in lenticel cavity; D, dead cells;
C, cambium.
Fig. 4 (bottom) .- Section through epidermis on shaded face of the same apple
shown in Fig. 3. W, wax plates.

EXPERIMENTAL
Fate of pre-harvest lenticel spot on stored fruit
In commercial practice, apples with pre-harvest lenticel spot are
usually discarded if the crop is to be stored. It is widely believed that
the spots are caused by Gloeosporium perennans Zeller & Childs,
G. album Osterw., or G . fructigenum Berk., and that they will develop
into rots if the fruit is held.
During summer of 1966-7. 35 Sturmer trees in Nelson were sprayed
differentially with four fungicides. seven trees being used for each,
and seven having no fungicides (Experiment C. Table 2). Two cases
 240 Pre-harvest lenticel spot of Jlpples

of fruit were taken from each tree on 31 May 1967. The fruit was
sorted into apples with pre-harvest 1enticel spot and those with none.
The clean apples and those with pre-harvest lenticel spot were stored in
separate boxes in the orchard shed until 11 August. On 7 July a record
was made of rotted fruits and these were removed from the boxes.
A second record was made on 11 August.
Mean air temperatures at the orchard during the storage period
were 7"c for June, 6.S o e for July, and lODe for the first 11 days of
August.
Results are shown in Table 1. Figures for rots in the table are
combined totals from the two recording dates.

TABLE I. Amolmt of rottillg amollgst Sturmer apples held 72 days ill shed
storage, Nelson, 1967. The fruit was from Experiment C, TC/ble 2

Percentage apples with rots by

~ean No.
Fruit condition I No. apples II August
Orchard spots per
treatment at harvest, I stor d apple,
31 ~ay e
31 ~ay
I

Fungicides~ Clean 5,270 Nil 0.47% 0.21% 0.26%

Lenticel spot 1,027 6 1.75% 1.17% 0.58%

-I
No fungicide I Clean 805 Nil 0.99% 0.62% 0.37%
Lenticel spot 752 26 2.39% 2.13% 0.26%

1 Gloeosporium rots caused by G. fructigenum, G. perennans, and G. album.

2 Fruit from trees sprayed in summer with Bordeaux mixture, captan, thiram,
or captafol is grouped in this table.

The general level of rotting in the stored fruit was low, but there
were more rots amongst apples which had pre-harvest lenticel spot
lhan amongst those which entered store clean. However, of the fruit
which had entered store with lenticel spot, 98% of the sprayed and
97% of the unsprayed had no rots after 72 days. That is, approxi-
mately 19,000 pre-harvest lenticel spots on the unsprayed apples
remained in an unchanged condition throughout the storage period.

Orchard fungicide programmes and pre-harvest lenticel spot

Spray programmes in which only the fungicides were variable were
applied to Sturmer in Nelson in three years, and to Golden Delicious
in Hawke's Bay in one year. After lenticel spot appeared, successive
fruit samples were picked and sorted as clean or having lenticel spot.
No differentiation was made in these records between apples with few
spots and those with many. In 1967 counts of spots on individual
apples were also made.
 P. J. BROOK 241
TABLE 2. Development of pre-harvest lenticel spot on apples sprayed under
programmes in which only the summer fungicides were variable. Sturmer trees
were in Nelson, Golden Delicious in Hawke's Bay

I
Differential summer Percentage fruits with pre-harvest Mean No.
fungicide treatments lenticel spot on: spots per
fruit on:
I

I I
A. Sturmer, 1960--1. 9 May 6 June I 27 June
10 sprays, 30 Nov.-
16 June
INo record
Bordeaux mixture 1/4/100 1 I
18 66
Captan 0.05% 20 57 69
Nil 55 87 99

I I
B. Sturmer, 1961-2. 7 April 5 May 5 June
8 sprays, 5 Dec.-
24 April I
I
I No record
Bordeaux mixture 1/4/100 Nil 3 51
Captan 0.05% 1 42 83
Nil 11 i 57 88
I

I
C. Sturmer, 1966-7. 22 May 26 June 10 July 26 June
10 sprays, 17 Dec.-
22 June
I

Bordeaux mixture 1/4/100 3 46 76 4

Captan 0.05% 21 59 I 60 9
Thiram 0.08% 34 55 73 10
Captafol 0.08% 7 24 I 32 3
Nil I 48 85 99 34
I I I

D. Golden Delicious 1966-7. 21 Feb. 20 Apr. j

1 June I 1 June
5 sprays, 14 Dec.-
22 Feb.
I ..
Captan 0.05% 30 57 99 16 .
Thiram 0.08% 47 59 94 i
14
Captafol 0.08% 34 51 69 2
Nil 47 60 93 15
i
Note. Normal harvest time for Sturmer is from the end of March until early
in May; and for Golden Delicious is from late in February until mil-April.

A summary of the experiments and results is given in Table 2.

The lower incidence of pre-harvest lenticel spot when fruit was
sprayed with fungicides, particularly captafol or Bordeaux mixture.
indicated that the agents causing the spots were fungi.
Fungi associated with pre-harvest lenticel spot
Pre-harvest lenticel spots were cultured in the following way. Apples
were washed under the tap. plunged in 0.25% sodium hypochlorite
solution for about one minute. and finally rinsed in sterile water. Whole
spots. including cuticle. were then transferred to nutrient agar.
242 Pre-harvest lenticel spot of apples

Spots were cultured from Nelson Sturmer harvested on 29 May

and 19 June 1967 (365 spots). and from Hawke's Bay Golden Delicious
harvested on 18 May 1967 (200 spots). Fungi emerged from 79%
of the Sturmer spots. and from 93% of the Golden Delicious spots.
Table 3 lists the species. Of the 26 undetermined sterile isolates from
Sturmer. 20 were identical colonies of predominantly black mycelium.

TABLE 3. Fungi isolated from pre-harvest lenticel spots on Nelson Sturmer

harvested 29 May and 19 June, and on Hawke·s Bay Golden Delicious harvested<
18 May

Sturmer Golden
(288 isolates) Delicious
! (186 isolates)
%
I %
Ulocladium consortiale (ThUrn.) Simmons 33 83
Stemphylium botryosum Wallr. 45
Epicoccum purpurascens Ehrenb. ex Schlecht. 3 7

Cladosporium herbarum Fr. 2

Phoma exigua Desrn. 3
Diaporthe perniciosa March. 4

Gloeosporium fructigenum Berk. < 6

G. perennans Zeller & Childs <
Undertermined (sterile) 9 4

Pre-harvest lenticel spots were also examined microscopically for

fungi. Hyphae were rarely seen penetrating amongst the dead apple
cells. On the other hand, hyphae. and spores, were common in lenticel
cavities. although development of mycelium was limited (Figs. 3. 5).

Apple fruit surface and pre-harvest lenticel spot

Pre-harvest lenticel spots are almost entirely confined to the apple
face exposed to the sun (Figs, 1, 2). Lenticels are, however. distributed
over the entire apple surface.
Sturmer apples from Nelson harvested on 19 June were examined
for differences between exposed and shaded surfaces. Sections of
epidermis were cut. at right angles to the surface. from the exposed
and the shaded faces of 10 apples. Cuticle thickness was measured above
10 epidermal cells from each face, from the inner side of the cell wall
to the outer boundary of the cuticle.
Mean cuticle thickness on the shaded face was 18 p., and. on the
exposed face, 15 p.. The mean difference between pairs of faces on
individual apples was 3 p. (significant at P < 0.001) .
 P. J. BROOK 243

A. Underhill
Fig. 5. -Surface view of lenticcl spot on Golden Delicious apple harvested
I June. The dead apple cells are not in focus. but hyphae and spores are
visible in the lenticel cavity.

The difference in cuticle thickness was largely due to the presence

of a layer of wax plates 2- 3 JL thick on the shaded face. This layer
was absent from the exposed face. Figs. 3 and 4, of sections cut fmm
opposite faces of the same apple, illustrate this difference.

DISCUSSION
The results described here showed !hat few pre-harvest lenticel
spots developed into rots when apples were held in store. In addition.
pathogens such as Gloeosporium spp. and Diaporthe perniciosa were
isolated from only a small proportion of the spots. It appears. therefore.
that the great majority of lenticel spots which develop on Sturmer and
Golden Delicious prior to harvest are not caused by pathogenic fungi.
The effectiveness of some fungicides in reducing the amount of
pre-harvest lenticel spot is evidence. however. that the spots are in
fact , caused by fungi. The conclusion drawn from the present work
244 Pre-harvest lenticel spot of apples

is that members of the apple surface microftora, notably Stemphylium

botryosum and Ulocladillm cOllSortiale, are causal agents of the spotting.
Hyphae were rarely found amongst or within the dead apple cells, and
it is assumed that cell death is caused by substances released from the
fungi growing in the lenticel cavities.
Lenticels and surface-inhabiting fungi are distributed over the
entire apple fruit, but pre-harvest lenticel spot is virtually confined
to the apple face exposed to the sun. It would appear, therefore, that
there is a difference in susceptibility between the exposed and the
shaded faces.
Clements (1935) described lenticels on apple fruits as cavities which
develop at stomata and where breaks occur in the epidermis as the
fruit expands. As the apple swells, ruptures occur between cells lining
the cavities, and the breaks are repaired by cutinisation of newly exposed
walls. Clements (loc. cit.) found that a distinct phellogen rarely
develops in lenticels on apple fruits, and he pointed out that apple
fruit lenticels are not similar to lenticels of woody stems.
The readiness with which ruptures are repaired in the epidermis and
in lenticel cavities probably determines whether or not superficial
fungi cause death of underlying apple cells. It was found that the
exposed face of the Sturmer apples lacked the layer of wax plates which
covered the shaded face. This indicates that direct exposure to the
sun's heat caused drying and hardening of cuticular wax. It is
suggested, therefore, that the susceptibility of the exposed face to
pre-harvest lenticel spot is due to reduced plasticity of cuticular wax
and consequent slowing down of cell wall repair at rupture points.

ACKNOWLEDGMENTS
Thanks are due to Messrs M. van Geldermalsen and 1. A. MacLean,
Fruit Research Division, D.S.LR., who carried out the field work in
Nelson and Hawke's Bay respectively.

REFERENCES
CARNE, W. M. 1948: The non-parasitic disorders of apple fruits in
Australia. Bull. Commonw. scient. indo Res. Org. 238. 83 pp.
CLEMENTS, H. F. 1935: Morphology and physiology of the pome
lenticels of Pyrus malus. Bot. Gaz. 97: 101-17.
PADFIELD, C. A. S. 1954: The storage of apples and pears. Bull. N.Z.
Dep. scient. indo Res. 111. 96 pp.